207
A su ey o he spide genus Lipoc ea Tho ell, 1878 (A aneae,
A aneidae) om Guiyang Ci y, Sou hwes China: An in eg a ed
mo phological and molecula app oach
Jianshuang Zhang1* , Chengwen Zhang1* , Yuanqian Xing1, Hao Yu1, Xiaoqi Mi2
1 The S a e Key Labo a o y o Sou hwes Ka s Moun ain Biodi e si y Conse a ion o Fo es y Adminis a ion, School o Li e Sciences, Guizhou No mal
Uni e si y, Guiyang, Guizhou, China
2 College o Ag icul u e and Fo es y Enginee ing and Planning, Guizhou P o incial Key Labo a o y o Biodi e si y Conse a ion and U iliza ion in he Fanjing
Moun ain Region, Tong en Uni e si y, Tong en, Guizhou, China
Co esponding au ho : Xiaoqi Mi ([email p o ec ed])
Copy igh : © Jianshuang 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
A su ey was unde aken o s udy he spide genus Lipoc ea Tho ell, 1878, om Guiyang
Ci y, Guizhou P o ince, sou hwes China. A o al o wo species is he e add essed based
on mo phology and i e me hods o molecula species delimi a ion, comp ising L. gui-
yang J. Zhang, Yu & Mi, sp. no . and L. usi o mis (Tho ell, 1877), he ype species o he
genus as well as a new eco d o mainland China. These wo species a e dis ibu ed
in Huaxi Dis ic and Kaiyang Coun y o Guiyang, espec i ely, p o iding he i s o mal
eco d o his genus om mainland China.
Key wo ds: Biodi e si y, DNA ba coding, molecula species delimi a ion, mo phology,
new species, o b-wea e
In oduc ion
Lipoc ea Tho ell, 1878 is a small spide genus ha was o iginally es ablished o
include h ee species: L. ph hisica (L. Koch, 1871) and L. abida (L. Koch, 1872)
om he Aus alasian Region, and he ype species L. usi o mis (Tho ell, 1877)
om he O ien al Region. An addi ional O ien al species, L. dilu a Tho ell, 1887
was subsequen ly desc ibed. Howe e , he alidi y o Lipoc ea as a dis inc ge-
nus was no ecognized by Simon (1887: 187), who ea ed hese species unde
he genus La inia Simon, 1874. O he a o emen ioned species, L. ph hisica has
since been ans e ed o La inia sensu s ic o (G assho 1970). The emain-
ing species, oge he wi h La inia longissima (Simon, 1881) om he E hiopian
Region, we e subsequen ly emo ed om La inia and placed in a newly e ec ed
genus, La inopa G assho (1970), by G assho (1970: 226). Al hough he name
Lipoc ea emains a ailable unde he ules o zoological nomencla u e, i has
been la gely o e looked (Le y 1986). The genus Lipoc ea, as ede ined by G ass-
ho unde he name La inopa, is p ima ily dis inguished om La inia by he
s uc u e o he geni alia, especially he mo phology o he male palpal o gan.
Academic edi o : C is ina Rheims
Recei ed:
12 May 2025
Accep ed:
25 Augus 2025
Published:
10 Oc obe 2025
ZooBank: h ps://zoobank.
o g/07FB18BD-9211-413B-A660-
05B73FF82A0F
Ci a ion: Zhang J, Zhang C, Xing
Y, Yu H, Mi X (2025) A su ey o
he spide genus Lipoc ea Tho ell,
1878 (A aneae, A aneidae) om
Guiyang Ci y, Sou hwes China:
An in eg a ed mo phological and
molecula app oach. ZooKeys 1255:
207–237. h ps://doi.o g/10.3897/
zookeys.1255.158340
ZooKeys 1255: 207–237 (2025)
DOI: 10.3897/zookeys.1255.158340
* These au ho s con ibu ed equally o his wo k.
208
ZooKeys 1255: 207–237 (2025), DOI: 10.3897/zookeys.1255.158340
Jianshuang Zhang e al.: A su ey o Lipoc ea om Guiyang
Cu en ly, he WSC (2025) lis s i e species unde Lipoc ea, none o which
a e known om he Chinese mainland (WSC 2025). Howe e , in se e al publi-
ca ions, some o hese species a e conside ed o belong o La inia. Fo exam-
ple, L. phosop (Tanikawa, In o & Pe cha ad, 2023) was o iginally desc ibed as
La inia has no ye been o mally ans e ed o Lipoc ea. Ne e heless, due o
i s s ong mo phological simila i y o he ype species o Lipoc ea, L. phosop
has been p o isionally placed in Lipoc ea by he WSC (2025). E en he ype
species, L. usi o mis, has equen ly been placed in La inia and edesc ibed
as such by a ious au ho s, including Tanikawa (1989, 2007, 2009), Okuma e
al. (1993), Ba ion and Li singe (1995), and Chang and Tso (2004). In iew o
he abo e-men ioned, he alidi y o he genus Lipoc ea and he cu en gene ic
placemen o i s cons i uen species emains in dispu e.
A p elimina y genus-le el axonomic molecula analysis o G assho ’s (1970)
‘La inia g oup’ was ca ied ou based on all a ailable COI sequences o he La inia
g oup and ela ed gene a (69 om NCBI and 9 newly sequenced he e) (Suppl. ma-
e ial 1: able S1). Acco ding o he esul s (Suppl. ma e ial 1: ig. S1): (1) he mono-
phyly o he genus Lipoc ea is well suppo ed; (2) La inia is polyphyle ic, which
is consis en wi h he esul s o Scha e al. (2020). Based on his p elimina y
esul and conside ing ha a o mal comp ehensi e e ision o he La inia g oup is
lacking, we ag ee wi h F amenau and Cas anhei a (2022) ha he La inia g oup e-
qui es u he sys ema ic s udy ha includes La inia linea a Lucas, 1846 ( he ype
species o he genus). A e iew o he La inia g oup is no wi hin he scope o his
wo k. Consequen ly, he p esen s udy assigns he wo species ea ed he e o he
monophyle ic genus Lipoc ea, a he han o he polyphyle ic La inia sensu la o.
Guiyang, he p o incial capi al o Guizhou P o ince (Fig. 1A), is he i s ci y
o ecei e he accolade o ‘ he na ional o es ci y’ in China, amous o mo e
han 55% o es co e age (Zhong 2018). Guiyang is also known as he ‘ci y o
a housand ga dens’, wi h 1025 u ban pa ks, and is conside ed one o China’s
mos biodi e se p o incial capi als (Yang 2020; Cheng and Chen 2022). How-
e e , spide s can be ega ded as being poo ly ep esen ed in Guiyang, wi h only
64 species om 25 amilies eco ded o desc ibed o da e (JZ and HY, unpubl.
da a). O hese, 21 species a e endemic, 16 we e newly desc ibed, and 15 we e
epo ed as new eco ds in ecen yea s (Yu e al. 2018; Yu and Zhang 2019;
Xin e al. 2020; Yan e al. 2021; Long e al. 2022; Zhang e al. 2022a, b, 2023;
He e al. 2023; Li e al. 2023; Yang e al. 2023a, b, c; Ding e al. 2024; Pan e al.
2024; Jiang e al. 2025; Zhang e al. 2025). This es ima e o spide di e si y is
assumed o be a om he ue di e si y wi hin his ci y.
As men ioned abo e, spide s a e poo ly s udied in Guiyang, including A a-
neidae. Only i e species ha e been eco ded: Cy a achne bu o (Bösenbe g &
S and, 1906), Hypsosinga pygmaea (Sunde all, 1831), Neoscona xishanensis
Yin, Wang, Xie & Peng, 1990, Nephila pilipes (Fab icius, 1793) and P onoides
b unneus Schenkel, 1936. Recen ly, sho bu in ensi e ield collec ions in Gui-
yang ha e been conduc ed by he s a o he Guizhou No mal Uni e si y. Du ing
hese su eys, we ha e ound some Lipoc ea specimens ha belong o a leas
wo mo phospecies (Figs 1–3): one is new o science, and he o he one has
been iden i ied as L. usi o mis, a new eco d o mainland China. The sym-
pa ic dis ibu ion wi h L. usi o mis and he high in aspeci ic mo phological
a ia ion in emales o he new species pose signi ican challenges o sexual
pai ing and species iden i ica ion. We he e o e gene a ed DNA ba code da a
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Jianshuang Zhang e al.: A su ey o Lipoc ea om Guiyang
Figu e 1. Locali y o Guiyang Ci y (A) and dis ibu ion eco ds o Lipoc ea species in Guiyang (B).
A
B
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Jianshuang Zhang e al.: A su ey o Lipoc ea om Guiyang
Figu e 2. Li ing specimens o Lipoc ea guiyang J. Zhang, Yu & Mi, sp. no . A. Male; B. Female. Pho og aphs by Q Lu
(Shenzhen).
A
B
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Jianshuang Zhang e al.: A su ey o Lipoc ea om Guiyang
Figu e 3. Li ing specimens o Lipoc ea usi o mis (Tho ell, 1877). A. Male; B, C. Female. Pho og aphs by Q Lu (Shenzhen).
AB
C
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Jianshuang Zhang e al.: A su ey o Lipoc ea om Guiyang
o de ise a specimen phylogeny and used i e molecula species delimi a ion
me hods o es mo phology-based species iden i ica ion.
The goal o his pape is o: 1) use he consensus esul s o an in eg a -
ed mo phological and molecula app oach o delimi Lipoc ea spide s om
Guiyang; 2) desc ibe he new species unde he name o Lipoc ea guiyang J.
Zhang, Yu & Mi, sp. no .; 3) e-illus a e L. usi o mis based on new ma e ial
om Guiyang, including supplemen a y mic og aphs; and 4) p o ide a dis ibu-
ion map o Lipoc ea species in Guiyang Ci y.
Ma e ials and me hods
Taxon sampling
Specimens in his s udy we e collec ed by hand and by bea ing ege a ion, and
di ec ly ixed in absolu e e hanol. The igh legs we e emo ed and s o ed a
−80 °C o subsequen DNA ex ac ion. The emainde o he specimens was
p ese ed in 80% e hanol o iden i ica ion and mo phological examina ion. A
o al o 11 adul s we e ob ained, examined, and p ocessed o DNA ex ac ion
bu only nine indi iduals yielded useable DNA (Table 1). All ouche specimens
(including ypes o he new species) a e deposi ed in he Museum o Guizhou
No mal Uni e si y, Guiyang, China (MGNU).
Molecula p o ocols
To al genomic DNA was ex ac ed using he Cell & Tissue Genomic DNA Isola-
ion Ki (Bio eke, Beijing, China) ollowing he manu ac u e ’s p o ocols. We am-
pli ied cy och ome c oxidase subuni I (COI) using he p ime pai s LCO1490/
HCO2198 (Folme e al. 1994) and s anda d polyme ase chain eac ion (PCR)
se ings (Wheele e al. 2016). PCR p oduc s we e anspo ed o he Beijing
Tsingke Bio ech Co., L d. (Chongqing, China) o sequencing using he same
PCR p ime s. We manually edi ed he sequences using Geneious P ime 2024
(Kea se e al. 2012), ansla ed nucleo ide sequences in o amino acids o check
o s op codons, and ensu ed he p ope con igu a ion o codon posi ions.
Phylogene ic analyses
Fo phylogene ic analyses, he COI da a o 17 specimens o Lipoc ea we e used
as he in-g oup, including eigh sequences o L. phosop downloaded om Gen-
Bank (Tanikawa e al. 2023; NCBI 2025). To oo he ee, we included wo spec-
imens o La inia joei Tanikawa & Pe cha ad, 2021 om NCBI (2025) as he ou -
g oup (Table 1). We pe o med maximum-likelihood (ML) analyses using IQ-TREE
. 2.3.1 (Minh e al. 2020) based on he bes COI subs i u ion model (GTR+I+G) in
jMODELTEST . 2.1.10 (Da iba e al. 2012). B anch suppo was es ima ed wi h
ul a as boo s apping wi h 1000 eplica es (Hoang e al. 2018). Bayesian-in e -
ence (BI) was pe o med wi h M Bayes . 3.2.1 (Ronquis e al. 2012) using one
independen chain o 50 million gene a ions. The i s 10% o ees om each un
we e disca ded as bu n-in. Finally, we used FigT ee . 1.4.4 (Rambau 2012) o i-
sualize and manipula e ees and used Pho oshop CC 2018 o summa ize hem.
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Jianshuang Zhang e al.: A su ey o Lipoc ea om Guiyang
Table 1. Samples used in species delimi a ion: specimen label, axon name, sample collec ion locali y wi h coo dina es,
and GenBank accession numbe s.
Specimen code Genus/Species Sex Locali y Coun y Coo dina es Ele a ion
(m a.s.l.)
COI GenBank
accession
YHGY208 Lipoc ea guiyang sp. no . ♂Nanjiang G and Canyon, Kaiyang Coun y,
Guiyang Ci y, Guizhou P o .
China 26.94°N, 106.97°E 861 PX230067
YHGY209 Lipoc ea guiyang sp. no . ♀Nanjiang G and Canyon, Kaiyang Coun y,
Guiyang Ci y, Guizhou P o .
China 26.94°N, 106.97°E 861 PX230068
YHGY428 Lipoc ea guiyang sp. no . ♀Nanjiang G and Canyon, Kaiyang Coun y,
Guiyang Ci y, Guizhou P o .
China 26.94°N, 106.97°E 861 PX230069
YHGY431 Lipoc ea usi o mis ♂Dangwu Town, Huaxi Dis ic , Guiyang
Ci y, Guizhou P o .
China 26.38°N, 106.60°E 1138 PX230061
YHGY432 Lipoc ea usi o mis ♀Uni e si y Town, Huaxi Dis ic , Guiyang
Ci y, Guizhou P o .
China 26.38°N, 106.65°E 1173 PX230062
YHGY433 Lipoc ea usi o mis ♀Dangwu Town, Huaxi Dis ic , Guiyang
Ci y, Guizhou P o .
China 26.38°N, 106.60°E 1138 PX230063
YHGY495 Lipoc ea usi o mis ♂Uni e si y Town, Huaxi Dis ic , Guiyang
Ci y, Guizhou P o .
China 26.38°N, 106.65°E 1173 PX230065
YHGY507 Lipoc ea usi o mis ♀Uni e si y Town, Huaxi Dis ic , Guiyang
Ci y, Guizhou P o .
China 26.38°N, 106.65°E 1173 PX230066
YHGY508 Lipoc ea usi o mis ♀Uni e si y Town, Huaxi Dis ic , Guiyang
Ci y, Guizhou P o .
China 26.38°N, 106.65°E 1173 PX230064
AT5343 Lipoc ea phosop ♂Mai Khe , Mueang P achinbu i Dis ic ,
P achinbu i P o .
Thailand 14.10°N, 101.33°E LC756453
AT5337 Lipoc ea phosop ♀Mai Khe , Mueang P achinbu i Dis ic ,
P achinbu i P o .
Thailand 14.10°N, 101.33°E LC756454
AT5338 Lipoc ea phosop ♀Mai Khe , Mueang P achinbu i Dis ic ,
P achinbu i P o .
Thailand 14.10°N, 101.33°E LC756455
AT5339 Lipoc ea phosop ♀Mai Khe , Mueang P achinbu i Dis ic ,
P achinbu i P o .
Thailand 14.10°N, 101.33°E LC756456
AT5340 Lipoc ea phosop ♀Mai Khe , Mueang P achinbu i Dis ic ,
P achinbu i P o .
Thailand 14.10°N, 101.33°E LC756457
AT5341 Lipoc ea phosop ♀Mai Khe , Mueang P achinbu i Dis ic ,
P achinbu i P o .
Thailand 14.10°N, 101.33°E LC756458
AT5342 Lipoc ea phosop ♂Mai Khe , Mueang P achinbu i Dis ic ,
P achinbu i P o .
Thailand 14.10°N, 101.33°E LC756459
AT5345 Lipoc ea phosop ♂Ban Pa hum, Sam Khok Dis ic , Pa hum
Thani P o .
Thailand 14.08°N, 100.58°E LC756460
LJO01 La inia joei ♂Khlong Sam, Pa hum Thani P o . Thailand 14.16°N, 100.66°E LC597525
LJO02 La inia joei ♀Khlong Sam, Pa hum Thani P o . Thailand 14.16°N, 100.66°E LC597526
Molecula species delimi a ion
To delimi h ee mo phospecies o Lipoc ea based on an accompanying mo -
phological s udy o he genus, we used wo gene ic dis ance-based me hods:
he DNA ba coding gap (Ba e and Hebe 2005) and ABGD (Puilland e e al.
2012), as well as h ee me hods based on he in e ed ee, GMYC (Pons e al.
2006), P ID (Libe al), and mPTP (Kapli e al. 2017).
Because he P ID (Libe al) and DNA ba coding gap (Ba e and Hebe 2005)
me hods equi e a p io i designa ion, we assigned 17 Lipoc ea indi iduals o
h ee pu a i e species based on a combina ion o phylogene ic opology and
mo phological cha ac e is ics. Wi h he DNA ba coding gap, we used he o e -
lap be ween he in e species and in aspecies Kimu a wo-pa ame e (K2P)
and unco ec ed p-dis ance o each candida e species calcula ed in MEGA X
(Kuma e al. 2018). The P ID (Libe al) me hod es s species delimi a ion by
214
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Jianshuang Zhang e al.: A su ey o Lipoc ea om Guiyang
elying on de ining he pu a i e species g oups. We used he BI ee as a guide
o es species hypo hesis (Xu e al. 2017).
The o he h ee me hods ha we used do no equi e e minals o be a p io-
i assigned o pu a i e species. ABGD calcula es all pai wise dis ances in he
da a se , e alua es in aspeci ic di e gences, and hen so s he e minals in o
candida e species wi h calcula ed P alues. We pe o med he ABGD analysis
on a web se e (h ps://bioin o.mnhn. /abi/public/abgd/) using h ee di e -
en models: Jukes-Can o (JC69; Jukes and Can o 1969), K2P (Kimu a 1980),
and simple dis ance (p-dis ance; Nei and Kuma 2000). We analyzed he da a
using wo di e en alues o he pa ame e s Pmin (0.001 and 0.0001), Pmax (0.1
and 0.2), and ela i e gap wid h (X = 1.5 o 2), wi h he o he pa ame e s se o
de aul alues. We used he BI ee as a guide o es he species hypo hesis
(Xu e al. 2017). Two uns o 100 million s eps we e used o he mPTP logging
e e y 1 million s eps, disca ding he i s 2 million s eps. Each un was s a ed
om a andom delimi a ion. The GMYC me hodology (Pons e al. 2006) anal-
ysis was conduc ed using he single- h eshold model in he “spli s” package
(Eza d e al. 2009) o R 4.2.2 (R De elopmen Co e Team 2024). BEAST 2.6.7
(Bouckae e al. 2014) was used o p oduce an ul ame ic ee o he GMYC
analysis. Analyses we e un o 50 million s eps wi h 10% o he ees in each
chain disca ded as bu n-in.
Mo phological p o ocols
Specimens we e examined using an Olympus SZX7 s e eomic oscope. Fu he
de ails we e s udied unde a CX41 compound mic oscope. Male and emale
copula o y o gans we e examined and illus a ed a e dissec ion. Epigynes
we e emo ed and clea ed in lac ic acid o a wa m 10% po assium hyd oxide
(KOH) solu ion. Images we e cap u ed wi h a Canon EOS 70D digi al came a
(20.2 megapixels) moun ed on an Olympus CX41 compound mic oscope and
assembled using Helicon Focus . 6.80 image-s acking so wa e. All measu e-
men s we e ob ained using an Olympus SZX7 s e eomic oscope and a e gi en
in millime e s. Eye diame e s we e measu ed a he wides pa . The o al body
leng h does no include he chelice ae o spinne e s. Leg leng hs a e gi en as
o al leng h ( emu , pa ella+ ibia, me a a sus, a sus). The e minology used in
he ex and igu e legends ollows G assho (1970), F amenau and Scha
(2008), F amenau and Cas anhei a (2022), Tanikawa and Pe cha ad (2021),
and Tanikawa e al. (2023).
Re e ences o igu es in he ci ed pape s a e lis ed in lowe case ( ig. o igs);
igu es om his pape a e no ed wi h an ini ial capi al (Fig. o Figs). The abb e-
ia ions used in he ex a e:
AER an e io eye ow;
ALE an e io la e al eyes;
AME an e io median eyes;
C conduc o ;
CD copula o y duc ;
CO copula o y opening;
Cy cymbium;
Em embolus;
FD e ilisa ion duc ;
HP hook-shaped p ocess o MA;
KP knob-like p ojec ion;
MA median apophysis;
MS median sep um;
MOQ median ocula quad angle;
Pc pa acymbium;
PLE pos e io la e al eyes;
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Jianshuang Zhang e al.: A su ey o Lipoc ea om Guiyang
PME pos e io median eyes;
R adix;
RA adix apophysis;
RER pos e io eye ow;
Sc scape;
Sp spe ma heca;
S sub egulum;
T egulum;
TA e minal apophysis;
TA I e minal apophysis I;
TA II e minal apophysis II;
TAA e minal apophysis appendix;
TE egula ex ension.
The dis ibu ion map was gene a ed wi h A cGIS . 10.5 (En i onmen al Sys-
ems Resea ch Ins i u e, Inc.).
Resul s and discussion
Based on adi ional mo phological cha ac e s and expe ience (ma ching o
males and emales we had hypo hesized mainly on he basis o co-occu ence
and compa ibili y o epigynes wi h male pedipalpal s uc u es), all examined
ma e ials could be iden i ied as a leas wo mo phospecies: one belongs o
an undesc ibed species new o science: L. guiyang sp. no .; he o he one was
iden i ied as L. usi o mis (G assho 1970; Chang and Tso 2004; Tanikawa
e al. 2023). Howe e , some mo phological a ia ion is exhibi ed in emales
o L. guiyang sp. no . Fi e molecula species delimi a ion me hods we e em-
ployed o es he alidi y o he mo phology-based iden i ica ion o he h ee
Lipoc ea species and he accu acy o he p oposed sex ma ching.
The COI ma ix o 17 Lipoc ea indi iduals analyzed in his s udy had a sequence
leng h o 629 bp, wi h 132 a iable and 116 pa simony-in o ma i e si es. Fo COI,
phylogene ic in e ence om BI and ML analyses yielded simila opologies wi h
high suppo (Fig. 4; pos e io p obabili y, PP = 1; boo s ap alue, BS = 100). The
ees clea ly di ided he samples in o ou deeply di e gen clades (Fig. 4).
When conside ing h ee species o Lipoc ea, in e speci ic dis ances we e
highe han in aspeci ic dis ances. In e speci ic dis ances ange om 3.27 o
11.69% o K2P (Fig. 5A) and om 3.18 o 10.65% o unco ec ed p-dis ance
(Fig. 5B). The lowes mean in e speci ic dis ance was 12.75% / 11.51% (K2P
/ unco ec ed p-dis ance) ound be ween L. phosop and L. usi o mis, and he
highes mean in aspeci ic dis ance (1.81% / 1.77% K2P / unco ec ed p-dis-
ance) was es ima ed o L. usi o mis. The ba coding gap ange iden i ied
h ee species, one o which is new (Fig. 4).
The ABGD esul s a ied based on di e en pa ame e combina ions o
bo h he ini ial and ecu si e pa i ions. Based on he ba coding gap esul s, a
dis inc gap was obse ed among he h ee Lipoc ea species, demons a ing
subs an ial gene ic di e gence. The e o e, he alue o X=2 was selec ed as
he ela i e gap wid h (Table 2). The ini ial h ee species pa i ion was consis-
en wi h he h ee mo phospecies, while he ecu si e pa i ion egime yielded
mo e species (Fig. 4; Table 2). Howe e , all analyses unde di e en assump-
ions e ealed h ee hypo he ical species which is en i ely consis en wi h he
mo phological iden i ica ion (Fig. 4).
Ou P ID (Libe al) esul s e ealed high P ID (Libe al) alues > 0.95 (0.86–1.0)
(Table 3), he eby also suppo ing he axonomy o h ee pu a i e species (Fig. 4).
The esul s o ou mPTP analysis indica ed ha , when only monophyle ic
species a e conside ed, ou h ee hypo he ical species we e iden i ied as an ic-
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Jianshuang Zhang e al.: A su ey o Lipoc ea om Guiyang
Figu e 7. Lipoc ea guiyang J. Zhang, Yu & Mi, sp. no ., emale pa a ypes, YHGY428 (A–C) and YHGY209 (D–F), habi us.
A, D. Do sal; B, E. Ven al; C, F. La e al. Scale ba s: 1 mm.
A B C
D E F
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Jianshuang Zhang e al.: A su ey o Lipoc ea om Guiyang
de ed wi h yellow lines and bea ing a p ominen la ge black spo an e io ly;
la e ally wi h wo dis inc yellow longi udinal lines, each line accompanied by
app oxima ely ou small black spo s; en e g ayish, wi hou dis inc pa e n;
spinne e s yellow.
Palp (Figs 8A–D, 9A–E). Cymbium (Cy) na icula , ~2.2× longe han wide,
do sally wi h spa se, long se ae (all de ached in e hanol), baso e ola e ally
wi h a humb-like pa acymbium (Pc). Pc mode a ely la ge, abou 1/5 leng h o
cymbium, apex blun , sligh ly cu ed and poin ing e ola e o-dis ally. Tegulum
(T) disc-shaped, sligh ly wide han cymbium, wi h dis inc spe m duc along
an e io ma gin, p oximally co e ed by b oad sub egulum (S ). Tegula ex en-
sion (TE) lamina , ex ending do sally, almos comple ely concealed by conduc-
o (C) in en al iew. S ~1/2 cymbium leng h, pa ly memb anous, su ace
w inkled and ibbed, wi h nume ous diagonal idges. Radix (R) lea shaped, ~½
he wid h o he sub egulum leng h, dis ally wi h a iangula apophysis (RA). RA
hyaline, nea ly as long as adix, apex sha p and poin ing dis ally. Median apoph-
ysis (MA) hea ily scle o ized, loca ed p ola e ally o egulum, consis ing o a
b oad base and a hook-shaped p ocess (HP); base na icula , ~2/5 he wid h
o he sub egulum in leng h; HP nea ly as long as base, apex sha p, dis inc ly
cu ed and poin ing e ola e ally. Te minal apophysis (TA) hidden behind eg-
ulum, ex ending dis ally, apex su passing he egulum and bi u ca ing in o wo
apophyses, o ming a C-shape in an e io iew; bo h e minal apophysis I (TA I)
and e minal apophysis II (TA II) hea ily scle o ized, wi h blun apices poin ing
p ola e ally; TA I ela i ely la ge, i s leng h nea ly equal o he wid h o he egu-
lum; TA II smalle and humble, ~½ he leng h o TAI. Te minal apophysis appen-
dix (TAA) memb anous, digi i o m, accompanied by e minal apophysis, hidden
behind egulum, ex ending dis ally. C o igina ing om do sal-an e io po ion
o egulum, p oximally used o weakly scle o ized TE; ip dis inc ly cu ed,
shaped like an ox ho n, wi h a sha p apex poin ing do so-dis ally. Embolus (Em)
spine-shaped, nea ly as long as he hook-shaped p ocess o he median apoph-
ysis, o igina ing cen ally in an e io iew, ex ending dis ally, su ounded by he
RA, MA, TA, and C.
Female (YHGY428). To al leng h 7.30. Ca apace 2.83 long, 1.81 wide. Abdo-
men 4.96 long, 2.67 wide. S e num 1.25 long and 0.84 wide. Labium 0.28 long
and 0.49 wide. Endi es 0.61 long and 0.43wide. Clypeus heigh 0.10. Bo h ma -
gins o chelice ae wi h ou ee h. Eye sizes and in e dis ances: AME 0.15, ALE
0.13, PME 0.13, PLE 0.11, AME–AME 0.23, ALE–AME 0.19, PME–PME 0.02,
PME–PLE 0.35. MOQ 0.44 long, an e io wid h 0.48, pos e io wid h 0.27. Leg
measu emen s: I 14.53 (3.33, 5.36, 4.54, 1.30), II 13.15 (3.26, 4.89, 3.86, 1.14),
III 6.96 (2.31, 2.27, 1.58, 0.80), IV 11.80 (3.28, 4.39, 3.14, 0.99).
Habi us (Fig. 7A–C, 10A). Simila o males, bu he do sum o he abdomen
lacks he p ominen an e io black spo .
Geni alia (Fig. 10B–F). Epigyne s ongly scle o ized wi h la ge pos e o-la -
e al lobes, dis inc ly wide han long, nea ly apezi o m in en al iew and in-
e ed iangula in la e al iew. Knob-shaped p ojec ion (KP) ep esen ed by
a small, pa ly memb anous ube cle, loca ed a an e io po ion o epigynal
pla e. Copula o y openings (CO) la ge, loca ed on he comma-shaped window
(o pocke s wi h chi inous pos e io ma gins) is a he pos e o-la e al po ion
o he epigynal pla e, sepa a ed by indis inc median sep um (MS). Copula o y
duc s (CD) sho , di e ging and ascending obliquely, o ming V-shaped cou se
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Figu e 8. Male palp o he holo ype o Lipoc ea guiyang J. Zhang, Yu & Mi, sp. no . A. Do sal; B. Ven al; C. P ola e al;
D. Re ola e al. Abb e ia ions: C = conduc o ; Cy = cymbium; Em = embolus; HP = hook-shaped p ocess o MA; MA =
median apophysis; Pc = pa acymbium; R = adix; RA = adix apophysis; S = sub egulum; T = egulum; TA I = e minal
apophysis I; TA II = e minal apophysis II; TAA = e minal apophysis appendix; TE = egula ex ension. Scale ba : 0.2 mm.
A B
C D
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Figu e 9. Male palpal bulb o he holo ype o Lipoc ea guiyang J. Zhang, Yu & Mi, sp. no . A. Do sal; B. Ven al; C. P o-
la e al; D. Re ola e al; E. An e io . Abb e ia ions: C = conduc o ; Em = embolus; HP = hook-shaped p ocess o MA; MA
= median apophysis; R = adix; RA = adix apophysis; S = sub egulum; T = egulum; TA I = e minal apophysis I; TA II =
e minal apophysis II; TAA = e minal apophysis appendix; TE = egula ex ension. Scale ba : 0.2 mm.
A B
C D
E
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Figu e 10. F on al iew o cephalo ho ax (A.) and mace a ed epigyne (B–F) o he pa a ype (YHGY428) o Lipoc ea
guiyang J. Zhang, Yu & Mi, sp. no . A. Female; B. La e al; C. Ven al (blue dashed line showing he la e al and pos e io
ma gins o he epigynal base); D. Do sal; E. Ven o-pos e io ; F. Do so-an e io . Abb e ia ions: CD = copula o y duc ; CO
= copula o y opening; FD = e iliza ion duc ; KP = knob-like p ojec ion; MS = median sep um; Sp = spe ma heca. Scale
ba s: 1 mm (A), 0.2 mm (B–F).
A B
C D
E F
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Figu e 11. Epigyne o he pa a ype (YHGY209) o Lipoc ea guiyang J. Zhang, Yu & Mi, sp. no . A. In ac , en al; B. Mac-
e a ed, la e al; C. Mace a ed, en al; D. Mace a ed, do sal; E. Mace a ed, en o-pos e io ; F. Mace a ed, do so-an e io .
Abb e ia ions: CD = copula o y duc ; CO = copula o y opening; FD = e iliza ion duc ; KP = knob-like p ojec ion; MS =
median sep um; Sp = spe ma heca. Scale ba : 0.2 mm.
A B
C D
E F
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Jianshuang Zhang e al.: A su ey o Lipoc ea om Guiyang
in do sal iew, inally en e ing an e io ly loca ed spe ma hecae. Spe ma hecae
(Sp) o al, ~1.2× longe han wide, ela i ely la ge, ~2/3 o epigyne leng h; wo
spe ma hecae close oge he , sepa a ed by ~2/3 o hei wid h. Fe iliza ion
duc s (FD) memb anous, ela i ely long, ~2/3 o spe ma hecae leng h, loca ed
on do sal-basal su ace o spe ma hecae.
Dis ibu ion. Known only om he ype locali y (Fig. 1).
E ymology. The speci ic epi he is de i ed om he name o he ype locali y;
noun in apposi ion.
Commen s. In spi e o he s able mo phology o he male palp and he consis-
en colo a ion o he male habi us, conside able mo phological a ia ion is ob-
se ed among emale indi iduals, p ima ily ela ed o epigynal s uc u es. These
a ia ions in ol e ea u es such as he p esence o absence o a knob-shaped
p ojec ion (KP), he shape o he copula o y openings (CO), and whe he he me-
dian sep um (MS) is dis inc o indis inc . Fo example, in some emales (e.g.,
YHGY428, as in Fig. 10B, C, E), he KP is dis inc , he CO is si ua ed on a com-
ma-shaped window (o wi hin pocke s wi h chi inous pos e io ma gins), and he
MS is indis inc . In con as , in o he indi iduals (e.g., YHGY209, as in Fig. 11A–C,
E), he KP is b oken o , he CO is posi ioned on a nea ly ci cula window (o wi h-
in pocke s bo de ed an e io ly, in e nally, and pos e io ly), and he MS is dis inc .
In addi ion, some a ia ion ela ed o he abdominal pa e n is also obse ed:
he do sum o he abdomen bea s a median band ex ending along i s en i e
leng h in some indi iduals (e.g., YHGY428), whe eas in o he s (e.g., YHGY209),
he median band is es ic ed o he pos e io qua e o he do sum (c . Fig. 7A
and Fig. 7D). Howe e , he mo phological a ia ion was de e mined o be in a-
speci ic a ia ion based on he molecula species delimi a ion analysis.
Lipoc ea usi o mis (Tho ell, 1877)
Figs 1, 3, 12–14
Me a usi o mis Tho ell, 1877: 431 (♀).
Lipoc ea usi o mis: Tho ell 1878: 6.
La inia quad ino a a Simon, 1889: 340 (ju .); Simon 1909: 105 (♀).
La inia lu escens Tho ell, 1898: 342 (♂♀).
La inopa usi o mis: G assho 1970: 231, igs 15a, b, 16a–e (♂♀, ans e om
La inia, synonym o La inia lu escens and L. quad ino a a); Tanikawa 1989:
35, igs 8–14 (♂♀); Chang and Tso 2004: 28, igs 5–8 (♂♀); Tanikawa e al.
2023: 56, igs 15–18.
No e. Fo ull lis o axonomic e e ences, see WSC (2025).
Ma e ial examined. • 1♂, 3♀ (YHGY432, YHGY495, YHGY507, YHGY508), Chi-
na: Guizhou P o .: Guiyang Ci y, Huaxi Dis ic , Uni e si y Town, 26.38°N, 106.65°E,
c. 1173 m, by bea ing, 7 VII 2022, Q. Jiang & Q. Du leg; • 1♂, 1♀ (YHGY431,
YHGY433), Guiyang Ci y, Huaxi Dis ic , Dangwu Town, 26.38°N, 106.60°E, c. 1138
m, by hand, 19 V 2022, H. Yu & Q. Lu leg; Guiyang Ci y, Kaiyang Co., Nanjiang
G and Canyon, 26.94°N, 106.97°E, c. 861 m, by hand, 7 VI 2022, H. Yu & Q. Lu leg.
Diagnosis and desc ip ion. See Tanikawa (1989). Li ing specimens as in
Fig. 3A–C, male habi us as in Fig. 12A–C, male palp as in Fig. 13A–D, emale
habi us as in Fig. 12D–F, geni alia as in Fig. 14A–F.
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Figu e 12. Lipoc ea usi o mis (Tho ell, 1877), male (YHGY431, A–C) and emale (YHGY432, D–F), habi us. A, D. Do sal;
B, E. Ven al; C, F. La e al. Scale ba s: 1 mm (A–C), 2 mm (D–F).
A B C
D E F
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Figu e 13. Male palp o Lipoc ea usi o mis (Tho ell, 1877). A. Do sal; B. Ven al; C. P ola e al; D. Re ola e al. Abb e i-
a ions: C = conduc o ; Cy = cymbium; Em = embolus; HP = hook-shaped p ocess o MA; MA = median apophysis; Pc =
pa acymbium; R = adix; RA = adix apophysis; S = sub egulum; T = egulum; TA = e minal apophysis; TAA = e minal
apophysis appendix; TE = egula ex ension. Scale ba : 0.2 mm.
A B
C D
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Figu e 14 Mace a ed epigynes o emales o Lipoc ea usi o mis (Tho ell, 1877), YHGY432 (A–C) and YHGY507 (D–F).
A. Ven al, wi h scape; B. La e al, wi h scape; C. Ven o-pos e io , wi h scape; D. Ven al, wi hou scape (blue dashed line
showing he la e al and pos e io ma gins o he epigynal base); E. Do sal, wi hou scape; F. Ven o-pos e io , wi hou
scape. Abb e ia ions: CD = copula o y duc ; CO = copula o y opening; FD = e iliza ion duc ; MS = median sep um; Sc =
scape; Sp = spe ma heca. Scale ba s: 0.2 mm (A, B, D–F).
A
B
C
D
E
F
