95
Taxonomic no es on he c ab-spide gene a Nyc imus Tho ell,
1877 and Zame opina Simon, 1909 (A aneae, Thomisidae) wi h
desc ip ions o six new species om Sou heas Asia
Su esh P. Benjamin1, Nau al U i Dhiya’ulhaq2,3 , Ch is a Deeleman-Reinhold4,†, Damayan i Bucho i5,6 ,
Pu nama Hidaya 5, S e an Scheu2,7 , Jochen D esche 2,8
1 Na ional Ins i u e o Fundamen al S udies, Han ana Road, Kandy 20000, S i Lanka
2 Animal Ecology, J.F. Blumenbach Ins i u e o Zoology and An h opology, Uni e si y o Gö ingen, Un e e Ka spüle 2, Gö ingen 37073, Ge many
3 Species Obscu a, Jl. Du en 1 No.2, Depok, Jawa Ba a 16434, Indonesia
4 Ne he lands
5 Depa men o Plan P o ec ion, Facul y o Ag icul u e, IPB Uni e si y, Kampus IPB D amaga Bogo , Jl. Raya D amaga, Wes Ja a 16680, Indonesia
6 Cen e o T ansdisciplina y and Sus ainabili y Sciences, IPB Uni e si y, Kampus IPB Ba anangsiang, Jl. Raya Pajaja an No.27, Wes Ja a 16127, Indonesia
7 Cen e o Biodi e si y and Sus ainable Land Use, Uni e si y o Gö ingen, Büsgenweg 1, Gö ingen 37077, Ge many
8 Senckenbe g Museum o Na u al Sciences Gö li z, Am Museum 1, 02826 Gö li z, Ge many
Co esponding au ho : Jochen D esche ([email p o ec ed])
Copy igh : © Su esh P. Benjamin 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
The c ab-spide gene a Zame opina Simon, 1909 and Nyc imus Tho ell, 1877 emain
poo ly s udied, wi h signi ican gaps in hei axonomic unde s anding. Nyc imus has
seen li le s udy since i s o iginal desc ip ion, and species assignmen s emain unce -
ain due o a lack o mode n diagnos ic da a. By con as , Zame opina, his o ically mono-
ypic, was ecen ly expanded wi h a second species (Z. wanliae Lin & Li, 2023) ollowing
a edesc ip ion o he ype species, Z. calcea a Simon, 1909. Ou s udy p esen s a e iew
o hese gene a, p ima ily based on specimens collec ed om opical Asia by he la e
Ch is a L. Deeleman-Reinhold as well as om he EFFo TS p ojec in he lowlands o Su-
ma a, Indonesia. The e iew includes he desc ip ion o he ollowing six new species:
Nyc imus alca us Benjamin & Dhiya’ulhaq, sp. no . (♂), Nyc imus kinabaluensis Benja-
min & Dhiya’ulhaq, sp. no . (♂♀), Nyc imus mu illoides Benjamin & Dhiya’ulhaq, sp. no .
(♂), Nyc imus quad ipunc a us Dhiya’ulhaq & Benjamin, sp. no . (♂♀), Nyc imus endang
Dhiya’ulhaq & Benjamin, sp. no . (♂♀), and Nyc imus saksang Dhiya’ulhaq & Benjamin,
sp. no . (♂). Nyc imus bis ia us Tho ell, 1877 is edesc ibed and new locali y eco ds
a e p o ided. Zame opina wanliae is eco ded o he i s ime ou side o Vie nam. Fu -
he , upda ed species desc ip ions o Nyc imus and Zame opina a e p o ided, along
wi h axonomic cla i ica ions o he gene a. This s udy emphasises he undamen al
impo ance o well-cu a ed, his o ical collec ions and ongoing ieldwo k in esol ing he
sys ema ics o lesse -known a achnid lineages.
Key wo ds: A achnida, biodi e si y, RTA-clade, axonomy
Academic edi o : F ancesco Balla in
Recei ed:
9 May 2025
Accep ed:
18 Augus 2025
Published:
9 Oc obe 2025
ZooBank: h ps://zoobank.
o g/584332C1-8B4B-41B9-A17D-
327F47B5616A
Ci a ion: Benjamin SP, Dhiya’ulhaq
NU, Deeleman-Reinhold C, Bucho i D,
Hidaya P, Scheu S, D esche J (2025)
Taxonomic no es on he c ab-spide
gene a Nyc imus Tho ell, 1877 and
Zame opina Simon, 1909 (A aneae,
Thomisidae) wi h desc ip ions o six
new species om Sou heas Asia.
ZooKeys 1255: 95–125. h ps://doi.
o g/10.3897/zookeys.1255.158380
ZooKeys 1255: 95–125 (2025)
DOI: 10.3897/zookeys.1255.158380
† Deceased.
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ZooKeys 1255: 95–125 (2025), DOI: 10.3897/zookeys.1255.158380
Su esh P. Benjamin e al.: Taxonomic no es on he c ab-spide gene a Nyc imus Tho ell, 1877 and Zame opina Simon, 1909
In oduc ion
Zame opina Simon, 1909, is a genus o c ab spide s belonging o he amily
Thomisidae. Re e ed o as c ab spide s, due o hei elonga ed i s wo leg
pai s and hei habi o mo ing sideways, Thomisidae a e among he mos
species- ich spide amilies wi h mo e han 2,100 species ac oss 171 gene a
(Wo ld Spide Ca alog 2025). Despi e i s inclusion in his di e se g oup, Zame-
opina emains ela i ely obscu e and axonomically un e ised. Ini ially, he ge-
nus comp ised only a single species, Zame opina calcea a Simon, 1909, which
had been inadequa ely unde s ood o decades due o a lack o illus a ions
in he o iginal desc ip ion. A signi ican upda e came wi h he wo k o Tang
e al. (2010), who e isi ed and edesc ibed he ype species. Th ough me ic-
ulous examina ion o he ype specimen, hey p o ided de ailed illus a ions
and desc ip ions, he eby cla i ying he diagnos ic ea u es o he genus and
aiding in i s axonomic ci cumsc ip ion. This e o signi ican ly enhanced he
unde s anding o Zame opina and se he s age o u u e axonomic s udies.
Recen ly, a second species, Zame opina wanliae Lin & Li, 2023, was desc ibed
in Lin e al. (2023), u he en iching he di e si y o he genus.
By con as , he genus Nyc imus Tho ell, 1877, has ecei ed limi ed axo-
nomic sc u iny since i s ini ial desc ip ion. His o ically, Nyc imus included wo
species: he ype species Nyc imus bis ia us Tho ell, 1877, om Indonesia
(Suma a and Sulawesi), and Nyc imus imeni (Simon, 1895) om Sou h A i-
ca. O iginally desc ibed as Zame opias imeni, he la e is likely misplaced,
indica ing a need o axonomic e ision. Addi ionally, Zame opias specula o
Tho ell, 1892, was ans e ed o Nyc imus and synonymised wi h N. bis ia us
by Leh inen (2016). Howe e , his ans e was made wi hou accompanying
illus a ions o a comp ehensi e e-examina ion o he ype ma e ial, lea ing
he axonomy o bo h species inadequa ely esol ed.
This pape aims o add ess some o hese gaps by e iewing he poo ly known
species o bo h Zame opina and Nyc imus, p ima ily d awing on specimens col-
lec ed om opical Asia by he la e Ch is a L. Deeleman-Reinhold, as well as
om he EFFo TS p ojec in he lowlands o Jambi P o ince, Suma a, Indonesia
(D esche e al. 2016). This e iew aims o cla i y he axonomy and p o ide up-
da ed desc ip ions, con ibu ing o a mo e obus unde s anding o hese gene a
and hei oles in he biodi e si y o he egions hey inhabi . The s udy highligh s
he impo ance o e isi ing his o ical collec ions and unde sco es he need o
con inued explo a ion and documen a ion o lesse -known spide axa.
Ma e ials and me hods
Sample collec ion
Specimens om he la e Ch is a L. Deeleman-Reinhold’s pe sonal collec ion,
housed a he esidence, we e selec ed by Su esh P. Benjamin (SPB) and ans-
e ed o he Zoological Resea ch Museum Alexande König (Bonn, Ge many,
ZFMK) o his s udy. These specimens, along wi h he es o he collec ion,
will ul ima ely be deposi ed a Na u alis Biodi e si y Cen e , Leiden, Ne he lands
(RMNH). Spide s om Jambi P o ince we e pa o a collec ion o mo e han hal
a million canopy a h opods (Pollie e e al. 2023) sampled by canopy ogging
97
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Su esh P. Benjamin e al.: Taxonomic no es on he c ab-spide gene a Nyc imus Tho ell, 1877 and Zame opina Simon, 1909
ac oss ain o es , jungle ubbe , and monocul u es o ubbe and oil palm in he
lowlands o Jambi, Suma a, Indonesia as pa o he Ge man-Indonesian EFFo TS
esea ch p ojec (D esche e al. 2016). O e all, mo e han 10,000 spide indi idu-
als om a leas 30 amilies and 400 species we e cap u ed (Ramos e al. 2022;
Dhiya’ulhaq e al. 2024). The ogging me hod is de ailed in Pollie e e al. (2023)
and he EFFo TS plo design and gene al s udy aims in D esche e al. (2016).
Iden i ica ion and pho og aphy
The gene al me hodology ollows Benjamin (2011) and Ilepe uma A achchi and
Benjamin (2019). Specimens used o habi us illus a ions we e placed in 70%
E OH and pho og aphed wi h a Zeiss AxioCam HRc came a moun ed on a dis-
sec ing mic oscope (Zeiss Disco e y V20) wi h op illumina ion and a magni-
ica ion o up o 150×. Images we e edi ed using he Zeiss ZEN P o so wa e
package. Unless o he wise s a ed, depic ed s uc u es a e o he le male palp.
Al e na i ely, some specimens we e examined unde a ZEISS S emi 2000 mic o-
scope. Female geni alia we e excised om he specimen’s body and hen clea ed
wi h 10% KOH o a leas one hou o examine he in e nal copula o y o gans.
Imaging he specimens was done using a Keyence VHX-7000 digi al mic oscope
sys em o a Zeiss AxioCam HRc came a moun ed on a dissec ing mic oscope
(Zeiss Disco e y V20). The desc ip ion o colou a ion is based on specimens
in e hanol. Measu emen s o legs a e gi en as o al leng h ( emu , pa ella, ibia,
me a a sus, a sus). Missing legs o leg segmen s a e ma ked wi h “-”, and legs
wi h missing segmen s do no ha e hei o al leng h eco ded. Unmen ioned
legs did no ha e hei leng h measu ed. All measu emen s a e in millime e .
Abb e ia ions
ALE an e io la e al eye
AME an e io median eye
CO copula o y opening
FD e iliza ion duc
PLE pos e io la e al eye
PME pos e io median eye
RTA e ola e al ibial apophysis o
male palp
VTA en al ibial apophysis o
male palp
Reposi o ies
GOET Animal Ecology G oup, J.-F. Blumenbach Ins i u e o Zoology and An-
h opology, Uni e si y o Gö ingen, Gö ingen, Ge many (Jochen D e-
sche , S e an Scheu)
MCSN Museo Ci ico di S o ia Na u ale Giacomo Do ia, Genoa, I aly (Ma ia
Ta ano)
MCZ Museum o Compa a i e Zoology, Massachuse s (Gonzalo Gi ibe )
MNHN Muséum Na ional d’His oi e Na u elle, Pa is (Ch is ine Rolla d)
MZB Museum Zoologicum Bogo iense, Bogo , Indonesia (Cahyo Rahmadi)
RMNH Na u alis Biodi e si y Cen e , Leiden (Ka en an Do p)
SMF Na u museum Senckenbe g, F ank u a. M., Ge many (Jana G üge ,
Pe e Jäge )
ZMH Zoologisches Museum Hambu g, Hambu g, Ge many (Nadine Dupé é,
Danilo Ha ms)
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Su esh P. Benjamin e al.: Taxonomic no es on he c ab-spide gene a Nyc imus Tho ell, 1877 and Zame opina Simon, 1909
Resul s
Family Thomisidae Sunde all, 1833
Nyc imus Tho ell, 1877
Type species. Nyc imus bis ia us Tho ell, 1877.
Diagnosis. Small homisids, 2–5 mm in body leng h. P osoma cuboid, as
long as wide, wi h g anula ed su ace and bea ing long mac ose ae (Fig. 1).
Body colou a ion da k wi h whi e/yellow ma kings, mos ly in he o m o spo s
and ba s. Opis hosoma wi h i e la ge sigillae a anged in a iangula o ma-
ion Embolus is weakly cu ed and possesses a hick base. Tegulum wi h a
“hood” (Fig. 3A). Tibial apophyses a e well de eloped, VTA p esen , ei he ee
(Fig. 3A) o hook-shaped and a ached o RTA (Fig. 7A). Epigynum highly a i-
able, spe ma heca ounded o o al, single pai (Fig. 3C, D).
Species composi ion. Nyc imus bis ia us Tho ell, 1877 (♂♀), Nyc imus al-
ca us Benjamin & Dhiya’ulhaq, sp. no . (♂), Nyc imus kinabaluensis Benjamin
& Dhiya’ulhaq, sp. no . (♂♀), Nyc imus mu illoides Benjamin & Dhiya’ulhaq, sp.
no . (♂), Nyc imus quad ipunc a us Dhiya’ulhaq & Benjamin, sp. no . (♂♀), Nyc-
imus endang Dhiya’ulhaq & Benjamin, sp. no . (♂♀), Nyc imus saksang Dhi-
ya’ulhaq & Benjamin, sp. no . (♂), Nyc imus imeni (Simon, 1895).
Nyc imus bis ia us Tho ell, 1877
Figs 1–3
Nyc imus bis ia us Tho ell, 1877: 499; Leh inen 2016: 166 (as Nyc imus bino-
a us, lapsus).
Zame opias specula o Tho ell, 1892: 123.
Type ma e ial. Holo ype. Indonesia – Sulawesi, Sou heas Sulawesi P o ince •
♂; Kenda i; 1874; O. Becca ii leg.; holo ype o Nyc imus bis ia us Tho ell, 1877;
MCSN. Examined.
O he ma e ial examined. B unei – Tu ong Dis ic • 1♂; Buki Sulang; 1982;
N. S o k leg.; canopy ogging in p ima y o es ; RMNH RMNH.ARA.17873.
Indonesia – Suma a, Jambi P o ince • 1♀; Sa olangun, Buki Duabelas Na-
ional Pa k; 01°58'55.2"S, 102°45'02.6"E; ele . 73 m; 7 Oc . 2013; J. D esche
leg.; canopy ogging in ain o es ; GOET 2013_BF2.1_A aThom051N_001 ( o
be ans e ed o MZB). Malaysia – Sabah S a e • 1♂; Kinabalu Na ional Pa k,
Po ing Ho Sp ings; 05°59'N, 116°42'E; ele . 600–700 m; 27 Feb. 1996; A. Flo-
en leg.; canopy ogging on Apo osa lagenoca pa ee, in p ima y o es ; RMNH
RMNH.ARA.17874. • 1♀; Kinabalu Na ional Pa k, Po ing Ho Sp ings; 05°59'N,
116°42'E; ele . 600–700 m; 28 Ma . 1998; A. Flo en leg.; canopy ogging on
Ba ing onia ee, in p ima y o es ; RMNH RMNH.ARA.17875. • 1♂; Kinabalu
Na ional Pa k, Po ing Ho Sp ings; 05°59'N, 116°42'E; ele . 600–700 m; 28 Ma .
1998; A. Flo en leg.; canopy ogging on Dac yodes laxa ee, in p ima y o es ;
RMNH RMNH.ARA.17876.
Diagnosis. Males can be easily dis inguished om all o he congene s by he
longe embolus, mo e han leng h o cymbium (in all congene s, he embolus
is a mos hal he leng h o cymbium; Fig. 3A, B) and he den icula e RTA. Fe-
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Su esh P. Benjamin e al.: Taxonomic no es on he c ab-spide gene a Nyc imus Tho ell, 1877 and Zame opina Simon, 1909
Figu e 1. Nyc imus bis ia us Tho ell, 1877 om Bo neo; habi us, do sal iew. A. Male (RMNH.ARA.17873); B. Female
(RMNH.ARA.17875). Scale ba s: 0.5 mm.
males can be easily dis inguished by he single CO; CD a ched, esembling a
ams’ ho n, mo e han wice he leng h o spe ma hecae (in all congene s, he
CO is pai ed, and he CD a e no longe han spe ma hecae; Figs 2C, D, 3C, D).
Desc ip ion. Male (RMNH.ARA.17873; Figs 1A, 3A, B). To al leng h 2.85. P o-
soma leng h 1.42; wid h 1.26. Opis hosoma leng h 1.43; wid h 1.17. Diame e
o eyes: AME 0.09; ALE 0.15; PLE 0.14 PME 0.07. In e dis ances be ween eyes:
AME–AME 0.50; AME–ALE 0.10; ALE–ALE 0.86; PME–PME 0.70; PME–PLE
0.20; ALE–PLE 0.18; AME–PME 0.13; PLE–PLE 1.09. Leg measu emen s: leg
I 3.3 (1.0, 0.5, 1.0, 0.8, 0.5). P osoma cuboid, almos as long as wide; do sal-
ly blackish b own, su ace g anula ed; la e ally wi h wo ows o long, cu ed
mac ose ae o igina ing on small ube cles. All eyes a e e y a apa , especial-
ly be ween he median eyes. Chelice ae, labium and s e num blackish b own;
s e num wi h small den s. Legs da k b own excep emo a III and IV, which a e
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Su esh P. Benjamin e al.: Taxonomic no es on he c ab-spide gene a Nyc imus Tho ell, 1877 and Zame opina Simon, 1909
Figu e 2. Nyc imus bis ia us Tho ell, 1877 emale om Suma a (2013 BF2.1 A aThom051N 001). A. Habi us, do sal
iew; B. P osoma, on al iew; C. Epigynum, en al iew; D. Epigynum, do sal iew, clea ed. Scale ba s: 1 mm (A);
0.5mm (B); 0.2 mm (C, D).
yellow; emo a and ibiae I–IV wi h wo long spines; pa ellae I–IV wi h one dis-
al long spine. Opis hosoma do sally g eyish black wi h many long se ae; i e
la ge sigillae p esen , a anged in a iangula o ma ion; en e g eyish black.
Palp (Fig. 3A, B): cymbium o al. Embolus long, whip-like, looping o e he
e o la e al side o bulb, hal o i s leng h co e ed by he cymbium. Tegula
hood small, loca ed close o he base o bulb. RTA la ge, oughly apezoid,
la e ally p o uding, dis al hal wi h den icles on all sides. VTA long, digi i o m;
apically lexed wi h ip en ally cu ed.
Female (RMNH.ARA.17875; Figs 1B, 3C, D). To al leng h 3.32. P osoma
leng h 1.15; wid h 1.30. Opis hosoma leng h 2.17; wid h 1.95. Diame e o eyes:
AME 0.08; ALE 0.13; PLE 0.16; PME 0.06. In e dis ances be ween eyes: AME–
AME 0.56; AME–ALE 0.10; ALE–ALE 0.86; PME–PME 0.64; PME–PLE 0.21;
ALE–PLE 0.18; AME–PME 0.16; PLE–PLE 1.09. Leg measu emen s: leg I 5.3
(1.6, 0.6, 1.3, 1.0, 0.8). Habi us as in male, excep opis hosoma do sally ligh e
han male and he p esence o whi e spo s as shown in Fig. 1B.
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Su esh P. Benjamin e al.: Taxonomic no es on he c ab-spide gene a Nyc imus Tho ell, 1877 and Zame opina Simon, 1909
Figu e 3. Nyc imus bis ia us Tho ell, 1877 om Bo neo. A. Male (RMNH.ARA.17873), le palp, en al iew; B. ibid.,
e o la e al iew; C. Female (RMNH.ARA.17875), epigynum; C. ibid., en al iew; D. ibid., do sal iew. Abb e ia ions: CD=
copula o y duc ; CO = copula o y opening; E = embolus; FD = e iliza ion duc ; RTA = e ola e al ibial apophysis; S =
spe ma heca; TH = egula hood; VTA = en al ibial apophysis. Scale ba s: 0.2 mm.
Epigynum (Figs 2C, D, 3C, D): ex e io pla e oughly pen agonal. A ium
inconspicuous. CO o al, single, la ge, CD long; di ided in o a e y wide, ams-
ho n-shaped ini ial pa and a na ow, C-shaped la e pa . Spe ma heca
globula , app oxima ely as wide as CO; an e io ly wi h a small bump; pos e-
io ly ex ended.
Dis ibu ion. B unei, Malaysia (Bo neo: Sabah), Indonesia (Suma a: Jambi;
Sulawesi: Sou heas Sulawesi) (Fig. 19).
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Su esh P. Benjamin e al.: Taxonomic no es on he c ab-spide gene a Nyc imus Tho ell, 1877 and Zame opina Simon, 1909
Nyc imus alca us Benjamin & Dhiya’ulhaq, sp. no .
h ps://zoobank.o g/8DF08156-9B26-4343-95A5-641924B7AC50
Fig. 4
E ymology. The species epi he alca us is de i ed om he La in wo d alx/ al-
cis, meaning “sickle” o “scy he.” I e e s o he dis inc ly cu ed, sickle-shaped
embolus o he male palp, a diagnos ic ea u e o he species.
Type ma e ial. Holo ype. Thailand – Phang Nga P o ince • ♂; Khao Lak Na-
ional Pa k; no mo e label da a; RMNH RMNH.ARA.17861.
Diagnosis. Males a e e y simila o N. saksang sp. no ., in ha ing a simi-
la sickle-shaped embolus, bu hey can be easily dis inguished om i by he
Figu e 4. Nyc imus alca us Benjamin & Dhiya’ulhaq, sp. no ., male holo ype (RMNH.ARA.17861). A. Habi us, do sal iew;
B. Le palp, en al iew; C. Le palpal ibia, e ola e al iew. Abb e ia ions: E = embolus; RTA = e ola e al ibial apoph-
ysis; TH = egula hood; VTA = en al ibial apophysis. Scale ba s: 0.5 mm (A); 0.2 mm (B, C).
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Su esh P. Benjamin e al.: Taxonomic no es on he c ab-spide gene a Nyc imus Tho ell, 1877 and Zame opina Simon, 1909
la ge, digi i o m VTA, a ached dis ally on he RTA ( he co esponding pa in N.
saksang, sp. no ., is much smalle and a ached medially on he en al lobe o
RTA; Fig. 4B s Figs 16C, 17A).
Desc ip ion. Male (holo ype RMNH.ARA.17861; Fig. 4). To al leng h: 2.36.
P osoma leng h 1.23; wid h: 1.05. Opis hosoma leng h 1.13; wid h 1.02. Diam-
e e o eyes: AME 0.08; ALE 0.15; PLE 0.14; PME 0.05. In e dis ances be ween
eyes: AME–AME 0.27; AME–ALE 0.08; ALE–ALE 0.59; PME–PME 026; PME–
PLE 0.27; ALE–PLE 0.18; AME–PME 0.14; PLE–PLE 0.79. P osoma cuboid, al-
mos as long as wide, da k eddish b own; su ace weakly g anula ed; la e ally
ado ned wi h long mac ose ae, each a ising om a small ube cle. Chelice ae
and s e num colou ed as p osoma. F on legs b own; back legs b own excep
o he whi e emo a III and IV. Opis hosoma ound, ligh b own; do sally co -
e ed wi h scu um; i e la ge sigillae p esen , a anged in a iangula o ma ion.
Palp (Fig. 4B, C): cymbium o al. Embolus sickle-shaped. Tegulum ound, wi h
a small iangula p ojec ion nex o he embolus; egula hood small. RTA wide
apezoid; VTA la ge, digi i o m, a ached dis ally on he RTA.
Female. Unknown.
Dis ibu ion. Only known om he ype locali y (Thailand: Phang Nga) (Fig. 19).
Rema ks. The specimen is pa ially damaged: bo h i s legs a e b oken and
incomple e, and he emaining legs a e mos ly missing o agmen ed.
Nyc imus kinabaluensis Benjamin & Dhiya’ulhaq, sp. no .
h ps://zoobank.o g/1F2D858B-2FED-4397-BE73-9B841300D782
Figs 5–7
E ymology. The speci ic epi he is de i ed om he name o he ype locali y.
Type ma e ial. Holo ype. Malaysia – Sabah S a e • ♂; Kinabalu Na ional
Pa k, Po ing Ho Sp ings; 05°59'N, 116°42'E; ele . 600–700 m; 27 Feb. 1996;
A. Flo en leg.; canopy ogging on Ba ing onia ee, in p ima y o es ; RMNH
RMNH.ARA.17862.
O he ma e ial examined. B unei – Tu ong Dis ic • 1♀; Lamunin, Buki Sulang;
1982; N. S o k leg.; canopy ogging; RMNH RMNH.ARA.17859. Indonesia – Suma-
a, Jambi P o ince • 1♀; Sa olangun, Buki Duabelas Na ional Pa k; 01°58'55.2"S,
102°45'02.6"E; ele . 73 m; 7 Oc . 2013; J. D esche leg.; canopy ogging in ain o -
es ; GOET 2013_BF2.1_A aThom004N_001 ( o be ans e ed o MZB).
Diagnosis. Males can be easily dis inguished om all o he congene s by he
embolic base ab up ly ben 90° owa ds he e ola e al side o he cymbium and
he ape ed, ben embolus (Fig. 7A, B). Females a e dis inguishable by he long,
o al spe ma heca (globula o sho -o al in all o he congene s; Figs 6C, D, 7C, D).
Desc ip ion. Male (holo ype RMNH.ARA.17862; Figs 5A, B, 7A, B). To-
al leng h 3.24. P osoma leng h 1.61; wid h 1.50. Opis hosoma leng h 1.63;
wid h 1.31. Diame e o eyes: AME 0.11; ALE 0.20; PLE 0.17; PME 0.08. In-
e dis ances be ween eyes: AME–AME 0.44; AME–ALE 0.11; ALE–ALE 0.81;
PME–PME 0.45; PME–PLE 0.36; ALE–PLE 0.21; AME–PME 0.17; PLE–PLE
1.15. Leg measu emen s: leg I 5.0 (1.5, 0.6, 1.3, 1.0, 0.6). P osoma cuboid, al-
mos as long as wide, da k eddish b own; su ace weakly g anula ed; la e ally
ado ned wi h long mac ose ae, each a ising om a small ube cle. Chelice ae
and s e num colou ed as p osoma. Endi es b own wi h whi e ip. F on legs
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Figu e 10. Nyc imus quad ipunc a us Dhiya’ulhaq & Benjamin, sp. no ., emale pa a ype (A aThom001N_2013_HF3.2_001).
A. Habi us, en al iew; B. Habi us, do sal iew; C. Epigynum, en al iew; D. Epigynum, do sal iew, clea ed. Scale ba s:
1 mm (A, B); 0.2 mm (C, D).
0.36); leg IV 2.74 (0.83, 0.27, 0.66, 0.52, 0.46). P osoma cuboid, almos as long
as wide, box-shaped, weakly g anula ed, highes a mid-poin , ligh b own; la e al
ma gins u nished wi h a ow o long mac ose ae. La e al eyes la ge, si ua ed
on ube cles on he la e al edge o he p osoma. Chelice ae, endi es, and labium
ligh b own. S e num ligh b own. Legs I ligh b own; o he s yellow-b own, wi h
spa se spines and black se ae. Opis hosoma o al, b own, wi h wo pai s o la ge
whi e spo s. Spinne e s b own.
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Figu e 11. Nyc imus quad ipunc a us Dhiya’ulhaq & Benjamin, sp. no . A. Male holo ype (A aThom001N_2013_HF3.2_001),
igh palp, mi o ed, en al iew; B. ibid., RTA, en o- e ola e al iew; C. Female pa a ype (A aThom001N_2013_
HF3.2_001), epigynum, en al iew; D. ibid., do sal iew. Abb e ia ions: CD = copula o y duc ; CO = copula o y opening; E=
embolus; RTA = e ola e al ibial apophysis; S = spe ma heca; TH = egula hood; VH = en al hook. Scale ba s: 0.2 mm.
Palp (Figs 9C, D, 11A, B): cymbium o al. Embolus sickle-shaped; base sligh ly
w inkled. Tegula hood e y wide. RTA la ge, ec angula , an il-shaped; do sally
bi u ca ed, ending in o wo long, acu e lobes, hidden behind a b ush o se ae;
VTA hook-shaped a ached medially on RTA.
Female (pa a ype 2013_HF3.2_A aThom001N_002; Figs 10, 11C, D). To al
leng h 4.81. P osoma leng h 1.95; wid h 1.77. Opis hosoma leng h 2.86; wid h
2.65. Diame e o eyes: AME 0.09; ALE 0.20; PLE 0.18; PME 0.07. In e dis ances
be ween eyes: AME–AME 0.53; AME–ALE 0.14; ALE–ALE 0.97; PME–PME
0.52; PME–PLE 0.39; ALE–PLE 0.23; AME–PME 0.24; PLE–PLE 1.37. Leg mea-
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Figu e 12. Nyc imus quad ipunc a us Dhiya’ulhaq & Benjamin, sp. no ., non- ype male specimen om Bo neo (RMNH.
ARA.17857). A. Habi us, do sal iew; B. Le palp, en al iew; C. Le palpal ibia, e ola e al iew. Abb e ia ions: E =
embolus; RTA = e ola e al ibial apophysis; TH = egula hood; VH = en al hook. Scale ba s: 0.5 mm (A); 0.2 mm (B, C).
su emen s: leg I 5.47 (1.66, 0.57, 1.34, 1.13, 0.77); leg II -; leg III 3.69 (1.22, 0.47,
0.87, 0.63, 0.50); leg IV 4.12 (1.36, 0.46, 0.96, 0.76, 0.58). Habi us as in males,
excep p osoma and leg I eddish b own.
Epigynum (Figs 10C, D, 11C, D): ex e io pla e la ge, hexagonal-shaped wi h
wide an e io side. A ium conspicuous, in e ed hea -shaped wi h a sepa a-
ion in he middle. CO diagonally o ien ed, pos e io ly acing. CD a ch-shaped.
Spe ma heca elonga ed, peanu -shaped, wi h a small bump an e io ly.
Dis ibu ion. Malaysia (Bo neo: Sabah); Indonesia (Suma a: Jambi) (Fig. 19).
Rema ks. Specimens om Bo neo show sligh di e ences in shape o em-
bolus and do sal ip o RTA (Fig. 12B, C).
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Nyc imus endang Dhiya’ulhaq & Benjamin, sp. no .
h ps://zoobank.o g/D674C659-6FE3-41A2-9484-300E4C2A918D
Figs 13–15
E ymology. The speci ic epi he e e s o a mea dish associa ed wi h he cui-
sine o he Minangkabau people om Wes Suma a. The mea is slow-cooked
in a ple ho a o spices, gi ing i a da k colou a ion. The da k colou o N. en-
dang, sp. no ., is eminiscen o he colou o his dish.
Type ma e ial. Holo ype. Indonesia – Suma a, Jambi P o ince • ♂; Sa olan-
gun, Ai Hi am, Desa Ba u; 02°01'49.5"S, 102°46'14.8"E; ele . 57 m; 12 Jul. 2013;
J. D esche leg.; canopy ogging in jungle ubbe plan a ion; GOET 2013_BJ6.1_
A aThom010N_001 ( o be ans e ed o MZB). Pa a ypes. Indonesia – Suma-
a, Jambi P o ince • 1♂1♀; same da a as holo ype; GOET 2013_BJ6.1_A aTh-
om010N_002, 003 ( o be ans e ed o MZB). • 2♂♂; Ba ang Ha i, Bajubang,
Singkawang; 01°47'07.9"S, 103°16'37.4"E; ele . 56 m; 18 Jun. 2013; J. D esche
leg.; canopy ogging in jungle ubbe plan a ion; GOET 2013_HJ4.2_A aTh-
om010N_001, 002 ( o be ans e ed o ZMH).
O he ma e ial examined. Indonesia – Suma a, Jambi P o ince • 1♀;
Ba ang Ha i, Hu an Ha apan Conse a ion A ea; 02°11'15.3"S, 103°20'36.0"E;
ele . 69 m; 7 Aug. 2013; J. D esche leg.; canopy ogging in ain o es ; GOET
2013_HF4.1_A aThom036N_001.
Diagnosis. Males a e simila o N. quad ipunc a us, sp. no ., in ha ing a sick-
le-shaped embolus and in he gene al shape o he bulb, bu he VTA is digi i o m
and ee (Fig. 15A, B; s hook-shaped, a ached o RTA in N. quad ipunc a us,
Fig. 11A, B) and he RTA possesses a pilla -shaped en al lobe and pai ed, hick
spines ( s bo h absen ). Females a e dis inc in ha ing he spe ma hecae wi h a
pos e io ex ension and a sho , ba ely no iceable CD (Fig. 11D).
Desc ip ion. Male (holo ype 2013_BJ6.1_A aThom010N_001; Figs 13, 15A,
B). To al leng h 2.41. P osoma leng h 1.16; wid h 1.03. Opis hosoma leng h
1.25; wid h 1.21. Diame e o eyes: AME 0.07; ALE 0.15; PLE 0.14; PME 0.05.
In e dis ances be ween eyes: AME–AME 0.35; AME–ALE 0.08; ALE–ALE 0.62;
PME–PME 0.32; PME–PLE 0.24; ALE–PLE 0.17; AME–PME 0.19; PLE–PLE
0.77. Leg measu emen s: leg I 3.36 (0.90, 0.29, 0.92, 0.75, 0.50); leg II 3.46
(1.04, 0.32, 0.84, 0.75, 0.51); leg III 2.11 (0.67, 0.20, 0.54, 0.38, 0.32); leg IV 2.25
(0.75, 0.18, 0.59, 0.42, 0.31). P osoma cuboid, almos as long as wide, da k ed-
dish b own; su ace weakly g anula ed; la e ally ado ned wi h long mac ose ae,
each a ising om a small ube cle. Chelice ae, and s e num colou ed as p o-
soma. Endi es b own wi h whi e ip. F on legs b own; back legs b own excep
o he whi e emo a. Opis hosoma ound, da k b own wi h pai ed whi e ba s in
he middle; do sally co e ed wi h scu um; i e la ge sigillae p esen , a anged
in a iangula o ma ion.
Palp (Figs 13C, D, 15A, B): cymbium o al. Embolus sickle-shaped. Tegulum
wi h a small iangula p ojec ion nex o he embolus; egula hood small. RTA
bilobed; en al lobe inge -shaped; do sal lobe wide apezoid, apically wi h a
pai o sho , hick spines. VTA long, inge -shaped, apically lexed.
Female (pa a ype A aThom010N_2013_BJ6.1_003; Figs 14, 15C, D). To al
leng h 3.86. P osoma leng h 1.67; wid h 1.43. Opis hosoma leng h 2.19; wid h
2.21. Diame e o eyes: AME 0.08; ALE 0.18; PLE 0.19; PME 0.07. In e dis ances
be ween eyes: AME–AME 0.55; AME–ALE 0.12; ALE–ALE 0.90; PME–PME
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Figu e 13. Nyc imus endang Dhiya’ulhaq & Benjamin, sp. no ., male holo ype (A aThom010N_2013_BJ6.1_001). A. Habi-
us, do sal iew; B. Habi us, en al iew; C. Le palp, en al iew; D. Le palp, e ola e al iew. Scale ba s: 1 mm (A, B);
0.2 mm (C, D).
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Figu e 14. Nyc imus endang Dhiya’ulhaq & Benjamin, sp. no ., emale pa a ype (A aThom010N 2013 BJ6.1 003). A. Habi-
us, do sal iew; B. Habi us, en al iew; C. Epigynum, en al iew; D. Epigynum, do sal iew; E. Epigynum, do sal iew,
clea ed. Scale ba s: 1 mm (A, B); 0.2 mm (C–E).
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Figu e 15. Nyc imus endang Dhiya’ulhaq & Benjamin, sp. no . A. Male holo ype (A aThom010N 2013 BJ6.1 001), le
palp, en al iew; B. ibid., le palp, e ola e al iew; C. Female pa a ype (A aThom010N_2013_BJ6.1_003), epigynum,
do sal iew; D. ibid., epigynum, en al iew. Abb e ia ions: CO = copula o y opening; E = embolus; PE = pos e io ex en-
sion o spe ma heca; RTA = e ola e al ibial apophysis; SP = spine; S = spe ma heca; TH = egula hood; VTA = en al
ibial apophysis. Scale ba s: 0.2 mm.
0.45; PME–PLE 0.34; ALE–PLE 0.23; AME–PME 0.23; PLE–PLE 1.11. Leg mea-
su emen s: leg I 4.85 (0.71, 1.01, 1.18, 0.49, 1.46); leg II -; leg III 3.05 (1.00, 0.30,
0.75, 0.54, 0.46); leg IV 3.41 (1.14, 0.32, 0.83, 0.65, 0.47). Habi us as in male,
excep opis hosoma no co e ed in scu um.
Epigynum (Fig. 14C, D, 15C, D): ex e io pla e wi h i egula bo de . A ium in-
conspicuous. CO semici cula , pos e io ly acing. CD e y sho . Spe ma heca
o al, an e io ly wi h a la ge bump, pos e io ly ex ended.
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Dis ibu ion. Indonesia (Suma a: Jambi) (Fig. 19).
Rema ks. One specimen shows ounded a he han o al spe ma hecae (Fig.
14E) and is, hus, en a i ely placed he e, as no males om he same locali y
we e ound.
Nyc imus saksang Dhiya’ulhaq & Benjamin, sp. no .
h ps://zoobank.o g/D731CC5A-13CF-480B-9894-01BCC9A42F9E
Figs 16, 17
E ymology. The speci ic epi he e e s o a mea -s ew dish associa ed wi h he
Ba ak people om No h Suma a. The mea is cooked in blood, gi ing i a da k
colou a ion. The da k colou o Nyc imus saksang Benjamin & Dhiya’ulhaq, sp.
no ., is eminiscen o he colou o his dish.
Type ma e ial. Holo ype. Indonesia – Suma a, Jambi P o ince • ♂; Ba ang
Ha i, Hu an Ha apan Conse a ion A ea; 02°09'48.9"S, 103°20'04.4"E; ele .
73 m; 6 Jul. 2013; J. D esche leg.; canopy ogging in ain o es ; GOET 2013_
HF2.1_A aThom036N_001 ( o be ans e ed o MZB).
Diagnosis. Males can be easily dis inguished om all o he congene s by he
e y wide, axe-shaped RTA which consis s o h ee la ge lobes (Figs 16C, D, 17).
Desc ip ion. Male (holo ype 2013_HF2.1_A aThom036N_001; Figs 16, 17).
To al leng h 3.11. P osoma leng h 1.50; wid h 1.25. Opis hosoma leng h 1.61;
wid h 1.40. Diame e o eyes: AME 0.08; ALE 0.17; PLE 0.14; PME 0.05. In e -
dis ances be ween eyes: AME–AME 0.40; AME–ALE 0.10; ALE–ALE 0.73;
PME–PME 0.38; PME–PLE 0.33; ALE–PLE 0.21; AME–PME 0.22; PLE–PLE
0.95. Leg measu emen s: leg I 4.20 (1.20, 0.44, 1.05, 0.89, 0.62); leg II 3.74
(1.10, 0.34, 0.91, 0.78, 0.61); leg III 2.78 (0.83, 0.37, 0.67, 0.52, 0.39); leg IV 2.87
(0.88, 0.26, 0.70, 0.60, 0.43). P osoma cuboid, almos as long as wide, da k ed-
dish b own; su ace weakly g anula ed; la e ally ado ned wi h long mac ose ae,
each a ising om a small ube cle. Chelice ae and s e num colou ed as p o-
soma. Endi es b own wi h whi e ip. F on legs b own; back legs b own excep
o he whi e emo a. Opis hosoma ound, da k b own wi h pai ed whi e ba s in
he middle ha a e almos ouching each o he ; an e io ly wi h a whi e bo de ;
pos e io ly wi h a whi e pa ch; do sally co e ed wi h scu um; i e la ge sigillae
p esen , a anged in a iangula o ma ion.
Palp (Figs 16C, D, 17): cymbium o al. Embolus elonga ed, ape ing; ip wi h a
sligh lange. Tegula hood small. RTA ilobed; en al lobed inge -shaped; me-
dian lobe iangula , wi h do sal lobe axe-shaped; do sal edge sligh ly olded.
VTA hook-shaped, small, a ached medially on he en al lobe o RTA.
Female. Unknown.
Dis ibu ion. Only known om he ype locali y (Indonesia: Suma a: Jambi)
(Fig. 19).
Zame opina Simon, 1909
Type species. Zame opina calcea a Simon, 1909.
Diagnosis. Zame opina is simila o Nyc imus in habi us, di e ing in he
p esence o a unique den icula e p ocess nex o he embolus on he male palp
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Figu e 16. Nyc imus saksang Dhiya’ulhaq & Benjamin, sp. no . male holo ype (A aThom036N 2013 HF2.1 001). A. Habi-
us, do sal iew; B. Habi us, en al iew; C. Le palp, en al iew; D. Le palp, e ola e al iew. Scale ba s: 1 mm (A, B);
0.2 mm (C, D).
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Figu e 17. Nyc imus saksang Dhiya’ulhaq & Benjamin, sp. no . male holo ype (A aThom036N 2013 HF2.1 001). A. Le
palp, en al iew; B. Le palp, e ola e al iew. Abb e ia ions: DL = do sal lobe; E = embolus; ML = medial lobe; RTA =
e ola e al ibial apophysis; TH = egula hood; VTA = en al ibial apophysis; VL = en al lobe. Scale ba : 0.2 mm.
(DP in Fig. 18B). No o he o ien al homisid genus possesses his s uc u e.
Fo a de ailed diagnosis see Tang e al. (2010).
Species composi ion. Zame opina calcea a Simon, 1909 and Zame opina
wanliae Lin & Li, 2023.
Zame opina calcea a Simon, 1909
Zame opina calcea a Simon, 1909: Simon 1909: 123; Tang e al. (2010): 66, igs
1−18.
Type ma e ial. Holo ype. Vie nam – Cao Bằng P o ince • ♂; “Boa Luc, dans
Hau -Tonkin, à l’Oues de Cao-Bang e p és la on iè e du Yun-Nan e du
Kuang-Si” (p obably Bảo Lạc Dis ic ); 1906 o 1907; de Pelaco leg.; MNHN
22937. Examined.
Diagnosis. See Lin e al. (2023).
Desc ip ion. See Tang e al. (2010).
Dis ibu ion. China (Yunnan), Vie nam (Cao Bằng) (Fig. 19).
Zame opina wanliae Lin & Li, 2023
Fig. 18
Zame opina wanliae Lin & Li, 2023 in Lin e al. (2023): 79, igs 68A, B, 69I.
