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Eu opean Jou nal o Taxonomy 1016: 1–139
h ps://doi.o g/10.5852/ej .2025.1016.3063
eu opeanjou nalo axonomy.eu
ISSN 2118-9773
2025 · Sha aie S. e al.
This wo k is licensed unde a C ea i e Commons A ibu ion License (CC BY 4.0)
Recei ed: 4 Ap il 2024 • Accep ed: 9 June 2025 • Published: 18 Sep embe 2025
Topic edi o : Magalie Cas elin • Sec ion edi o : A naud Hen a d • Desk edi o : K is iaan Hoedemake s
Monog aph
u n:lsid:zoobank.o g:pub:971686D8-3C1F-4F8F-9AA1-FBB8BC97E175
New da a on zoda iid spide s (A achnida, A aneae) in he Middle Eas
and Eas e n Medi e anean
Sepideh SHAFAIE 1 , S ano PEKÁR 2,* , Osman SEYYAR 3
1,2 Depa men o Bo any and Zoology, Facul y o Sciences, Masa yk Uni e si y, Ko lářská 2,
61137 B no, Czech Republic.
3 Depa men o Biology, Facul y o A & Sciences, Niğde Öme Halisdemi Uni e si y,
TR-51240 Niğde, Tu key.
* Co esponding au ho : [email p o ec ed]
1 Email: [email p o ec ed]
3 Email: [email p o ec ed]
Abs ac . Al hough axonomic esea ch on zoda iid spide s has a long his o y, he iden i ica ion o
many species has emained challenging due o insu icien documen a ion. In his s udy, we e isi
50 zoda iid species om he Middle Eas and Eas e n Medi e anean. Fo each axon examined, we
p o ide de ailed pho og aphic documen a ion o soma ic and copula o y s uc u es o acili a e u u e
axonomic in es iga ions. Se en new species om h ee di e en gene a a e desc ibed: Lachesana
ga ishae Sha aie & Peká sp. no . (♂♀, Is ael), L. lubinae Sha aie & Peká sp. no . (♂♀, Is ael),
Pax akilae Sha aie & Peká sp. no . (♂♀, Sy ia), Zoda ion jakubi Sha aie & Peká sp. no . (♂♀,
Oman), Z. le yi Sha aie & Peká sp. no . (♂♀, Is ael and Sy ia), Z. milani Sha aie & Peká sp. no . (♂,
Sy ia), and Z. zons eini Sha aie & Peká sp. no . (♂, Cyp us and Is ael). Re-examina ion o he Z. nicki
ype specimen con i ms i s dis inc ion om Z. ni idum, and we p o ide edesc ip ions o bo h sexes o
Z. ni idum. Addi ionally, he male o Z. nicki and he emale o Z. nam un a e desc ibed he e o he i s
ime. We e alida e Z. une iacum and designa e a neo ype based on newly examined ma e ial. Two new
synonymies a e also p oposed: Acan hinozodium c a e i e um syn. no . is synonymized wi h A. ibes i,
and Z. kun i syn. no . is synonymized wi h A. odem. Fu he mo e, we epo en new coun y eco ds:
A. odem (Sy ia), A. ibes i (Egyp ), Z. aban ense (G eece), Z. g anula um (Sy ia), Z. ka pa hos (Tu key),
Z. mo osum (Sy ia), Z. e icula um (Tu key), Z. hodiense (Cyp us, Sy ia), Z. honi (I an, G eece), and
Z. une iacum (Egyp ).
Keywo ds. An -ea ing spide s, Cen al Asia, No h A ica, new species.
Sha aie S., Peká S. & Seyya O. 2025. New da a on zoda iid spide s (A achnida, A aneae) in he Middle Eas and
Eas e n Medi e anean. Eu opean Jou nal o Taxonomy 1016: 1–139. h ps://doi.o g/10.5852/ej .2025.1016.3063
Eu opean Jou nal o Taxonomy 1016: 1–139 (2025)
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In oduc ion
Zoda iidae Tho ell, 1881, wi h a o al numbe o 1307 species, including ex an (Wo ld Spide Ca alog
2025) and ossil (Dunlop e al. 2023) ep esen a i es, is amongs he mos speciose spide amilies. The
amily has a wide dis ibu ion, wi h he highes species di e si y occu ing in opical and sub opical
egions o he Palea c ic, Aus alian, A ican, and Neo opical ealms. Despi e ema kable p og ess in
e ealing he biodi e si y o zoda iids o e he pas decade (Jocqué & Hen a d 2015; Bosmans e al.
2019; Coşa & Danişman 2021; Coşa e al. 2021, 2022; Lecigne & Hen a d, 2022; Zamani & Ma usik
2021, 2022), a la ge numbe o species emain poo ly documen ed. Fo ins ance, o e 41% o cu en ly
desc ibed species a e known only om a single sex o ju enile specimens. In addi ion, 10 o he 90
ecognized gene a a e mono ypic (Wo ld Spide Ca alog 2025).
The Middle Eas and Eas e n Medi e anean lie a he in e sec ion o i e majo biodi e si y ho spo s
(Mye s e al. 2000), sugges ing a high po en ial o species ichness (K upp e al. 2009). Despi e his,
cu en eco ds documen only 113 species om 10 gene a ac oss ou sub amilies – C yp o helinae
C.L. Koch, 1872; Lachesaninae Jocqué, 1991; S o eninae Simon, 1893; Zoda iinae Simon, 1890 – in
his as egion (Wo ld Spide Ca alog 2025), sugges ing ha he ac ual biodi e si y may be signi ican ly
unde es ima ed. No ably, a conside able numbe o zoda iid species ha e been epo ed om coun ies
such as G eece, Tu key, I an, and Is ael. In con as , many o he coun ies in he egion – including
Lebanon, Egyp and Yemen – epo ewe han nine species, while some – such as Oman and Kuwai
– lack any o mal occu ence eco ds. This dispa i y highligh s signi ican gaps in ou unde s anding
o zoda iid dis ibu ion and emphasizes he need o in ensi ied sampling and axonomic esea ch
h oughou he egion.
While species-le el iden i ica ion in zoda iid spide s adi ionally elies on di e ences in he copula o y
o gan o bo h sexes, compa a i e examina ions and habi us-based illus a ions emain la gely una ailable
o mos desc ibed species. This gap unde sco es a undamen al obs acle in zoda iid axonomy. Fo
ins ance, Zoda ion ex aneum Denis, 1935, o iginally desc ibed om a single emale specimen, was
synonymized by Bosmans (1994) due o he absence o dis inc copula o y o gan di e ences be ween
i s holo ype and specimens o Z. s yli e um (Simon, 1870). La e , Peká e al. (2003) ele a ed i o a
subspecies, Z. s yli e um ex aneum, based on di e ences in ch omosome numbe , ch omosome sys ems
and body colo a ion. In subsequen wo k, Peká e al. (2012) no ed ha , despi e pa ial ep oduc i e
isola ion and he absence o mi ochond ial DNA di e gence be ween Z. s. ex aneum (wi h he ligh es
colo a ion) and Z. s. s yli e um (wi h he da kes colo a ion), he wo popula ions displayed dis inc
ecological p e e ences, including di e ences in mic ohabi a and die . O iz e al. (2021) expanded on
hese indings by conduc ing a la ge-scale phylogeog aphic s udy o he Z. s yli e um complex ac oss
he Ibe ian Peninsula, e ealing wo allopa ic species – one p ima ily in Po ugal and he o he in Spain.
They explained he p e iously obse ed lack o mi ochond ial di e gence by iden i ying bo h ligh e and
da ke mo phs wi hin each species. Mos ecen ly, Sha aie e al. (2025), a e examining a la ge numbe
o specimens, con i med ha he wo species iden i ied by O iz e al. (2021) could be dis inguished by
colo a ion o emo a III and di e ences in male palp, he eby eins a ing Z. ex aneum as a alid species,
p edominan ly dis ibu ed in wes e n Ibe ia, especially in cen al-sou he n Po ugal.
Simila ly, a compa a i e s udy o Z. ni idum Audouin, 1826 by Peká e al. (2022) iden i ied ma ked
ecological and beha io al di e ences be ween he black and yellow mo pho ypes, including a ia ion
in ci cadian ac i i y, p ey p e e ences, and enom composi ion. These dis inc ions we e u he
suppo ed by e idence o pa ial o comple e p e- ep oduc i e isola ion, sugges ing a p obable case o
ecen sympa ic specia ion. Complemen ing hese indings, O iz e al. (2024) conduc ed a genomic
analysis ha s ongly suppo s he in e p e a ion ha he yellow and black mo phs ep esen dis inc
and e olu iona ily independen lineages, wi h no de ec able ecen gene low. No ably, phylogene ic
SHAFAIE S. e al., New da a on Middle Eas and Eas e n Medi e anean zoda iids (A aneae)
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econs uc ions indica e ha he yellow lineage is mo e closely ela ed o Z. luc uosum (O. Picka d-
Camb idge, 1872), a mo phologically dis inc species, han o he black lineage.
These s udies collec i ely suppo he hypo hesis ha addi ional species complexes may exis wi hin
he genus Zoda ion, wa an ing de ailed sc u iny o popula ions ac oss he genus o de ec consis en
mo phological di e ences. Such complexes o en comp ise axa ha exhibi p onounced gene ic
di e gence ye display li le mo phological di e en ia ion. Recognizing ha colo a ion may hold
axonomic alue compa able o o he mo phological ai s – and conside ing ha such ea u es ha e o en
been insu icien ly documen ed – we aim o expand he cu en axonomic da ase by including habi us
pho og aphs whe e such in o ma ion is lacking. Ou objec i e is o p o ide a comp ehensi e axonomic
e ision o zoda iid species om he Middle Eas and Eas e n Medi e anean egions. Gi en he high
species ichness o his amily in he egion, ou indings will be p esen ed ac oss mul iple olumes o
suppo and acili a e u u e axonomic wo k.
Ma e ial and me hods
All desc ip ions we e ob ained om specimens imme sed in e hanol. The ul a was dissec ed and ea ed
o 5–11 min by gen le boiling in 20% KOH. The specimens we e pho og aphed using a KEYENCE
VHX-5000 digi al mic oscope, and he images we e enhanced using Adobe Pho oshop CS2 Ex ended.
All measu emen s a e in millime e s. The leng hs o he palp and leg segmen s a e measu ed on he
do sal side and a e gi en as: o al ( emu , pa ella, ibia, me a a sus [no p esen in palp], and a sus). The
spina ion o legs I–IV is epo ed only o species o Lachesana. The e minology used ollows ha o
Jocqué (1991), Le y (1992), and Bosmans (1994).
Ins i u ional abb e ia ions and cu a o s
CMU = Collec ion o Masa yk Uni e si y, B no, Czech Republic (S. Peká )
HNHM = Hunga ian Na u al His o y Museum, Budapes , Hunga y (W. Wins)
HUJ = Heb ew Uni e si y o Je usalem, Je usalem, Is ael (E. Ga ish-Rege )
MMBC = Mo a ské Zemské Muzeum, B no, Czech Republic (J. Mi áček)
MNHN = Na ional Museum o Na u al His o y, Pa is, F ance (C. Rolland)
MSNB = Museo Ci ico di Scienze Na u ali “E. Ca i“, Be gamo, I aly (P. Pan ini)
NMBE = Na u al His o y Museum in Be n, Be n, Swi ze land (C. K op )
NMP = Na ional Museum in P ague, P ague, Czech Republic (P. Dolejš)
OUMNH = Ox o d Uni e si y Museum o Na u al His o y, Ox o d, Uni ed Kingdom (Z. Simmons)
SMF = Na u al His o y Museum o Senckenbe g, F ank u am Main, Ge many (P. Jäge )
ZMFUM = Zoological Museum o he Fe dowsi Uni e si y o Mashhad, Mashhad, I an (O. Mi shamsi)
ZMH = Zoological Museum o Na u e, Hambu g, Ge many (D. Ha ms)
ZMMU = Zoological Museum o he Moscow S a e Uni e si y, Moscow, Russia (K. G. Mikhailo )
ZMUT = Zoological Museum o he Uni e si y o Tu ku, Tu ku, Finland (V. Vah e a)
Abb e ia ions o mo phological e ms
ALE = an e io la e al eye
AME = an e io median eye
Ap = an e io pocke
B = base o e ola e al ibial apophysis
Co = conduc o
Cp = conduc o al p ong
Cx = coxa
d = do sal
Em = embolus
FD = e iliza ion duc
Eu opean Jou nal o Taxonomy 1016: 1–139 (2025)
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Fe = emu
ju . = ju enile(s)
MA = median apophysis
Ml = lamella ex ension o he median apophysis
Mp = median pla ele
Ms = median scape
M = me a a sus
p = p ola e al
Pa = pa ella
PLE = pos e io la e al eye
PMA = p ola e al p ocess o median apophysis
PME = pos e io median eye
PVS = pos e io en al spines
= e ola e al
RTA = e ola e al ibial apophysis
Sh = spe ma hecal head
Sm = spe ma hecal medial ube
Ss = spe ma hecal s alk
Ta = a sus
Ti = ibia
Ts = ibial spu
= en al
Va = en o- e ola e al apophysis
VTP = en al anspa en p o usion o he palpal ibia
Resul s
Class A achnida Cu ie , 1812
O de A aneae Cle ck, 1757
Family Zoda iidae Tho ell, 1881
Sub amily Lachesaninae Jocqué, 1991
Genus Lachesana S and, 1932
Type species
Lachesis pe e sa Audouin, 1826.
Diagnosis
Fo genus diagnosis, e e o Jocqué (1991).
Lachesana ga ishae Sha aie & Peká sp. no .
u n:lsid:zoobank.o g:ac :E314B50D-8742-4003-BC4B-13F516A87F71
Figs 1–3, 106; Tables 1–2
Diagnosis
The male o L. ga ishae sp. no . esembles hose o L. pe seus Zamani & Ma usik, 2021 in he gene al
shape o he e ola e al ibial apophysis, and L. insensibilis Jocqué, 1991 in he bulb de ails and he
mesop oximal spines on he cymbium. Howe e , he new species can be dis inguished om he o me
species by (1) he p esence o spines on he mesop oximal pa o cymbium ( s absen , c . Fig. 2A, D
SHAFAIE S. e al., New da a on Middle Eas and Eas e n Medi e anean zoda iids (A aneae)
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Fig. 1. Lachesana ga ishae Sha aie & Peká sp. no . A–B, E–F. Holo ype, ♂ (HUJINV-A 21312).
C–D. Pa a ype, ♀ (HUJINV-A 21313). A. Habi us, do sal iew. B. Habi us, en al iew. C. Habi us,
do sal iew. D. Habi us, en al iew. E. Ca apace, en al iew. F. Leg I, la e al iew. Scale ba s: A–E =
2 mm; F = 1 mm.
Eu opean Jou nal o Taxonomy 1016: 1–139 (2025)
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and Zamani & Ma usik 2021: ig. 3h, j); and (2) he uni o mly pale and pa e nless abdomen ( s wi h
dis inc pa e n, c . Fig. 1A and Zamani & Ma usik 2021: ig. 1e). The new species can be dis inguished
om L. insensibilis by ha ing a leas 1.5 imes as many spines on each leg (e.g., leg IV, 53 in L. ga ishae
s 20 in L. insensibilis) and he absence o a scu um (Fig. 1A). In addi ion, he new species migh be
dis inguished om all o he species o Lachesana by he smalles body size (7.8 mm s < 9.8 mm) and
by he e ola e al ibial apophysis wi h a a he sho and poin ed ip (Fig. 2C, G). The emale can be
dis inguished om hose o o he congene s by a hea -shaped epigynal pla e wi h la ge lobes ha a e
s ongly chi inized an e ola e ally (Fig. 3A, C) ( s no hea -shaped), as well as by a unique o m o
spe ma hecae, comp ising a sligh ly di e gen , upwa d-sloping s aigh spe ma hecal s alk ending in a
con e gen , globula spe ma hecal head ( s a cylind ical head in L. u i en is Simon, 1873; exclusi ely
di e gen in L. a abae i Zons ein & O chinniko , 1999; c . Fig. 3B, and Le y 1990: ig. 19 and
Zons ein & O chinniko 1990: ig. 4, espec i ely).
E ymology
The new species is named in hono o ou Is aeli colleague, E a Ga ish-Rege .
Fig. 2. Male palp o Lachesana ga ishae Sha aie & Peká sp. no ., holo ype (HUJINV-A 21312).
A. P ola e al iew. B. Ven al iew. C. Re ola e al iew. D. Do sal iew. E. P ola e al iew. F. Ven al
iew. G. Re ola e al iew. Abb e ia ions: see Ma e ial and me hods. Scale ba s: 0.2 mm.
SHAFAIE S. e al., New da a on Middle Eas and Eas e n Medi e anean zoda iids (A aneae)
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Type ma e ial
Holo ype
ISRAEL • ♂; Nege Dese , Mashabim; 31.007° N, 34.739° E; 27–30 Ma . 2007; S. Peká and
S. Hén iques leg.; sand; HUJINV-A 21312.
Pa a ypes
ISRAEL • 2 ♀♀; same collec ion da a as o holo ype; 20 Ma . 2007; HUJINV-A 21313, HUJINV-A
21314.
O he ma e ial examined
ISRAEL • 1 ♀, 9 ju .; same collec ion da a as o pa a ypes; CMU • 1 ju .; same collec ion da a as o
pa a ypes; 27–30 Ma . 2007; S. Peká and S. Hén iques leg.; CMU • 3 ju .; same collec ion da a as o
pa a ypes.; 4 Ap . 2009; J. K ál leg.; CMU.
Desc ip ion
Male
Habi us (Fig. 1A–B, E–F). Ca apace yellow; chelice ae, mou hpa s, s e num, and spinne e s pale;
labium a ow-shaped and conside ably longe han wide; gna hocoxae 1.5 imes as long as labium.
Chelice ae mesally densely co e ed wi h b own se ae. Legs en i ely di y yellow excep o en ally
pale coxae; legs wi h 214 spines (Table 1). Abdomen di y yellow on bo h sides, wi hou scu um.
Fig. 3. Female copula o y o gan o Lachesana ga ishae Sha aie & Peká sp. no ., pa a ype (HUJINV-
A 21313). A. In ac epigyne, en al iew. B. Vul a, en al iew. C. Epigyne, en al iew. D. Vul a,
en al iew. Abb e ia ions: see Ma e ial and me hods. Scale ba s: 0.2 mm.
Eu opean Jou nal o Taxonomy 1016: 1–139 (2025)
8
Measu eMen s. To al leng h 7.85; ca apace 4.4 long, 2 wide a pa s cephalica, 3.15 wide a pa s ho acica.
Eye sizes and in e dis ances: AME: 0.19, ALE: 0.12, PME: 0.15, PLE: 0.17, AME–AME: 0.04, AME–
ALE: 0.06, ALE–ALE: 0.37, PME–PME: 0.06, PME–PLE: 0.19. Legs: I 13.55 (4.25, 1.56, 3.12, 2.81,
1.81), II 14.68 (3.81, 1.5, 3.06, 4.06, 2.25), III 13.33 (4.06, 0.09, 1.94, 4.87, 2.37), IV 16.47 (4.68, 1.75,
2.93, 5.68, 1.43). Palp: 4.1 (1.6, 0.7, 0.5, 1.3).
PalP (Fig. 2A–G). RTA dis ally oblique, bo h p o/ e ola e al edges pa allel, do sal pa ape ing blade-
like; RTA abou 0.6 imes as long as ibia, wides pa o e 3 imes as wide as s alk. Cymbium wi h spines
on mesop oximal side; bulb 1.5 imes as long as wide. Embolus ilamen ous, embola base la ge, abou
hal he leng h o egulum.
Female (pa a ype HUJINV-A 21313)
Habi us (Fig. 1C–D). Ca apace yellow an e io ly, pale pos e io ly; chelice ae yellowish-o ange co e ed
by black se ae. Mou hpa s, s e num and abdomen simila o hose in male; gna hocoxae iangula
shield-shaped, yellow p oximally and pale dis ally. Ca apace and abdomen wi hou pa e n. Spinne e s
pale p oximally and di y yellow dis ally. Legs en i ely yellow, wi h 97 spines (Table 2).
Measu eMen s. To al leng h 10.95; ca apace 5.4 long, 2.4 wide a pa s cephalica, 3.52 wide a pa s
ho acica. Eye sizes and in e dis ances: AME: 0.15, ALE: 0.12, PME: 0.12, PLE: 0.13, AME–AME: 0.06,
AME–ALE: 0.09, ALE–ALE: 0.46, PME–PME: 0.09, PME–PLE: 0.34. Legs: I 12.8 (4.1, 1.9, 2.9, 2.4,
1.5), II 12.5 (4.7, 1.8, 2.1, 2.5, 1.4), III 11.3 (3.3, 1.8, 1.5, 3, 1.7), IV 14.1 (3.9, 1.8, 2.5, 3.9, 2).
ePigyne (Fig. 3A–D). Epigynal pla e abou 1.3 imes as wide as long, including hea -shaped pla e, bo h
la ge lobes da kly chi inized an e ola e ally; wo di e gen copula o y duc s ex ended la e ally. An e io
pocke s s ongly sinuous, ouching each o he . Spe ma hecae si ua ed along epigynal pla e and con ining
edges o epigynal pocke s, including a he long, di e gen , upwa d-sloping s aigh spe ma hecal s alks
ending in globula and con e gen spe ma hecal heads; spe ma hecal heads sepa a ed by abou hei
diame e .
Size a ia ion
To al body leng h 10.95–13.77, ca apace leng h/wid h 5.4–6.87/2.4–3.5 in emales (N = 3).
Dis ibu ion
The species is cu en ly only known om he ype locali y in Is ael.
Table 1. Male leg spina ion o Lachesana ga ishae Sha aie & Peká sp. no . Abb e ia ions: d = do sal;
Fe = emu ; M = me a a sus; p = p ola e al; Pa = pa ella; = e ola e al; Ti = ibia; = en al.
Fe Pa Ti M To al
Id4 p5 6 1 p1 1 p4 2 10 p2 5 9 50
II d4 p7 5 1 d1 p2 1 p4 3 12 p4 2 11 57
III d3 p6 4 5 d1 p2 1 d2 p3 4 6 d3 p3 2 9 54
IV d4 p6 4 3 d2 p1 1 d2 p3 4 6 d3 p5 2 7 53
SHAFAIE S. e al., New da a on Middle Eas and Eas e n Medi e anean zoda iids (A aneae)
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Lachesana lubinae Sha aie & Peká sp. no .
u n:lsid:zoobank.o g:ac :EF76D45C-001C-4947-B30C-D43AD30FF63C
Figs 4–6, 106; Tables 3–4
Diagnosis
This new species is mos simila o L. ga ishae sp. no . in e ms o body colo a ion and o e all palp
mo phology. The male o he new species can be ecognized by a wide e ola e al ibial apophysis
apical wid h/basal wid h a io (3.8 s 3.1); and di e en numbe o spines on leg segmen s and cymbium
(i.e., me a a sus spines 89 s 57 and cymbium spines 9 s 16). The emale can be dis inguished by a
ema kably spa ula ed, mush oom-shaped epigynal pla e ( s hea -shaped, c . Fig. 6A, C and Fig. 3A,
C), and con e gen spe ma hecal s alks ending in kidney-shaped spe ma hecal heads ( s pa allel
spe ma hecal s alks, wi h globula spe ma hecal heads, c . Fig. 6B, D and Fig. 3B, D). Fu he mo e, he
o al numbe o leg spines can sepa a e bo h adul and ju enile specimens o L. lubinae sp. no . om
hose o L. ga ishae (> 130 spines s < 97).
E ymology
This species is named in hono o ou colleague, he Is aeli a achnologis , Yael Lubin.
Type ma e ial
Holo ype
ISRAEL • ♂; Nege Dese , Mashabim; 31.007° N, 34.739° E; 19 Oc . 2004; J. K ál leg.; sands;
HUJINV-A 21315.
Pa a ype
ISRAEL • 1 ♀; same collec ion da a as o holo ype; 20 Ma . 2007; S. Peká leg.; HUJINV-A 21316.
O he ma e ial examined
ISRAEL • 6 ju .; same collec ion da a as o pa a ype; CMU • 7 ju .; same collec ion da a as o pa a ype;
27–30 Ma . 2007; S. Peká and S. Hén iques leg.; CMU • 2 ju .; same collec ion da a as o pa a ype; 4
Ap . 2009; J. K ál leg.; CMU.
Desc ip ion
Male
Habi us (Fig. 4A–B, E, G). Ca apace da k yellow, wi h se e al pale s ipes adia ing om o ea;
chelice ae, gna hocoxae, and a ow-shaped labium pale, labium as long as wide; chelice ae mesally
densely co e ed wi h hin, b own se ae; gna hocoxae 1.6 imes as long as labium. S e num pale. Fe, Pa,
Table 2. Female leg spina ion o Lachesana ga ishae Sha aie & Peká sp. no . Abb e ia ions: d =
do sal; Fe = emu ; M = me a a sus; p = p ola e al; Pa = pa ella; = e ola e al; Ti = ibia; =
en al.
Fe Pa Ti M To al
Id4 p1 1 0 p2 1 p0 0 4 p0 0 10 23
II d4 p1 0 0 P1 0 p2 0 2 p1 0 10 21
III d4 p1 0 0 p3 2 d0 p3 4 0 d1 p3 2 6 29
IV d3 p0 0 0 p3 1 d0 p3 2 0 d1 p4 2 5 24
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Measu eMen s. To al leng h 7.56; ca apace 3.95 long, 1.01 wide a pa s cephalica, 2.65 wide a pa s
ho acica, scu um leng h 2.59. Eye sizes and in e dis ances: AME 0.33, ALE 0.39, PME 0.34, PLE 0.39,
AME–AME 0.17, AME–ALE 0.28, ALE–ALE 1.29, PME–PME 0.22, PME–PLE 0.73. Legs: I 8.44
(2.37, 1.12, 1.8, 1.74, 1.41), II 7.99 (2.59, 1.12, 1.52, 1.58, 1.18), III 7.98 (2.3, 1.12, 1.41, 2.03, 1.12),
IV 10.29 (2.7, 1.18, 2.24, 2.59, 1.58). Palp: 4.11 (1.58, 0.34, 0.28, 1.91).
PalP (Fig. 8A–F). RTA wi h an oblong ex ension, comp ising a a he long, o wa d poin ed ibial spu
and a obus hump. Median pla ele ex ended mesally and an e io ly; conduc o al p ong inge -shaped;
median apophysis ec angula basally and na owed apically, wi h a olded ip. Embolus e y long,
o igina ing a a 4 o’clock posi ion, na owed sligh ly and olded a ound bulb.
Female (pa a ype NMP P6A 7515)
Habi us (Fig. 7C–D). Ca apace, chelice ae, s e num, mou hpa s simila o hose in male. Legs yellow
excep o sligh ly da ke Cx, g eyish Fe and ligh g ey ips o M ; spinne e s pale, palp segmen s g eyish,
wi h da ke Ta. Abdomen blackish-b own do sally, wi h no pa e n, da k b own en ally, wi h h ee sub-
pa allel yellow s ipes in he middle, la e al s ipes wide han middle s ipe.
Measu eMen s. To al leng h 8.29; ca apace 3.5 long, 1.24 wide a pa s cephalica, 2.37 wide a pa s
ho acica. Eye sizes and in e dis ances: AME 0.33, ALE 0.39, PME 0.45, PLE 0.39, AME–AME 0.11,
AME–ALE 0.33, ALE–ALE 0.99, PME–PME 0.17, PME–PLE 0.56. Legs: I 8.44 (2.37, 1.12, 1.8, 1.74,
1.41), II 7.99 (2.59, 1.12, 1.52, 1.58, 1.18), III 7.71 (2.03, 1.12, 1.41, 2.03, 1.12), IV 10.29 (2.7, 1.18,
2.24, 2.59, 1.58). Palp: 3.03 (1.01, 0.45, 0.45, 1.12).
Fig. 9. Female copula o y o gan o Pax akilae Sha aie & Peká sp. no ., pa a ype (NMP P6A 7515).
A. In ac epigyne, en al iew. B. Epigyne, en al iew. C. Vul a, en al iew. D. Vul a, en al iew.
Abb e ia ion: see Ma e ial and me hods. Scale ba s: 0.4 mm.
SHAFAIE S. e al., New da a on Middle Eas and Eas e n Medi e anean zoda iids (A aneae)
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ePigyne (Fig. 9A–D). Epigyne abou 1.6 imes as wide as long, comp ising h ee longi udinal ubes: one
da k medial and wo ligh , sub-la e al ubes accompanied by a pai o da k ma ginal ubes. Spe ma hecal
ubes pa allel comp ising a pai o elonga ed, yellow s uc u es, si ua ed medially, ex e nally a ached o
a pai o eddish- b own, la ge s uc u es ending in wis ed coils.
Size a ia ion
To al body leng h 6.5–7.62, ca apace leng h/wid h 3.5–3.95/2.01–2.75 in males (N = 3). To al body
leng h 7.12–8.29, ca apace leng h/wid h 3–3.5/2.24–2.3 in emales (N = 5).
Dis ibu ion
The species is cu en ly only known om he ype locali y in Sy ia.
Pax engediensis Le y, 1990
Fig. 10
Pax engediensis Le y, 1990: 356, igs 64–67.
Diagnosis
Re e o Le y (1990).
Type ma e ial examined
Holo ype
ISRAEL • ♀; En Gedi; 26 Ma . 1957; A. Shulo leg.; HUJ 14216.
Desc ip ion
Fo a de ailed desc ip ion o emale, e e o Le y (1990).
Dis ibu ion
Is ael.
Pax islami a (Simon, 1873)
Figs 11–12
Hab ones es islami a Simon, 1873: 75.
S o ena islami a – Simon 1893: 247.
Pax islami a – Le y 1990: 358, igs 68–71. — Seyya e al. 2008: 52, igs 1–11.
Diagnosis
Re e o Le y (1990).
Ma e ial examined
ISRAEL • 3 ♀♀, 9 ju .; Adullam; 31.641° N, 34.931° E; 25 Ma . 2001; S. Peká , S. Hén iques and
M. Řezáč leg.; CMU • 2 ♂♂, 6 ♀♀, 2 ju .; same collec ion da a as o p eceding; 5 Ap . 2017; S. Peká
and S. Ko enko leg.; CMU • 1 ♂; Moun Ca mel; 13 Ap . 2009; J. K ál leg.; CMU • 1 ♂; Ca mel;
32.733° N, 35.05° E; 6 May 2023; M. Řezáč leg.; CMU.
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TURKEY • 1 ♂; SE Tu key; 28 Ap . 2004; M. Řezáč leg.; CMU • 1 ♂, 1 ♀; Top akkale; 37.061° N,
36.148° E; 67 m a.s.l.; 1 May 2009; CMU • 1 ♂; Me sin, 12 km o Anamu , 1 km o Ala B idge;
36.174° N, 32.894° E; 93 m a.s.l.; 14 May 2009; R. Kaya leg.; ZMUT.
Desc ip ion
Fo a de ailed desc ip ion, e e o Le y (1990).
Dis ibu ion
Tu key, Is ael, Sy ia, Lebanon.
Fig. 10. Pax engediensis Le y, 1990, holo ype, ♀ (HUJ 14216). A. Habi us, do sal iew. B. Habi us,
en al iew. C. In ac epigyne, en al iew. Scale ba s: A–B = 1 mm; C = 0.4 mm.
SHAFAIE S. e al., New da a on Middle Eas and Eas e n Medi e anean zoda iids (A aneae)
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Fig. 11. Habi us o Pax islami a (Simon, 1873) (CMU). A–B. ♂. C–D. ♀. A. Do sal iew. B. Ven al
iew. C. Do sal iew. D. Ven al iew. Scale ba s: 1 mm.
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20
Fig. 12. Copula o y o gans o Pax islami a (Simon, 1873) (CMU). A–G. Male palp. H–I. Female
copula o y o gan. A. P ola e al iew. B. Ven al iew. C. Ven o- e ola e al iew. D. Re ola e al iew.
E. P ola e al iew. F. Ven al iew. G. Re ola e al iew. H. In ac epigyne, en al iew. I. Vul a, en al
iew. Scale ba s: 0.4 mm.
SHAFAIE S. e al., New da a on Middle Eas and Eas e n Medi e anean zoda iids (A aneae)
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Pax palmonii Le y, 1990
Figs 13–14
Pax palmonii Le y, 1990: 352, igs 60–63.
Diagnosis
Re e o Le y (1990).
Type ma e ial examined
Holo ype
ISRAEL • ♂; Ma’oz Hayyam; 30 May 1943; A. Shulo leg.; HUJ 14215.
Pa a ype
ISRAEL • 1 ♀; same collec ion da a as o holo ype; HUJ 14215.
Desc ip ion
Fo a de ailed desc ip ion, e e o Le y (1990).
Dis ibu ion
Is ael.
Sub amily Zoda iinae Simon, 1890
Genus Acan hinozodium Denis, 1966
Type species
Acan hinozodium spinulosum Denis, 1966.
Diagnosis
Fo genus diagnosis, e e o Jocqué & Hen a d (2015).
Acan hinozodium odem (Le y, 2007)
Figs 15–16, 105
Zoda ion odem Le y, 2007: 15, igs 38–39.
Zoda ion kun i Coşa e al., 2021: 49, igs 5a– , 6a–d, 7a–c, 8a–c, 9a–b, 10a–b. Syn. no .
Acan hinozodium odem – Zons ein 2025: 392, igs 1–21.
Diagnosis
Re e o Zons ein (2025).
Type ma e ial examined
Holo ype
ISRAEL • ♀; Odem Fo es , Golan Heigh s (219/290 Is. G .); 950 m a.s.l.; May 1996; S. Rake e leg.;
HUJ 15220.
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Fig. 13. Habi us o Pax palmonii Le y, 1990. A–B. Holo ype, ♂ (HUJ 14215). C–D. Pa a ype, ♀ (HUJ
14215). A. Do sal iew. B. Ven al iew. C. Do sal iew. D. Ven al iew. Scale ba s: 1 mm.
SHAFAIE S. e al., New da a on Middle Eas and Eas e n Medi e anean zoda iids (A aneae)
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Fig. 14. Copula o y o gans o Pax palmonii Le y, 1990. A–D. Holo ype, male palp (HUJ 14215).
E. Pa a ype, emale copula o y o gan (HUJ 14215). A. Palp, p ola e al iew. B. Palp, en al iew.
C. Palp, en o- e ola e al iew. D. Palp, e ola e al iew. E. In ac epigyne, en al iew. Scale ba s:
0.2 mm.
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Fig. 15. Habi us o Acan hinozodium odem (Le y, 2007). A–B. ♂ (CMU). C–D. ♀ (CMU). A. Do sal
iew. B. Ven al iew. C. Do sal iew. D. Ven al iew. Scale ba s: 1 mm.
Fig. 16 (nex page). Copula o y o gans o Acan hinozodium odem (Le y, 2007). A–G. Male palp (CMU).
H–J. Female copula o y o gan (CMU), specimens om Sy ia (H) and Is ael (I–J). A. P ola e al iew.
B. Ven al iew. C. Re ola e al iew. D. P ola e al iew. E. Ven al iew. F. Re ola e al iew, a ow
indica es ci cula pi . G. Embola ip, en al iew. H. In ac epigyne, en al iew. I. In ac epigyne,
en al iew. J. Vul a, en al iew. Abb e ia ion: see Ma e ial and me hods. Scale ba s: 0.2 mm.
SHAFAIE S. e al., New da a on Middle Eas and Eas e n Medi e anean zoda iids (A aneae)
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Fig. 21. Habi us o Palaes ina expoli a O. Picka d-Camb idge, 1872 (CMU). A–B. ♂. C–D. ♀. A. Do sal
iew. B. Ven al iew. C. Do sal iew. D. Ven al iew. Abb e ia ion: see Ma e ial and me hods. Scale
ba s: 0.2 mm.
SHAFAIE S. e al., New da a on Middle Eas and Eas e n Medi e anean zoda iids (A aneae)
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Palaes ina e emica Le y, 1992
Figs 23–24
Palaes ina e emica Le y, 1992: 74, igs 19–26.
Diagnosis
Re e o Le y (1992).
Type ma e ial examined
Holo ype
EGYPT • ♂; Sinai, S Ca he ine Monas e y, (049/775 Is .g id); 14 Ap . 1987; V.D. Ro h leg.; HUJ 14250.
Fig. 22. Copula o y o gans o Palaes ina expoli a O. Picka d-Camb idge, 1872 (CMU). A–C. Male palp.
D–E. Female copula o y o gan. A. P ola e al iew. B. Palp wi h b oken median apophysis, en al iew.
C. Re ola e al iew. D. In ac epigyne, en al iew. E. Vul a, en al iew. Abb e ia ions: see Ma e ial
and me hods. Scale ba s: 0.2 mm.
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Fig. 23. Habi us o Palaes ina e emica Le y, 1992. A–B. Holo ype, ♂ (HUJ 14250). C–D. Pa a ype, ♀
(HUJ 14251). A. Do sal iew. B. Ven al iew. C. Do sal iew. D. Ven al iew. Scale ba s: 0.2 mm.
SHAFAIE S. e al., New da a on Middle Eas and Eas e n Medi e anean zoda iids (A aneae)
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Pa a ype
EGYPT • 1 ♀; same collec ion da a as o holo ype; HUJ 14251.
Desc ip ion
Fo a de ailed desc ip ion, e e o Le y (1992).
Dis ibu ion
Egyp , Is ael.
Fig. 24. Copula o y o gans o Palaes ina e emica Le y, 1992. A–C. Holo ype, male palp (HUJ 14250).
D. Pa a ype, emale (HUJ 14251). A. Palp, p ola e al iew. B. Palp, en al iew. C. Palp, e ola e al
iew. D. In ac epigyne, en al iew. Scale ba s: 0.2 mm.
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36
Genus Pa azoda ion O chinniko , Ahmad & Gu ko, 2009
Type species
Zoda ion addei Simon, 1889.
Diagnosis
Fo genus diagnosis, e e o O chinniko e al. (2009).
Pa azoda ion addei (Simon, 1889)
Figs 25–26, 106
Zoda ion addei Simon, 1889: 383.
Zoda ion laso i Sy she skaja in Vlasso & Sy she skaja, 1937: 249, igs 1–3.
Zoda ion addei – Denis 1958: 111, igs 42–44. — Fe 1985: 274. — Ma usik & Koponen 2001: 46,
ig. 27.
Pa azoda ion addei – O chinniko e al. 2009: 471, ig. 1.1–6. — Zamani & Ma usik 2021: 182,
ig. 26a–c.
Diagnosis
Re e o O chinniko e al. (2009).
Ma e ial examined
AFGHANISTAN • 1 ♀; Badghis, Bala Mu ghab; 18 Jul.–4 Sep. 1964; O. Jakeš leg.; MMBC 7633 (5601/
En ).
IRAN • 5 ♀♀; Taybad, Dowqa un; 8 No . 2014; P. Rashidi leg.; ZMFUM-TA 1183.
Desc ip ion
Fo a de ailed desc ip ion, e e o O chinniko e al. (2009).
Rema ks
The single emale specimen om A ghanis an exhibi s sligh di e ences compa ed o he I anian
popula ion, pa icula ly in he heigh o he spe ma hecal bodies, which a e a anged in descending
o de ( s mo e o less equally sized, c . Fig. 26D and Fig. 26B), and in he ligh e p oximal pa s o Fe I
and II ( s en i ely da k, c . Fig. 25C and Fig. 25A). Gi en he unlikely p ospec o ob aining addi ional
ma e ial om A ghanis an in he nea u u e, we ea ed his single specimen as P. addei.
Dis ibu ion
A ghanis an, I an, Kazakhs an, Uzbekis an, Tu kmenis an, Tajikis an, Uni ed A ab Emi a es.
SHAFAIE S. e al., New da a on Middle Eas and Eas e n Medi e anean zoda iids (A aneae)
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Fig. 25. Habi us o Pa azoda ion addei (Simon, 1889), ♀. A–B. Specimen om I an (ZMFUM-TA
1183). C–D. Specimen om A ghanis an (MMBC 7633 (5601/En ). A. Do sal iew. B. Ven al iew.
C. Do sal iew. D. Ven al iew. Scale ba s: 1 mm.
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Genus T yge us Simon, 1882
Type species
T yge us sexocula us O. Picka d-Camb idge, 1872.
Diagnosis
Fo genus diagnosis, e e o Le y (1992).
T yge us sexocula us (O. Picka d-Camb idge, 1872)
Figs 27–28
Palaes ina sexocula a O. Picka d-Camb idge, 1872: 270.
T yge us senocula us – Simon 1893: 439.
T yge us sexocula us – Jocqué 1991: 149, igs 378–383. — Le y 1992: 77, igs 27–38.
Diagnosis
Re e o Jocqué (1991) and Le y (1992).
Fig. 26. Female copula o y o gans o Pa azoda ion addei (Simon, 1889). A–B. Specimen om I an
(ZMFUM-TA 1183). C–D. Specimen om A ghanis an (MMBC 7633 (5601/En ). A. Epigyne, en al
iew. B. Vul a, en al iew. C. In ac epigyne, en al iew. D. Epigyne, en al iew. Scale ba s:
0.2 mm.
SHAFAIE S. e al., New da a on Middle Eas and Eas e n Medi e anean zoda iids (A aneae)
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Ma e ial examined
ISRAEL • 2 ♂♂; Saye Shaked; 1999; J. Le y leg.; CMU • 1 ♀; Leha im; 31.364° N, 34.823° E; 12
Ap . 2001; S. Peká leg.; CMU • 2 ♀♀; same collec ion da a as o p eceding; 20 Ap . 2001; S. Peká
leg.; CMU • 1 ♂; Halukim; 30.854° N, 34.768° E; 21 Ma . 2007; S. Peká leg.; CMU • 8 ♂♂, 3 ♀♀, 1
ju .; same collec ion da a as o p eceding; 7 Ap . 2019; S. Peká leg.; CMU • 2 ♂♂; same collec ion da a
as o p eceding; 29 Ma . 2023; S. Peká leg.; CMU • 1 ♀; Wadi Pa an; 8 Ap . 2019; S. Peká leg.; CMU.
Desc ip ion
Fo a de ailed desc ip ion, e e o Le y (1992).
Dis ibu ion
Egyp , Is ael.
Fig. 27. Habi us o T yge us sexocula us (O. Picka d-Camb idge, 1872) (CMU). A–B. ♂. C–D. ♀.
A. Do sal iew. B. Ven al iew. C. Do sal iew. D. Ven al iew. Scale ba s: 1 mm.
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Genus Zoda ion Walckenae , 1826
Type species
Enyo ni ida Audouin, 1826.
Diagnosis
Fo genus diagnosis, e e o Le y (1992).
Fig. 28. Copula o y o gans o T yge us sexocula us (O. Picka d-Camb idge, 1872) (CMU). A–C. Male
palp. D–E. Female copula o y o gan. A. P ola e al iew. B. Ven al iew. C. Re ola e al iew. D. In ac
epigyne, en al iew. E. Vul a, en al iew. Scale ba s: 0.2 mm.
SHAFAIE S. e al., New da a on Middle Eas and Eas e n Medi e anean zoda iids (A aneae)
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Zoda ion ni idum Audouin, 1826
Figs 29–31, 87, 106
Enyo ni ida Audouin, 1826: 350, pl. 3 ig. 7.
Enyo longipes Audouin, 1826: 351, pl. 3 ig. 8.
Zoda ion camb idgei Denis, 1938: 387.
Clo ho ni ida – Walckenae 1837: 639.
Clo ho longipes – Walckenae 1837: 640.
A gus longipes – Walckenae 1841: 347, pl. 16 ig. 6.
Zoda ion longipes – Walckenae 1847: 563.
Enyo ni ida – Walckenae 1847: 563. — Tho ell 1869: 108. — F.O. Picka d-Camb idge 1876: 559.
Enyo luisan e – Simon 1864: 160.
Zoda ium longipes – Tho ell 1869: 107.
Zoda ium ni idum – Simon 1885: 29.
Zoda ion ni idum – Simon 1893: 432, igs 403–407; 1908: 425. — Denis 1937: 6, pl. 1 ig. 1; 1962:
31, ig. 2. — Le y 1992: 86, igs 53–61. — Zamani & Ma usik 2021: 188, ig. 28a–c; 2022: 162,
ig. 11a–c.
Me a gus longipes – F. Picka d-Camb idge 1902: 8.
Diagnosis
Zoda ion ni idum specimens a e mos simila o Z. nicki in he shape o hei copula o y o gans, bu he
o me species is dis inguished by a b ownish ca apace and yellow coxae ( s black ca apace and black
o g ey coxae; c . Fig. 29A, C and Fig. 84A, C). Fu he mo e, males o Z. ni idum possess uni o mly
yellow emo a ( s en i ely black emo a in Z. nicki; c . Fig. 29A and Fig. 84A).
Ma e ial examined
EGYPT • 1 ♀; Alexand ia; MNHN-AR2854 • 2 ♀♀; Cai o o Aswan, Lake Ma iou ; MNHN-AR2847.
ISRAEL • 1 ♂; Mid eshe Sde Boke ; 30.857° N, 34.781° E; 1 Ap . 2001; S. Peká leg.; HUJINV-A
21301 • 1 ♀, same collec ion da a as o p eceding; HUJINV-A 21302 • 1 ju .; Mid eshe Sde Boke ;
10 Ma . 2001; S. Peká leg.; CMU • 1 ♂; A a a A ea, 12 km N o Ein Yaha Mosha , Ha ze a Mosha ;
30.754° N, 35.269° E; 13 Ma . 2001; S. Peká leg.; CMU • 1 ♀, 3 ju .; s. loc.; Ap . 2001; S. Peká leg.;
CMU • 12 ♂♂, 9 ♀♀, 50 ju .; same collec ion da a as o p eceding; Feb.–Ma . 2018; L. Sen enská leg.;
CMU • 1 ♀, 2 ju .; same collec ion da a as o p eceding; 6 Ap . 2019; S. Peká leg.; CMU • 9 ♂♂ 3 ♀♀
1 ju .; Halukim; 30.854° N, 34.784° E; 16 Ap . 2001; S. Peká leg.; CMU • 1 ♀; same collec ion da a as
o p eceding; 27 Ma . 2007; S. Peká leg.; CMU • 3 ♂♂, 1 ju .; same collec ion da a as o p eceding;
18 Ma . 2017; O. Macháč leg.; CMU • 6 ♂♂; same collec ion da a as o p eceding; 9 Ap . 2019; S.
Peká leg.; CMU • 1 ♀; Mashabim; 31.000° N, 34.757° E; 4 Sep. 2002; Y. Lubin leg.; CMU • 2 ♀♀;
same collec ion da a as o p eceding; Sep. 2004; J. K ál leg.; CMU • 1 ♂, 1 ♀; same collec ion da a as
o p eceding; 1 Ap . 2016; S. Ko enko leg.; CMU • 1 ♂; same collec ion da a as o p eceding; 1 Ap .
2017; S. Ko enko leg.; CMU • 1 ju .; same collec ion da a as o p eceding; 6 Ap . 2019; S. Peká leg.;
CMU • 1 ♀; same collec ion da a as o p eceding; 28 Ma . 2023; S. Peká and V. Opa o á leg.; CMU •
1 ♀; same collec ion da a as o p eceding; 29 Ma . 2023; S. Peká leg.; CMU • 1 ♂; Bee sheba, be ween
Bee She a and Mi zpe Ramon; 6–29 May 2003; M. Řezáč leg.; CMU • 1 ♀; Mei a ; 29 Sep. 2007;
J. K ál leg.; CMU • 2 ♂♂, 2 ju .; Ye oham; 30.971° N, 34.97° E; 3 Ma . 2016; S. Peká , S. Ko enko
and E. Lízna o á leg.; CMU • 18 ♂♂, 11 ♀♀; same collec ion da a as o p eceding; 5 Ap . 2017; S.
Peká and S. Ko enko leg.; CMU • 2 ♀♀, 8 ju .; same collec ion da a as o p eceding; 18 May 2017;
O. Macháč leg.; CMU • 3 ju .; same collec ion da a as o p eceding; 5 Ap . 2019; S. Peká leg.; CMU •
1 ju .; Ramon C a e , Nahal Pa an; 30.333° N, 34.968° E; 8 Ap . 2019; S. Peká leg.; CMU • 6 ♀♀,
2 ju .; s. loc.; May 2019; S. Peká , L. Sen enská and S. Ko enko leg.; CMU • 2 ju .; Ha i a; 30.872° N,
34.816° E; 28 Ma . 2023; S. Peká and V. Opa o á leg.; CMU • 1 ♂, 1 ♀, 4 ju .; Bo o Lo z; 30.511° N,
34.612° E; 29 Ma . 2023; S. Peká and V. Opa o á leg.; CMU.
Eu opean Jou nal o Taxonomy 1016: 1–139 (2025)
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Fig. 33. Zoda ion aculea um Chyze , 1897, holo ype, ♀ (HNHM 1187). A. P osoma, do sal iew.
B. P osoma, en al iew. C. Abdomen, do sal iew. D. Abdomen, en al iew. E. Epigyne, en al
iew. Scale ba s: A–D = 1 mm; E = 0.2 mm.
SHAFAIE S. e al., New da a on Middle Eas and Eas e n Medi e anean zoda iids (A aneae)
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Zoda ion ba ba ae Bosmans, 2009
Figs 34–35
Zoda ion ba ba ae Bosmans, 2009: 248, igs 80–83, 138–139.
Zoda ion ba ba ae – Bosmans e al. 2019: 32, ig. 20a–g. — Özkü ük 2022: 241, igs 1–15. — Coşa &
Danışman 2024: 268, igs 21–22.
Fig. 34. Habi us o Zoda ion ba ba ae Bosmans, 2009. A–B. Holo ype, ♂ (NMBE-AR4147).
C–D. Pa a ype, ♀ (NMBE-AR4147). A. Do sal iew. B. Ven al iew. C. Do sal iew. D. Ven al iew.
Scale ba s: 1 mm.
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50
Diagnosis
Re e o Bosmans (2009).
Type ma e ial examined
Holo ype
GREECE • ♂; Peloponnisos, O os Taige os, Moni Panaghi Gia issa; 1100 m a.s.l.; 30 Sep. 1991;
B. Kno lach and K. Thale leg.; NMBE-AR4147.
Pa a ypes
GREECE • 2 ♀♀; same collec ion da a as o holo ype; NMBE-AR4147.
Fig. 35. Copula o y o gans o Zoda ion ba ba ae Bosmans, 2009. A–C. Holo ype, male palp (NMBE-
AR4147). D–E. Pa a ype, emale copula o y o gan (NMBE-AR4147). A. P ola e al iew. B. Ven al
iew. C. Re ola e al iew. D. In ac epigyne, en al iew. E. Vul a, en al iew. Scale ba s: 0.2 mm.
SHAFAIE S. e al., New da a on Middle Eas and Eas e n Medi e anean zoda iids (A aneae)
51
O he ma e ial examined
CYPRUS • 1 ♂, 14 ju .; Nicosia, Le ka Dam; 35.063° N, 32.837° E; 288 m a.s.l.; 25 Ap . 2022; S. Peká ,
S. Sha aie, V. Šol ýso á and O. Michálek leg.; CMU.
TURKEY • 1 ♂, 1 ♀ 14 ju .; Konakli, Macchie Hill, 10 km W o Alanya; 20 May 2005; J. Dolanský
leg.; CMU.
Desc ip ion
Fo a de ailed desc ip ion, e e o Bosmans (2009).
Dis ibu ion
G eece, Cyp us, Tu key.
Zoda ion be oni Komneno & Cha zaki, 2016
Figs 36–37
Zoda ion be oni Komneno & Cha zaki in Komneno e al., 2016: 41, igs 97–111.
Diagnosis
Re e o Komneno e al. (2016).
Ma e ial examined
GREECE • 1 ♂; 20 km a e Dadia; 660 m a.s.l.; 22 Jun.–21 Jul. 2015; M. Cha zaki leg.; CMU • 6 ♂♂,
1 ♀; 2 km a e Pe ka/Re ziki Subu b; 121 m a.s.l.; 22 Jun.–21 Jul. 2015; M. Cha zaki leg.; CMU • 2 ♂♂;
be ween Aisymi-Mega De eio, 500 m W.; 389 m a.s.l.; 22 Jul.–26 Aug. 2015; M. Cha zaki leg.; CMU.
Desc ip ion
Fo a de ailed desc ip ion, e e o Komneno e al. (2016).
Dis ibu ion
G eece.
Zoda ion blagoe i Bosmans, 2009
Fig. 38
Zoda ion blagoe i Bosmans, 2009: 249, igs 84–87, 140–141.
Zoda ion blagoe i – Del she e al. 2022: 248, igs 2, 6, 10, 14, 20–22, 30, 34, 38, 42, 46, 49, 52, 55.
Diagnosis
Re e o Bosmans (2009).
Ma e ial examined
GREECE • 10 ♂♂; Aladjagiola We land (N o Ch isoupoli); 41.01° N, 24.703° E; 25 Jun. 2006;
M. Sch öde leg.; CMU.
Desc ip ion
Fo a de ailed desc ip ion, e e o Bosmans (2009).
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52
Fig. 36. Habi us o Zoda ion be oni Komneno & Cha zaki, 2016 (CMU). A–B. ♂. C–D. ♀. A. Do sal
iew. B. Ven al iew. C. Do sal iew. D. Ven al iew. Scale ba s: 0.4 mm.
SHAFAIE S. e al., New da a on Middle Eas and Eas e n Medi e anean zoda iids (A aneae)
53
Dis ibu ion
Bulga ia, G eece.
Zoda ion ch is ae Bosmans, 2009
Figs 39–40
Zoda ion ch is ae Bosmans, 2009: 284, igs 186–187, 196–197.
Zoda ion ch is ae – Coşa e al. 2021: 46, igs 1a– , 2a–e, 3a–b.
Diagnosis
Re e o Bosmans (2009).
Fig. 37. Copula o y o gans o Zoda ion be oni Komneno & Cha zaki, 2016 (CMU). A–C, F. Male
palp, wi h o a ed bulb inside. D–E. Female copula o y o gan. A. P ola e al iew. B. Ven al iew.
C. Re ola e al iew. D. In ac epigyne, en al iew. E. Vul a, en al iew. F. De ail o embolus,
en al iew. Scale ba s: 0.2 mm.
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54
Fig. 38. Zoda ion blagoe i Bosmans, 2009 (CMU), ♂. A. Habi us, do sal iew. B. Habi us, en al iew.
C. Palp, p ola e al iew. D. Palp, en al iew. E. Palp, e ola e al iew. Scale ba s: A–B = 1 mm;
C–E = 0.2 mm.
SHAFAIE S. e al., New da a on Middle Eas and Eas e n Medi e anean zoda iids (A aneae)
55
Fig. 39. Habi us o Zoda ion ch is ae Bosmans, 2009 (CMU). A–B. ♂. C–D. ♀. A. Do sal iew.
B. Ven al iew. C. Do sal iew. D. Ven al iew. Scale ba s: 1 mm.
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56
Ma e ial examined
TURKEY • 1 ♂, 1 ♀; Ispa a, Aksu, Dedegöl Moun ain; 37.656° N, 31.291° E; 8 Jul. 2003; M. Řezáč
leg.; CMU.
Desc ip ion
Fo a de ailed desc ip ion, e e o Bosmans (2009).
Rema ks
Due o damage sus ained by he median apophyses in he male palps and he epigyne du ing he clea ing
p ocess, i was no possible o include pho og aphs o hese s uc u es in his s udy.
Dis ibu ion
G eece, Tu key.
Fig. 40. Copula o y o gans o Zoda ion ch is ae Bosmans, 2009 (CMU). A–C. Male palp. D–E. Female
copula o y o gan. A. P ola e al iew. B. Ven al iew. C. Re ola e al iew. D. Epigyne, en al iew.
E. Vul a, en al iew. Scale ba s: 0.2 mm.
SHAFAIE S. e al., New da a on Middle Eas and Eas e n Medi e anean zoda iids (A aneae)
57
Zoda ion c ewsae Coşa , Danışman & Kun , 2022
Figs 41–42, 106
Zoda ion c ewsae Coşa e al. 2022: 229, igs 1–16.
Diagnosis
Re e o Coşa e al. (2022).
Ma e ial examined
TURKEY • 1 ♂; Tunceli, Munzu Ri e ; 38.957° N, 39.561° E; 22 Jun. 2002; M. Řezáč leg.; CMU.
Desc ip ion
Fo a de ailed desc ip ion, e e o Coşa e al. (2022).
Rema ks
A igu e-based compa ison o he single specimen om eas e n Tu key wi h specimen om he sou he n
popula ion e ealed ha he eas e n specimen exibi ed a signi ican ly sho e dis al b anch o he median
apophysis, and a ligh e emu I and ca apace. Howe e , o accu a ely classi y his popula ion, addi ional
specimens a e needed o e alua e whe he i ep esen s a new species.
Dis ibu ion
Tu key.
Fig. 41. Habi us o Zoda ion c ewsae Coşa , Danışman & Kun , 2022 (CMU), ♂. A. Do sal iew.
B. Ven al iew. Scale ba s: 1 mm.
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64
Zoda ion ena um Simon, 1885
Figs 48–49
Zoda ion ena um Simon, 1885: 336.
Zoda ion ena um – Fage 1921: 174, ig. 2. — Denis 1937: 39, pl. 8 igs 65–68. — Wunde lich 1980:
240. — Bosmans 1997: 271, igs 12–13, 85–86; 2009: 286, igs 192–193, 198–199. — Dimi o
2024: 114, igs 10–14.
Zoda ion c e icum – Roewe 1928: 119, pl. 2 ig. 37.
Zoda ion hodiense – Capo iacco 1948: 47, ig. 6.
Fig. 48. Habi us o Zoda ion ena um Simon, 1885 (CMU). A–B. ♂. C–D. ♀. A. Do sal iew. B. Ven al
iew. C. Do sal iew. D. Ven al iew. Scale ba s: 1 mm.
SHAFAIE S. e al., New da a on Middle Eas and Eas e n Medi e anean zoda iids (A aneae)
65
Zoda ion hodiense nig i emu – Capo iacco 1948: 49.
Zoda ion ionicum – B ignoli 1984: 314, igs 41–42.
Diagnosis
Re e o Bosmans (1997).
Type ma e ial examined
Pa a ype
GREECE • 1 ju .; Naxos; E. Keyseling leg.; MNHN-AR2850.
Fig. 49. Copula o y o gans o Zoda ion ena um Simon, 1885 (CMU). A–C. Male palp. D–E. Female
copula o y o gan. A. P ola e al iew. B. Ven al iew. C. Re ola e al iew. D. In ac epigyne, en al
iew. E. Vul a, en al iew. Scale ba s: 0.2 mm.
Eu opean Jou nal o Taxonomy 1016: 1–139 (2025)
66
O he ma e ial examined
GREECE • 1 ♀, 1 ju .; Volo (S .); MNHN-AR2851 • 1 ♀; 1.5 km a e Dadia; 660 m a.s.l.; 14–22 May
2015; M. Cha zaki leg.; CMU • 3 ♂♂, 2 ju .; same collec ion da a as o p eceding; 22 Jun.–21 Jul. 2015;
M. Cha zaki leg.; CMU • 2 ♂♂, 1 ♀, 1 ju .; 2 km a e Pe ka/Re ziki Subu b; 121 m a.s.l.; 16 May–22
Jun. 2015; M. Cha zaki leg.; CMU • 1 ♀; same collec ion da a as o p eceding; 22 Jul.–24 Aug. 2015;
M. Cha zaki leg.; CMU • 1 ♂; 1.5 km N o Polia; 50 m a.s.l.; 22 Jul.–26 Aug. 2015; M. Cha zaki leg.;
CMU.
Desc ip ion
Fo a de ailed desc ip ion, e e o Bosmans (1997).
Rema ks
E en hough Bosmans (1997) did no examine he ype o Z. ena um, he epo ed ha he holo ype ube
(MNHN-AR2850) con ained one male and one ju enile. Howe e , ou examina ion o he ube e ealed
only a single ju enile specimen, sugges ing ha he holo ype male is p obably los . Fu he mo e, while
Fage (1921) and Bosmans (1997) eco ded a single emale in ube MNHN-AR2851, we iden i ied wo
specimens, including one emale and one ju enile.
Dis ibu ion
I aly, Albania, No h Macedonia, Bulga ia, G eece, Tu key.
Zoda ion geshu Le y, 2007
Figs 50–51
Zoda ion geshu Le y 2007: 13, igs 33–37.
Diagnosis
Re e o Le y (2007).
Type ma e ial examined
Holo ype
ISRAEL • ♂; Geshu , S Golan (217/240 .3. J); May 1998; R. Sha on leg.; HUJ 15215.
Pa a ype
ISRAEL • 1 ♀; same collec ion da a as o holo ype; HUJ 15215.
Desc ip ion
Fo a de ailed desc ip ion, e e o Le y (2007).
Rema ks
Al hough Le y (2007) no ed ha he ype specimens we e collec ed in May 1996, he label indica es
May 1998.
Dis ibu ion
Is ael.
SHAFAIE S. e al., New da a on Middle Eas and Eas e n Medi e anean zoda iids (A aneae)
67
Fig. 50. Habi us o Zoda ion geshu Le y, 2007. A–B. Holo ype, ♂ (HUJ 15215). C–D. Pa a ype, ♀
(HUJ 15215). A. Do sal iew. B. Ven al iew. C. Do sal iew. D. Ven al iew. Scale ba s: 1 mm.
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68
Fig. 51. Copula o y o gans o Zoda ion geshu Le y, 2007. A–C. Holo ype, male palp (HUJ 15215).
D. Pa a ype, emale in ac epigyne (HUJ 15215). A. P ola e al iew. B. Ven al iew. C. Re ola e al
iew. D. Ven al iew. Scale ba s: 0.2 mm.
SHAFAIE S. e al., New da a on Middle Eas and Eas e n Medi e anean zoda iids (A aneae)
69
Zoda ion g anula um Kulczyński, 1908
Figs 52–53, 105
Zoda ion g anula um Kulczyński, 1908: 59, pl. 2 igs 4, 10.
Zoda ion g anula um – Le y 1992: 92, igs 72–84. — Bosmans 2009: 275, igs 170–171, 178–179. —
Lecigne & Hen a d 2022: 37, ig. 5e, h. — Bosmans & Ga alas 2023: 33, ig. 21a–e. — Dimi o
2024: 114, ig. 18.
Zoda ion pa ashi – Wunde lich 2022: 48, igs 105–106.
Diagnosis
Re e o Bosmans (2009) and Lecigne & Hen a d (2022).
Type ma e ial examined
Holo ype (Zoda ion pa ashi)
GREECE • ♂; Eas e n Aegean Island Naxos; 25.57° N, 37.0.37° E; 20 Jul. 1982; L. Pa ashi leg.; ZMH
A0014216.
Ma e ial examined
CYPRUS • 1 ♂ (subadul ); Nicosia, Olympus, Mou oulas Moun ain; 34.977° N, 32.828° E; 800 m a.s.l.;
25 Ap . 2022; S. Peká , S. Sha aie, V. Šol ýso á and O. Michálek leg.; CMU.
GREECE • 1 ♂; Aladjagiola We land (N Ch isoupoli); 28 Jun. 2008; M. Sch öde leg.; CMU.
SYRIA • 1 ♂; Aln Al-Za ah, nea Idlib, Canyon o O on hes; 18 Ma . 2001; M. Řezáč leg.; CMU • 1 ♀;
Mash a Al-Helu, Odem Fo es ; 3 Ap . 2001; M. Řezáč leg.; CMU.
Desc ip ion
Fo a de ailed desc ip ion, e e o Bosmans (2009).
Rema ks
The leng h and a angemen o he PVS di e be ween males and emales. Male specimens possess
signi ican ly longe PVS, which a e a anged in a somewha adia ing pa e n (Fig. 52B), whe eas
emales ha e sho e PVS a anged pa allel o each o he (Fig. 52D). Mo eo e , he single male specimen
om Sy ia exhibi s a smalle PMA compa ed o he specimen om G eece, along wi h a mo e slende
embola ip.
Dis ibu ion
This species was p e iously eco ded in G eece, Cyp us, Tu key, Lebanon, and Is ael (Wo ld Spide
Ca alog 2025). This is he i s epo o his species om Sy ia
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70
Fig. 52. Habi us o Zoda ion g anula um Kulczyński, 1908 (CMU). A–B. ♂. C–D. ♀. A. Do sal iew,
a ow indica es g anula ions. B. Ven al iew. C. Do sal iew. D. Ven al iew. Scale ba s: 1 mm.
SHAFAIE S. e al., New da a on Middle Eas and Eas e n Medi e anean zoda iids (A aneae)
71
Zoda ion hause i B ignoli, 1984
Figs 54–55
Zoda ion hause i B ignoli, 1984: 312, ig. 40.
Zoda ion hause i – S e ano ska e al. 2008: 37, igs 5–9. — Bosmans 2009: 251, igs 92–95, 144–145.
— Naumo a e al. 2017: 480, igs 28–30.
Diagnosis
Re e o Bosmans (2009).
Ma e ial examined
GREECE • 1 ♂; Aladjagiola We land (N o Ch isoupoli); 25 May 2008; M. Sch öde leg.; CMU •
5 ♂♂; E ga ikes; 37.721° N, 24.04° E; 18 May 2023; D. O iz and S. Peká leg.; CMU • 1 ♂; Oulin zi;
37.946° N, 23.834° E; 18 May 2023; D. O iz and S. Peká leg.; CMU • 3 ♂♂, 3 ♀♀; Ano Vakalopoulo;
Fig. 53. Copula o y o gans o Zoda ion g anula um Kulczyński, 1908 (CMU). A–C, F. Male palp.
D–E. Female copula o y o gan. A. P ola e al iew. B. Ven al iew. C. Re ola e al iew. D. In ac
epigyne, en al iew. E. Vul a, en al iew. F. De ail o embolus, en al iew. Scale ba s: 0.2 mm.
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72
37.996° N, 23.922° E; 18 May 2023; D. O iz and S. Peká leg.; CMU • 2 ♂♂, 1 ♀; Li adia; 38.448° N,
22.866° E; 24 May 2023; D. O iz and S. Peká leg.; CMU.
Desc ip ion
Fo a de ailed desc ip ion, e e o Bosmans (2009).
Fig. 54. Habi us o Zoda ion hause i B ignoli, 1984 (CMU). A–B. ♂. C–D. ♀. A. Do sal iew. B. Ven al
iew. C. Do sal iew. D. Ven al iew. Scale ba s: 1 mm.
SHAFAIE S. e al., New da a on Middle Eas and Eas e n Medi e anean zoda iids (A aneae)
73
Dis ibu ion
No h Macedonia, Bulga ia, G eece.
Fig. 55. Copula o y o gans o Zoda ion hause i B ignoli, 1984 (CMU). A–C. Male palp. D–E. Female
copula o y o gan. A. P ola e al iew. B. Ven al iew. C. Re ola e al iew. D. In ac epigyne, en al
iew. E. Vul a, en al iew. Scale ba s: 0.2 mm.
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80
Rema ks
The o e all body size, mo phology, and male palp s uc u e o Z. judaeo um closely esemble hose
obse ed in he genus Palaes ina. Howe e , Z. judaeo um lacks he coa se chelice al den i ion ha se es
as he dis inguishing ea u e o Palaes ina. Fu he in es iga ion is necessa y o cla i y he phylogene ic
ela ionship be ween Z. judaeo um and Palaes ina.
Dis ibu ion
Is ael.
Fig. 61. Copula o y o gans o Zoda ion judaeo um Le y, 1992. A–C. Holo ype, male palp (HUJ 14565).
D. Pa a ype, emale in ac epigyne (HUJ 14564). A. P ola e al iew. B. Ven al iew. C. Re ola e al
iew. D. Ven al iew. Scale ba s: 0.2 mm.
SHAFAIE S. e al., New da a on Middle Eas and Eas e n Medi e anean zoda iids (A aneae)
81
Zoda ion ka pa hos Bosmans, 2009
Figs 62, 105
Zoda ion ka pa hos Bosmans, 2009: 253, igs 96–99.
Diagnosis
Re e o Bosmans (2009).
Type ma e ial examined
Holo ype
GREECE • ♂; Dodekanisa, Ka pa hos, Volada; 20 Sep. 2000; B. Kno lach & K. Thale leg.; NMBE-
AR4144.
Pa a ype
GREECE • 1 ♂; same collec ion da a as o holo ype; NMBE-AR4144.
O he ma e ial examined
TURKEY • 2 ♂♂; Ispa a, Eğ idi , su ounding o Eğ idi Lake; 37.921° N, 30.918° E; 946 m a.s.l.; 10
Jun. 2007; O. Seyya leg.; CMU.
Desc ip ion
Fo a de ailed desc ip ion o male, e e o Bosmans (2009).
Dis ibu ion
This species was p e iously eco ded in G eece (Wo ld Spide Ca alog 2025). This is he i s epo o
his species om Tu key, and ep esen s, he e o e, he eas e nmos occu ence o his species.
Zoda ion killini Bosmans, 2009
Fig. 63
Zoda ion killini Bosmans, 2009: 289, igs 164–165.
Diagnosis
Re e o Bosmans (2009).
Type ma e ial examined (holo ype no examined)
Pa a ypes
GREECE • 2 ♀♀; Peloponnisos, Ko in hia, O os Killini, Pheneos Pass; 880 m a.s.l.; 19 Sep. 1993; B.
Kno lach leg.; NMBE-AR4158.
Pa a ype
GREECE • 1 ♂ (subadul ); same collec ion da a as o holo ype; NMBE-AR4158.
Desc ip ion
Fo a de ailed desc ip ion o emale, e e o Bosmans (2009).
Dis ibu ion
G eece.
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82
Fig. 62. Zoda ion ka pa hos Bosmans, 2009, holo ype, ♂ (NMBE-AR4144). A. Habi us, do sal iew.
B. Habi us, en al iew. C. Palp, p ola e al iew. D. Palp, en al iew. E. Palp, e ola e al iew. Scale
ba s: A–B = 1 mm; C–E = 0.2 mm.
SHAFAIE S. e al., New da a on Middle Eas and Eas e n Medi e anean zoda iids (A aneae)
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Zoda ion kon adi Bosmans, 2009
Figs 64–65
Zoda ion g aecum B ignoli, 1984: 311, igs 37–39.
Zoda ion kon adi Bosmans, 2009: 268, igs 132–133, 160–161.
Diagnosis
Re e o Bosmans (2009).
Fig. 63. Zoda ion killini Bosmans, 2009, pa a ype, ♀ (NMBE-AR4158). A. Habi us, do sal iew.
B. Habi us, en al iew. C. In ac epigyne, en al iew. D. Vul a, en al iew. Scale ba s: A–B =
1 mm; C–D = 0.2 mm.
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84
Type ma e ial examined
Holo ype
GREECE • ♂; Peloponnisos, Lakonia, O os Taige os, Aghios Elias; 2400 m a.s.l.; 26 Sep. 1985;
B. Kno lach and K. Thale leg.; NMBE-AR3668.
Fig. 64. Habi us o Zoda ion kon adi Bosmans, 2009. A–B. Holo ype, ♂ (NMBE-AR3668).
C–D. Pa a ype, ♀ (NMBE-AR3668). A. Do sal iew. B. Ven al iew. C. Do sal iew. D. Ven al iew.
Scale ba s: 1 mm.
SHAFAIE S. e al., New da a on Middle Eas and Eas e n Medi e anean zoda iids (A aneae)
85
Pa a ype
GREECE • ♀; O os Taige os, Moni Panaghia Gia issa; 1100 m a.s.l.; 30 Sep. 1991; B. Kno lach and
K. Thale leg.; NMBE-AR3668.
O he ma e ial examined
GREECE • 1 ♀; Co u, Agios Geo gios; 8 Ap . 2004; F. Šťáhla ský leg.; CMU.
Desc ip ion
Fo a de ailed desc ip ion, e e o Bosmans (2009).
Fig. 65. Copula o y o gans o Zoda ion kon adi Bosmans, 2009. A–C. Holo ype, male palp (NMBE-
AR3668). D. Pa a ype, emale in ac epigyne (NMBE-AR3668). A. P ola e al iew. B. Ven al iew.
C. Re ola e al iew. D. Ven al iew. Scale ba s: 0.2 mm.
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86
Rema ks
As he p osoma o he specimen was missing, i could no be pho og aphed.
Dis ibu ion
No h Macedonia, G eece.
Zoda ion kossamos Bosmans, 2009
Fig. 66
Zoda ion kossamos Bosmans, 2009: 253, igs 100–103, 146–147.
Zoda ion kossamos – Lecigne 2011: 75, igs P120–125, S4–5.
Fig. 66. Zoda ion kossomas Bosmans, 2009 (CMU), ♂. A. Habi us, do sal iew. B. Habi us, en al
iew. C. Palp, p ola e al iew. D. Palp, en al iew. E. Palp, e ola e al iew. Scale ba s: A–B = 1 mm;
C–E = 0.2 mm.
SHAFAIE S. e al., New da a on Middle Eas and Eas e n Medi e anean zoda iids (A aneae)
87
Diagnosis
Re e o Bosmans (2009).
Ma e ial examined
GREECE • 1 ♂; Kos, Ke alos; 36.744° N, 26.970° E; 100 m a.s.l.; 18 Aug. 2010; A. Exne o á leg.;
CMU.
TURKEY • 2 ♂♂; An alya, Akseki, su ounding o Al ınbeşik Ca e; 37.034° N, 31.635° E; 471 m a.s.l.;
17 Ap . 2008; O. Seyya leg.; CMU.
Desc ip ion
Fo a de ailed desc ip ion, e e o Bosmans (2009).
Dis ibu ion
G eece, Tu key.
Zoda ion le yi Sha aie & Peká sp. no .
u n:lsid:zoobank.o g:ac :F5D64ED9-3627-4905-A456-134F362A4231
Figs 67–69, 106
Diagnosis
This new species is mos simila o Z. be oni by he small body size and asymme ically bi-conca e
embola ip (Fig. 68G s Fig. 37F) bu can be dis inguished by he ollowing cha ac e s: (1) a mo e
ex ended p ola e al p ocess o median apophysis ( s elonga ed, c . Fig. 68B, E and Fig. 37B); and (2)
sligh ly con e gen embola a ms ( s di e gen a ms, c . Fig. 68G and Fig. 37F). Females o Z. le yi
sp. no . can be ecognized om he o he membe s o he honi-g oup by an ex ended median sep um
(Fig. 69A–B), whe eas i is less ex ended in Z. be oni (Fig. 37D), Z. g anula um (Fig. 53D), Z. nam un
(Fig. 82A–B), Z. e icula um (Fig. 90D), Z. hodiense (Fig. 92D), and Z. honi (Fig. 96D).
E ymology
This species is named a e he la e Ge shom Le y, an Is aeli a achnologis .
Type ma e ial
Holo ype
ISRAEL • ♂; Odem Fo es , Golan Heigh s; 1 May 1996; J. Le y leg.; HUJINV-A 21303.
Pa a ype
ISRAEL • 1 ♀; same collec ion da a as o holo ype; HUJINV-A 21304.
O he ma e ial examined
ISRAEL • 1 ♀; same collec ion da a as o holo ype; CMU • 3 ♀♀; Odem Fo es ; 33.206° N, 35.736° E;
29 Ma . 2023; S. Peká and V. Opa o á leg.; CMU • 1 ♀; same collec ion da a as o p eceding; 4 May
2023; M. Řezáč leg.; CMU • 1 ♂, 1 ♀; Ha He mon; 33.288° N, 35.764° E; 5 May 2023; M. Řezáč leg.;
CMU.
SYRIA • 1 ♂; W Homs, Al-Za ah; 28 Ma . 2001; O. Řezáč leg.; CMU.
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Desc ip ion
Male (holo ype HUJINV-A 21303)
Habi us (Fig. 67A–B). Ca apace and chelice ae ligh b own; mou hpa s basally ligh b own and apically
pale; s e num yellow. All legs yellow. Palpal segmen s yellow. Abdomen do sally black, wi h a shiny,
black scu um, en e o abdomen and spinne e s sligh ly di y yellow; PVS p esen .
Fig. 67. Habi us o Zoda ion le yi Sha aie & Peká sp. no . A–B. Holo ype, ♂ (HUJINV-A 21303).
C–D. Pa a ype, ♀ (HUJINV-A 21304). A. Do sal iew. B. Ven al iew. C. Do sal iew. D. Ven al
iew. Scale ba s: 1 mm.
SHAFAIE S. e al., New da a on Middle Eas and Eas e n Medi e anean zoda iids (A aneae)
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Measu eMen s. To al leng h 2.44; ca apace 1.25 long, 0.89 wide. Clypeus 0.14 high. Eye sizes and
in e dis ances: AME 0.13, ALE = PME 0.09, PLE 0.08, AME–AME 0.05, AME–ALE 0.03, ALE–ALE
0.29, AME–PME 0.05, PME–PME 0.33, PME–PLE 0.05, PLE–PLE 0.26, ALE–PLE 0.1. Legs: I 3.55
(0.99, 0.39, 0.72, 0.89, 0.56), II 2.97 (0.82, 0.36, 0.52, 0.75, 0.52), III 2.89 (0.79, 0.33, 0.56, 0.75, 0.46),
IV 4.31 (1.15, 0.36, 0.92, 1.22, 0.66). Palp: 0.89 (0.29, 0.13, 0.05, 0.42).
P
alP
(Fig. 68A–G). RTA b oad, wi h a claw-shaped ex ension. Median apophysis b oad, ec angula , wi h
an o al lea -shaped PMA. Embolus sho , wi h a bi u ca ed ip including wo con e gen a ms.
Fig. 68. Male palp o Zoda ion le yi Sha aie & Peká sp. no ., holo ype (HUJINV-A 21303).
A. P ola e al iew. B. Ven al iew. C. Re ola e al iew. D. P ola e al iew. E. Ven al iew.
F. Re ola e al iew. G. Embola ip, en al iew. Scale ba s: 0.2 mm.
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Fig. 74. Habi us o Zoda ion lu ipes (O. Picka d-Camb idge, 1872) (CMU). A–B. ♂. C–D. ♀. A. Do sal
iew. B. Ven al iew. C. Do sal iew. D. Ven al iew. Scale ba s: 1 mm.
SHAFAIE S. e al., New da a on Middle Eas and Eas e n Medi e anean zoda iids (A aneae)
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Fig. 75. Copula o y o gans o Zoda ion lu ipes (O. Picka d-Camb idge, 1872) (CMU). A–C. Male palp.
D–E. Female copula o y o gan. A. P ola e al iew. B. Ven al iew. C. Re ola e al iew. D. In ac
epigyne, en al iew. E. Vul a, en al iew. Scale ba s: 0.2 mm.
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98
Zoda ion milani Sha aie & Peká sp. no .
u n:lsid:zoobank.o g:ac :6E2ADB03-2768-49EC-8744-4B9488A6DCB7
Figs 76–77, 106
Diagnosis
Males o his new species a e simila o hose o Z. hodiense in body size and he colo a ion o he
p osoma and legs, bu hey can be dis inguished by hei la ened embola ip and he spine-like appendix
(Fig. 77G s Fig. 92F).
E ymology
The species is named a e ou colleague, Milan Řezáč, who collec ed he specimens.
Type ma e ial
Holo ype
SYRIA • ♂; Mash a Al-Helu; 3 Ap . 2001; M. Řezáč leg.; NMP P6A 7509.
O he ma e ial examined
SYRIA • 1 ♂; Al-Qadmus, Al-Kah Cas le; 5 Ap . 2001; M. Řezáč leg.; CMU.
Desc ip ion
Male (holo ype NMP P6A 7509)
Habi us (Fig. 76A–B). Ca apace and chelice ae b own; chelice ae wi h a longi udinal black s ipe.
S e num and mou hpa s b ownish. Cx g eyish, black dis ally; Fe black, wi h wo pai s o longi udinal
yellow s ipes; Pa I, Ti I–III, M and Ta I–IV pale. Pa II–IV and Ti IV pale en ally and da k la e ally.
Palpal emu black, wi h wo pai s o o ange s ipes, pa ella and ibia pale en ally and da k la e ally,
and cymbium b own, wi h yellow ip. Abdomen do sum black, wi h an o al scu um, en e black excep
o b ownish epiand ous; PVS p esen . Spinne e s black basally and pale apically.
Measu eMen s. To al leng h 4.55; ca apace 2.34 long, 1.58 wide. Clypeus 0.26 high. Eye sizes and
in e dis ances: AME 0.15, ALE = PME 0.1, PLE 0.08, AME–AME 0.06, AME–ALE 0.01, ALE–ALE
0.4, AME–PME 0.07, PME–PME 0.45, PME–PLE 0.1, PLE–PLE 0.27, ALE–PLE 0.2. Legs: I 7.77
(1.98, 0.72, 1.65, 2.14, 1.28), II 7 (1.91, 0.72, 1.38, 1.94, 1.05), III 7.01 (1.91, 0.69, 1.28, 2.14, 0.99), IV
9.64 (2.70, 0.75, 2.24, 2.8, 1.15). Palp: 2.37 (0.85, 0.39, 0.16, 0.95).
Fig. 76. Habi us o Zoda ion milani Sha aie & Peká sp. no ., holo ype, ♂ (NMP P6A 7509). A. Do sal
iew. B. Ven al iew. Scale ba s: 1 mm.
SHAFAIE S. e al., New da a on Middle Eas and Eas e n Medi e anean zoda iids (A aneae)
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PalP (Fig. 77A–G). RTA b oad, wi h a claw-shaped ex ension. Median apophysis ec angula , wi h a
beak-shaped PMA, companied by a blun p ola e al ex ension. Embolus wi h a la ened embola ip
accompanied by a la e al spine-like appendix.
Fig. 77. Male palp o Zoda ion milani Sha aie & Peká sp. no ., holo ype (NMP P6A 7509). A. P ola e al
iew. B. Ven al iew. C. Re ola e al iew. D. P ola e al iew. E. Ven al iew. F. Re ola e al iew.
G. Embola ip, en al iew. Scale ba s: 0.2 mm.
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100
Female
Unknown.
Size a ia ion
To al body leng h 4.55–4.57, ca apace leng h/wid h 2.34–2.35/1.58–1 in males (N = 2).
Dis ibu ion
A p esen , known only om he ype locali y in Sy ia.
Zoda ion mo osum Denis, 1935
Figs 78–79, 105
Zoda ion mo osum Denis, 1935b: 78, igs 22–24.
Zoda ion aegaeum Denis, 1935b: 81, ig. 25.
Zoda ion mo osum – Ka ol 1969: 201, igs 1–11. — Del she 1987: 19, ig. 1.1–2. — Ko blyuk 2003:
177, igs 1–8, 14–16. — Bosmans 2009: 236, igs 20–22, 54–55. — Lecigne 2011: 75, igs P125–
128, S6–7. — Se opian e al. 2023: 293, igs 136–137. — Nae 2023: 56, igs 1–6.
Diagnosis
Re e o Bosmans (2009).
Ma e ial examined
GREECE • 2 ♀♀; E Olympos, Lep oka ya, Olympic Ri e ; 9 Jun. 1996; J. Dolanský leg.; CMU • 1 ♀;
same collec ion da a as o p eceding; 22 Jun. 2008; M. Sch öde leg.; CMU • 1 ♂; Aladjagiola We land
(N o Ch isoupoli); 41.01° N, 24.703° E; 18 Jun. 2004; S. Buchholz leg.; CMU • 1 ♀; same collec ion
da a as o p eceding; 22 Jun. 2008; M. Sch öde leg.; CMU • 4 ♂♂, 1 ju .; Les os; 18 May 2006; W.
Nen wig leg.; CMU • 14 ♂♂, 2 ♀♀, 1 ju .; same collec ion da a as o p eceding; 15 Jun. 2006; W.
Nen wig leg.; CMU.
SYRIA • 1 ♀; La akia; MNHN-AR2857.
Desc ip ion
Fo a de ailed desc ip ion, e e o Bosmans (2009).
Rema ks
Examina ion o he ma e ial om he Simon collec ion (MNHN-AR2857) e ealed a emale specimen
o Z. mo osum.
Dis ibu ion
This species was o me ly eco ded om No h Macedonia, Bulga ia, Albania, G eece, Tu key, Uk aine,
and Russia (Wo ld Spide Ca alog 2025); his is he i s epo o his species om Sy ia.
SHAFAIE S. e al., New da a on Middle Eas and Eas e n Medi e anean zoda iids (A aneae)
101
Fig. 78. Habi us o Zoda ion mo osum Denis, 1935 (CMU). A–B. ♂. C–D. ♀. A. Do sal iew. B. Ven al
iew. C. Do sal iew. D. Ven al iew. Scale ba s: 1 mm.
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102
Fig. 79. Copula o y o gans o Zoda ion mo osum Denis, 1935 (CMU). A–C. Male palp. D–E. Female
copula o y o gan. A. P ola e al iew. B. Ven al iew. C. Re ola e al iew. D. In ac epigyne, en al
iew. E. Vul a, en al iew. Scale ba s: 0.2 mm.
SHAFAIE S. e al., New da a on Middle Eas and Eas e n Medi e anean zoda iids (A aneae)
103
Zoda ion nam un Dimi o , 2024
Figs 80–82, 106
Zoda ion nam un Dimi o , 2024: 116, igs 15–17, 19–20.
Fig. 80. Habi us o Zoda ion nam un Dimi o , 2024. A–B. ♂ (NMP P6A 7503). C–D. ♀ (NMP P6A
7505). A. Do sal iew. B. Ven al iew. C. Do sal iew. D. Ven al iew. Scale ba s: 1 mm.
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104
Diagnosis
Fo male diagnosis, e e o Dimi o (2024). The gene al epigynal shape o Z. nam un is simila o
ha o Z. honi, excep o he spe ma hecae s ongly con e ging pos e io ly ( s sligh ly con e ging
pos e io ly, c . Fig. 82A–B and Fig. 96D) and he size o dis al spe ma hecal duc s (smalles s wi h he
same size, c . Fig. 82C and Fig. 96E).
Ma e ial examined
TURKEY • 2 ♂♂; Kah amanma aş, Göksun, Pü en Mou ain Pass; 37.945° N, 36.511° E; 1518 m a.s.l.;
17 Jun. 2008; O. Seyya leg.; NMP P6A 7503, NMP P6A 7504 • 1 ♀; same collec ion da a as o
p eceding; NMP P6A 7505 • 1 ♂; same collec ion da a as o p eceding; CMU.
Fig. 81. Male palp o Zoda ion nam un Dimi o , 2024 (NMP P6A 7503). A. P ola e al iew. B. Ven al
iew. C. Re ola e al iew. D. P ola e al iew. E. Ven al iew. F. Re ola e al iew. G. Embola ip,
en al iew. Scale ba s: 0.2 mm.
SHAFAIE S. e al., New da a on Middle Eas and Eas e n Medi e anean zoda iids (A aneae)
105
Desc ip ion
Fo a de ailed desc ip ion o male, e e o Dimi o (2024).
Female (NMP P6A 7505)
H
abi us
(Fig. 80C–D). Ca apace dis ally yellowish-b own, an e io ly b own; clypeus and chelice ae
b own; mou hpa s yellow basally and pale apically; s e num g eyish. All legs yellow excep o en i ely
da k Fe I and II. Palpal segmen s yellow. Abdomen do sum black, wi h no pa e n; en e mos ly da k
excep o a pai o pale, con e ging pa ches and a ligh a ea a ound he copula o y o gan; PVS p esen .
Spinne e s ligh g ey basally, whi ish apically.
Measu eMen s. To al leng h 5.01; ca apace 2.21 long, 1.48 wide. Clypeus 0.33 high. Eye sizes and
in e dis ances: AME 0.12, ALE 0.1, PME = PLE 0.08, AME–AME 0.07, AME–ALE 0.02, ALE–ALE
0.41, AME–PME 0.07, PME–PME 0.45, PME–PLE 0.1, PLE–PLE 0.27, ALE–PLE 0.21. Legs: I 6.68
(1.74, 0.66, 1.45, 1.81, 1.02), II 5.96 (1.61, 0.66, 1.12, 1.65, 0.92), III 6.21 (1.81, 0.69, 1.08, 1.81, 0.82),
IV 8.25 (2.21, 0.77, 1.94, 2.44, 0.89). Palp: 1.87 (0.69, 0.33, 0.33, 0.52).
ePigyne (Fig. 82A–D). Epigyne wi h s ongly con e ging spe ma hecae; ul a comp ising h ee globula ,
oluminous and con e gen bodies, a anged by size om la ges o smalles .
Size a ia ion
To al body leng h 3.56–4.12, ca apace leng h/wid h 1.8–1.84/1.3–1.42 in males (N = 3).
Dis ibu ion
A p esen , known only om Tu key.
Fig. 82. Female copula o y o gan o Zoda ion nam un Dimi o , 2024 (NMP P6A 7505). A. In ac
epigyne, en al iew. B. Epigyne, en al iew. C. Vul a, en al iew. D. Vul a, en al iew. Scale
ba s: 0.2 mm.
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112
Diagnosis
Re e o Bosmans (2009).
Ma e ial examined
GREECE • 1 ♂; Aladjagiola We land (N o Ch isoupoli); 41.007° N, 24.703° E; 8 May 2004; S. Buchholz
leg.; CMU • 1 ♂; same collec ion da a as o p eceding; 8 July 2008; M. Sch öde leg.; CMU.
Desc ip ion
Fo a de ailed desc ip ion, e e o Bosmans (2009).
Dis ibu ion
Bulga ia, G eece.
Zoda ion e icula um Kulczyński, 1908
Figs 89–90, 105
Zoda ion e icula um Kulczyński, 1908: 61, pl. 2 ig. 16.
Zoda ion e icula um – Le y 1992: 93. — Bosmans 2009: 280, igs 174–175, 182–183. — Lecigne &
Hen a d 2022: 37, ig. 5d, g.
Diagnosis
Re e o Bosmans (2009) and Lecigne & Hen a d (2022).
Ma e ial examined
CYPRUS • 1 ♂, 1 ♀; Olympus Moun , Apliki; 34.941° N, 33.123° E; 720 m a.s.l.; 27 Ap . 2022;
S. Peká , S. Sha aie, V. Šol ýso á and O. Michálek leg.; CMU • 1 ♂ (subadul ); Olympus Moun ,
Mou oulas Moun ain; 34.977° N, 32.828° E; 800 m a.s.l.; 25 Ap . 2022; S. Peká , S. Sha aie, V. Šol ýso á
and O. Michálek leg.; CMU • 3 ju .; Le ka Dam; 35.063° N, 32.837° E; 288 m a.s.l.; 25 Ap . 2022;
S. Peká , S. Sha aie, V. Šol ýso á and O. Michálek leg.; CMU.
GREECE • 1 ♀; G e ena, Kalami si; 15 Aug. 1999; CMU.
TURKEY • 2 ♂♂, 1 ♀; Konakli, Macchie Hill, 10 km W o Alanya; 20 May 2005; J. Dolanský leg.;
CMU • 2 ♂♂, 1 ♀; same collec ion da a as o p eceding; 19 May 2005; J. Dolanský leg.; CMU • 1 ♀;
s. loc.; 21 Jun. 2003; CMU • 1 ♀; E zincan, Çağlayan; 39.589° N, 39.705° E; 1541 m a.s.l.; 24 Jun.
2002; M. Řezáč leg.; CMU.
Desc ip ion
Fo a de ailed desc ip ion, e e o Lecigne & Hen a d (2022).
Rema ks
As in Z. g anula um, males o Z. e icula um possess longe PVS, a anged in a somewha adia ing
pa e n (Fig. 89B), whe eas emales exhibi sho e , mo e pa allel PVS s uc u es (Fig. 89D).
Dis ibu ion
P e ious s udies eco ded Z. e icula um in Cyp us and G eece (Wo ld Spide Ca alog 2025). This is he
i s epo o his species om Tu key and ep esen s, he e o e, he no hwes e nmos occu ence o
his species ac oss i s dis ibu ion ange.
SHAFAIE S. e al., New da a on Middle Eas and Eas e n Medi e anean zoda iids (A aneae)
113
Fig. 89. Habi us o Zoda ion e icula um Kulczyński, 1908 (CMU). A–B. ♂. C–D. ♀. A. Do sal iew.
B. Ven al iew. C. Do sal iew. D. Ven al iew. Scale ba s: 1 mm.
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114
Zoda ion hodiense Capo iacco, 1948
Figs 91–92, 105
Zoda ion hodiense Capo iacco, 1948: 47, ig. 5.
Zoda ion hodiense – B ignoli 1984: 317, igs 50–53. — Dimi o 2024: 113, igs 1–9.
Zoda ion hodiense nig i emu – Capo iacco 1948: 49.
Fig. 90. Copula o y o gans o Zoda ion e icula um Kulczyński, 1908 (CMU). A–C. Male palp.
D–E. Female copula o y o gan. A. P ola e al iew. B. Ven al iew. C. Re ola e al iew. D. In ac
epigyne, en al iew. E. Vul a, en al iew. F. De ail o embolus, en al iew. Scale ba s: 0.2 mm.
SHAFAIE S. e al., New da a on Middle Eas and Eas e n Medi e anean zoda iids (A aneae)
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Zoda ion nig i emu – Bosmans 2009: 279, igs 172–173, 180–181. — Lecigne & Hen a d 2022: 33,
igs 1a–d, 2a–g, 3a–e, 4a–h, 5a. — Se opian e al. 2023: 293, igs 138–140. — Dimi o 2024: 113,
igs 1–9.
Zoda ion honi – Lecigne 2021: 34, ig. 55a–b.
Diagnosis
Re e o Lecigne & Hen a d (2022) and Dimi o (2024).
Fig. 91. Habi us o Zoda ion hodiense Capo iacco, 1948 (CMU). A–B. ♂. C–D. ♀. A. Do sal iew.
B. Ven al iew. C. Do sal iew. D. Ven al iew. Scale ba s: 1 mm.
Eu opean Jou nal o Taxonomy 1016: 1–139 (2025)
116
Ma e ial examined
CYPRUS • 4 ♀♀, 5 ju .; Ky enia Moun ain ange, Sain Hila ion Cas le, Sysklipos (Akcicek); 35.309° N,
33.202° E; 329 m a.s.l.; 27 Ap . 2022; S. Peká , S. Sha aie, V. Šol ýso á and O. Michálek leg.; CMU.
GREECE • 2 ♀♀, 1 ju .; Les os; 39.220° N, 26.291° E; 967 m a.s.l.; 15 Jun. 2006; W. Nen wig leg.;
CMU.
SYRIA • 2 ♀♀; Hanna A ea; 11 Ap . 2001; M. Řezáč leg.; CMU.
TURKEY • 1 ♀; Ispa a, Aksu, Dedegöl Moun ain; 37.656° N, 31.291° E; 8 Jul. 2003; M. Řezáč leg.;
CMU.
Desc ip ion
Fo a de ailed desc ip ion, e e o Lecigne & Hen a d (2022).
Fig. 92. Copula o y o gans o Zoda ion hodiense Capo iacco, 1948 (CMU). A–C, F. Male palp.
D–E. Female copula o y o gan. A. P ola e al iew. B. Ven al iew. C. Re ola e al iew. D. In ac
epigyne, en al iew. E. Vul a, en al iew. F. De ail o embolus, en al iew. Scale ba s: 0.2 mm.
SHAFAIE S. e al., New da a on Middle Eas and Eas e n Medi e anean zoda iids (A aneae)
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Dis ibu ion
This species was p e iously epo ed om Geo gia, G eece and Tu key; hese a e he i s epo s o his
species om Cyp us and Sy ia, and ep esen , he e o e, he sou he nmos occu ences o his species
ac oss i s dis ibu ion ange.
Zoda ion spiniba bis Wunde lich, 1973
Figs 93–94
Zoda ion spiniba bis Wunde lich, 1973: 173, igs 4–10.
Zoda ion spiniba bis – B ignoli 1984: 319, ig. 45.
Zoda ion spiniba be – Bosmans 2009: 244, igs 72–75, 136–137. — Coşa e al. 2022: 232, igs 22–32.
— Coşa & Danışman 2024: 268, igs 23–24.
Fig. 93. Habi us o Zoda ion spiniba bis Wunde lich, 1973 (CMU). A–B. ♂. C–D. ♀. A. Do sal iew.
B. Ven al iew. C. Do sal iew. D. Ven al iew. Scale ba s: 1 mm.
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118
Diagnosis
Re e o Bosmans (2009).
Ma e ial examined
GREECE • 1 ♀; Ky hi a, Agia Pelagia; 36.325° N, 22.982° E; 383 m a.s.l.; 28 May 2003; J. Dolanský
leg.; CMU.
TURKEY • 1 ♂, 1 ♀; Kah amanma aş, Andi an, Geben, Deği mende e; 37.889° N, 36.448° E;
1518 m a.s.l.; 15 May 2008; O. Seyya leg.; CMU.
Desc ip ion
Fo a de ailed desc ip ion, e e o Bosmans (2009).
Dis ibu ion
G eece and Tu key.
Zoda ion honi Nosek, 1905
Figs 95–96, 105
Zoda ion honi Nosek, 1905: 128, ig. 10.
Re e o Wo ld Spide Ca alog 2025.
Fig. 94. Copula o y o gans o Zoda ion spiniba bis Wunde lich, 1973 (CMU). A–C. Male palp, wi h
o a ed bulb. D–E. Female copula o y o gan. A. P ola e al iew. B. Ven al iew. C. Re ola e al iew.
D. In ac epigyne, en al iew. E. Vul a, en al iew. Scale ba s: 0.2 mm.
SHAFAIE S. e al., New da a on Middle Eas and Eas e n Medi e anean zoda iids (A aneae)
119
Diagnosis
Re e o Bosmans (2009) and Lecigne & Hen a d (2022).
Ma e ial examined
CYPRUS • 2 ♂♂, 1 ♀; La chi, The Olympians La chi Beach Villas; 35.041° N, 32.413° E; 7 m a.s.l.;
7–27 Aug. 2021; K.G. Mikhailo leg.; ZMMU.
Fig. 95. Habi us o Zoda ion honi Nosek, 1905 (CMU). A–B. ♂. C–D. ♀. A. Do sal iew. B. Ven al
iew. C. Do sal iew. D. Ven al iew. Scale ba s: 1 mm.
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GREECE • 2 ♂♂; Kos, Ke alos; 36.744° N, 26.970° E; 100 m a.s.l.; 18 Aug. 2010; A. Exne o á leg.;
CMU.
IRAN • 1 ♂, 1 ♀; U mia, Band; 37.461° N, 44.932° E; 1400 m a.s.l.; 29 Jun. 2022; S. Sha aie leg.; CMU.
TURKEY • 2 ♀♀; Ag i, Dogubeyazi ; 39.547° N, 44.083° E; 1625 m a.s.l.; 16 May 1997; V. B yja leg.;
CMU • 1 ♂, 1 ♀; Tunceli, Munzu Ri e ; 38.957° N, 39.561° E; 22 Jun. 2002; J. Dolanský leg.; CMU •
1 ♂; Si as, Demi yu ; 39.883° N, 37.6° E; 1542 m a.s.l.; 28 Jun. 2002; J. Dolanský leg.; CMU • 2 ♂♂,
1 ♀; Toka , Paza , Ballica Ca e; 40.227° N, 36.301° E; 1085 m a.s.l.; 1 Jul. 2002; J. Dolanský leg.;
CMU • 1 ♀; Konakli, Macchie Hill, 10 km W o Alanya; 36.589° N, 31.875° E; 800 m a.s.l.; 20 May
2005; J. Dolanský leg.; CMU • 5 ♂♂; Kayse i, Yahyali, nea he Kapuzbaşı Wa e all; 26 May 2007;
O. Seyya leg.; CMU • 1 ♂, 1 ♀, 31 ju .; Canki i, Anka a-Canki i Highway; 40.389° N, 33.570° E;
689 m a.s.l.; 15 Sep. 2010; Y.M. Ma usik leg.; ZMUT • 4 ju .; Bilecik, Mezi ; 39.950° N, 29.701° E;
500 m a.s.l.; 26 Sep. 2010; Y.M. Ma usik leg.; ZMUT.
Fig. 96. Copula o y o gans o Zoda ion honi Nosek, 1905 (CMU). A–C, F. Male palp. D–E. Female
copula o y o gan. A. P ola e al iew. B. Ven al iew. C. Re ola e al iew. D. In ac epigyne, en al
iew. E. Vul a, en al iew. F. De ail o embolus, en al iew. Scale ba s: 0.2 mm.
SHAFAIE S. e al., New da a on Middle Eas and Eas e n Medi e anean zoda iids (A aneae)
121
Desc ip ion
Fo a de ailed desc ip ion, e e o Lecigne & Hen a d (2022).
Dis ibu ion
P e ious s udies eco ded his species in Eas e n Eu ope, Cyp us, Tu key, he Caucasus, and Lebanon.
The cu en s udy epo ed his species om G eece and I an o he i s ime.
Zoda ion une iacum S and, 1906
Figs 97–98, 106
Zoda ium une iacum S and, 1906: 669.
Zoda ium une iacum – S and 1908: 86.
Zoda ion une iacum – Denis 1937: 17, pl. 3 ig. 21.
Diagnosis
This species can be sepa a ed om Z. cy enaicum by he p esence o a ho izon ally cu ly b ace-shaped
p o ube ance in he middle o epigyne ( s absen in Z. cy enaicum; c . Fig. 98A–B and Fig. 46D).
Type ma e ial examined
Neo ype (he e designa ed)
EGYPT • ♀; Alexand ia; MNHN-AR2854.
Rema ks
Examina ion o ma e ial om he Simon collec ion (MNHN-AR2854) led o he iden i ica ion o a
emale specimen belonging o Z. une iacum. Al hough his species has no been lis ed as a nomen
Fig. 97. Habi us o Zoda ion une iacum S and, 1906. A–B. Neo ype, ♀ (MNHN-AR2854). A. Do sal
iew. B. Ven al iew. Scale ba s: 1 mm.
Eu opean Jou nal o Taxonomy 1016: 1–139 (2025)
128
O he ma e ial examined
CYPRUS • 1 ♂ (subadul ), 2 ju .; Agios Geo gios; 34.822° N, 32.897° E; 840 m a.s.l.; 29 Ap . 2022;
S. Peká , S. Sha aie, V. Šol ýso á and O. Michálek leg.; CMU.
ISRAEL • 16 ju .; same collec ion da a as o holo ype; CMU • 7 ju .; Geshu ; 32.816° N, 35.718° E;
30 Ma . 2023; S. Peká leg.; CMU • 1 ♂, 8 ju .; Kada im; 32.894° N, 35.478° E; 31 Ma . 2023; S. Peká
and V. Opa o á leg.; CMU • 1 ju .; Ha He on; 32.996° N, 35.410° E; 3 May 2023; M. Řezáč leg.; CMU.
Desc ip ion
Male (holo ype NMP P6A 7510)
Habi us (Fig. 103A–B). Ca apace eddish-b own companied by a ligh e , bu e ly-like ma k on behind
o o ea; clypeus, chelice ae eddish-b own; labium and gna hocoxae b ownish, wi h ligh e ips.
S e num yellow a he middle and sligh ly da ke a bo de s. Cx I and Fe I and II en i ely da k, Cx II
da k on la e al sides, Cx III and IV yellowish, Fe III and IV ligh b own p oximally and da ke dis ally,
wi h a pai o longi udinal, ligh s ipes. Pa and Ti o all legs yellow, wi h g eyish la e als, o he segmen s
en i ely yellow. Palpal segmen s yellow, wi h pale cymbium. Abdomen do sum en i ely da k, en e da k
excep o ligh e pa s a ound he epiand ous and PVS. Spinne e s pale, wi h da k basal pa .
M
easu eMen s
. To al leng h 4.49; ca apace 2.16 long, 1.76 wide. Eye sizes and in e dis ances: AME
0.15, ALE 0.11, PME = PLE 0.08, AME–AME 0.1, AME–ALE 0.02, ALE–ALE 0.36, PME–PME 0.23,
PLE–PLE 0.4, PME–PLE 0.03. Legs: I 9.59 (2.6, 0.8, 2.26, 2.53, 1.4), II 9.19 (2.53, 1, 2.13, 2.4, 1.13),
III 8.38 (2.26, 0.73, 1.66, 2.6, 1.13), IV 10.06 (3, 1, 2.53, 3.33, 1.2). Palp: 3.26 (1, 0.46, 0.2, 1.6).
PalP (Fig. 104A–C). Palp en i ely simila o ha o Z. lu ipes bu ma kedly la ge ( o a de ailed
desc ip ion, e e o Le y 1992: c . igs 67–69 and Bosmans 2009: c . igs 184–185).
Fig. 103. Habi us o Zoda ion zons eini Sha aie & Peká sp. no ., holo ype, ♂ (NMP P6A 7510).
A. Do sal iew. B. Ven al iew. Scale ba s: 1 mm.
SHAFAIE S. e al., New da a on Middle Eas and Eas e n Medi e anean zoda iids (A aneae)
129
Rema ks
Al hough no s uc u al di e ences we e de ec ed in he copula o y o gans o Z. zons eini sp. no . and
Z. lu ipes, he wo ep esen a i es exhibi no able dispa i ies in o e all body and copula o y o gan
dimensions. Pa icula ly, he ca apace and cymbium o Z. zons eini a e 1.31 and 1.76 imes longe ,
espec i ely, han hose o Z. lu ipes. Such di e gence in absolu e copula o y o gan dimensions may
lead o mechanical incompa ibili y be ween sexes, po en ially impeding success ul copula ion (Ramos
e al. 2005). Fo ins ance, he ela i ely la ge palpal o gans o Z. zons eini migh no be e ec i ely
pai ed wi h he smalle , hea ily scle o ized epigyne o Z. lu ipes. Mo eo e , he wo species di e in
soma ic mo phology, wi h Z. lu ipes exhibi ing a dis inc bicolo ed ca apace and ligh e legs compa ed
o Z. zons eini. Simila pa e ns o mo phological di e gence ha e p e iously been obse ed in
Z. ni idum and Z. nicki ( e e o Peká e al. 2022 and co esponding commen s in his s udy). Based
on hese consis en mo phological di e ences, we ea Z. zons eini and Z. lu ipes as sepa a e species.
Ne e heless, u he examina ion o addi ional ma e ial will be necessa y o ully esol e issues o
species delimi a ion and axonomic consis ency.
Dis ibu ion
A p esen , known only om Is ael and Cyp us.
Fig. 104. Male palp o Zoda ion zons eini Sha aie & Peká sp. no ., holo ype (NMP P6A 7510).
A. P ola e al iew. B. Ven al iew. C. Re ola e al iew. Scale ba s: 0.2 mm.
Eu opean Jou nal o Taxonomy 1016: 1–139 (2025)
130
Discussion
Al hough se e al a achnologis s ha e e ised he zoda iid species o he Middle Eas and Eas e n
Medi e anean (Le y 1992; Bosmans 1994, 1997, 2009; Zamani & Ma usik 2021), ou knowledge on
he axonomy o hese spide s in his egion emains agmen a y. The p ima y issue s ems om he as
geog aphical a ea ha hos s many species, and he ela i ely ew a achnologis s wo king in he egion.
Mo eo e , poli ical ins abili y in some coun ies u he hampe s p og ess in he esea ch.
Ano he signi ican issue lies in he o e eliance on speci ic diagnos ic cha ac e s. Ea ly axonomic
s udies in he 19
h
and ea ly 20
h
cen u ies p ima ily concen a ed on soma ic cha ac e s o zoda iids,
whe eas la e esea ch shi ed i s emphasis o copula o y ai s. This ansi ion ini ially led o a
p oli e a ion o ecognized species, ollowed by a subsequen axonomic educ ion. Al hough he
emphasis on copula o y o gan mo phology has p o en eliable, an exclusi e ocus on hese ai s – a
he expense o soma ic ai s and body dimensions – may hinde consensus on which cha ac e s should
ecei e p io i y and wha deg ee o a ia ion is accep able o de ining species bounda ies.
Fig. 105. Dis ibu ion map showing new coun y eco ds ac oss he Middle Eas and Eas e n
Medi e anean, one sign may e e o se e al poin s, each numbe may e e o se e al locali ies. 1 =
Acan hinozodium odem (Le y, 2007); 2 = A. ibes i Jocqué, 1991; 3 = Zoda ion aban ense Wunde lich,
1980; 4 = Z. g anula um Kulczyński, 1908; 5 = Z. ka pa hos Bosmans, 2009; 6 = Z. mo osum Denis,
1935; 7 = Z. e icula um Kulczyński, 1908; 8 = Z. hodiense Capo iacco, 1948; 9 = Z. honi Nosek,
1905.
SHAFAIE S. e al., New da a on Middle Eas and Eas e n Medi e anean zoda iids (A aneae)
131
Recen s udies ha ques ioned his long-held adi ional app oach was conduc ed by Peká e al. (2022)
and O iz e al. (2024). P e iously, Z. nicki has been conside ed a synonym o Z. ni idum, wi h hei
dis inc body colo a ion – blackish in Z. nicki and yellowish in Z. ni idum – in e p e ed as in aspeci ic
a ia ion. Howe e , Peká e al. (2022) and O iz e al. (2024) p o ided compelling e idence o he
impo ance o soma ic cha ac e s in dis inguishing hese wo species o Zoda ion. Al hough copula o y
o gans did no p o ide a clea di e en ia ion, molecula , ecological, and beha io al da a e ec i ely
sepa a ed he species, co obo a ing he dis inc ion based on soma ic ai s and a i ming he alidi y o
bo h species.
Ano he issue is he quali y o a ailable illus a ions. Many species ha e been desc ibed p ima ily using
copula o y o gan d awings. While hese illus a ions a e axonomically aluable, hey can a guably
in oduce inaccu acies and inconsis encies in species iden i ica ion, as hei in e p e a ion is o en
subjec i e and in luenced by he axonomis ’s pe spec i e.
In his s udy, we documen ed 50 zoda iid ep esen a i es, p o iding de ailed illus a ions and pho og aphs
o bo h soma ic and copula o y o gans. Following he e ised axonomy p esen ed he e, he zoda iid
auna o he Middle Eas and Eas e n Medi e anean now includes he ollowing gene a, wi h he numbe
o species known om each indica ed in pa en heses: Acan hinozodium (14), Cyd ela Tho ell, 1873 (1),
Dusmadio es Jocqué, 1987 (1), Indozodion (1), Lachesana (11), Palaes ina (3), Pa azoda ion (1), Pax
Fig. 106. Dis ibu ion eco ds ac oss he Middle Eas and Eas e n Medi e anean. 1 = Indozodion
lindbe gi (Roewe , 1960); 2 = Lachesana ga ishae Sha aie & Peká sp. no .; 3 = L. lubinae Sha aie &
Peká sp. no .; 4 = Pa azoda ion addei (Simon, 1889); 5 = Pax akilae Sha aie & Peká sp. no .;
6 = Zoda ion c ewsae Coşa , Danışman & Kun , 2022; 7 = Z. jakubi Sha aie & Peká sp. no .; 8 =
Z. le yi Sha aie & Peká sp. no .; 9 = Z. milani Sha aie & Peká sp. no .; 10 = Z. nam un Dimi o ,
2024; 11 = Z. nicki S and, 1914; 12 = Z. ni idum Audouin, 1826; 13 = Z. une iacum S and, 1906;
14 = Z. zons eini Sha aie & Peká sp. no .
Eu opean Jou nal o Taxonomy 1016: 1–139 (2025)
132
(8), T yge us (6), Zoda iellum And ee a & Tys shenko, 1968 (1), and Zoda ion (75). The highes species
ichness is obse ed in G eece (43), Tu key (40), Is eal (26), and I an (22) (Fig. 107).
In conclusion, his s udy ep esen s a ho ough su ey o he axonomy o zoda iid spide s in he Middle
Eas e n and Eas Medi e anean. To achie e a mo e p ecise es ima e o he ac ual numbe o species,
i is c ucial o in eg a e comp ehensi e mo phological documen a ion wi h da a de i ed om mul iple
complemen a y app oaches.
Acknowledgmen s
We wish o exp ess ou wa mes hanks o C. K op (Swi ze land) o p o iding us wi h he ype
specimens o Z. ba ba ae, Z. ka pa hos, Z. killini and Z. kon adi; o W. Wins and T. Szű s (Hunga y) o
he ype o Z. aculea um; o P. Jäge and J. Al mann (Ge many) o he ype o Z. nicki; o Z. Simmons
and R. Douglas (UK) o he ypes o Z. expe s, Z. luc uosum and Z. lu ipes; o E. Ga ish-Rege
(Is ael) o he ypes o P. e emica, P. engediensis, P. palmonii, Z. geshu , Z. judaeo um and Z. odem; o
C. Rolland and J. Pé illon (F ance) o o ganizing he i s au ho ’s s ay in Pa is and loaning impo an
ma e ial; o D. Ha ms and N. Dupé é (Museum o Na u e Hambu g, Ge many) o p epa ing he igu es
o Z. pa ashi; o J. Mi áček (Czech Republic), O. Mi shamsi (I an), P. Pan ini (I aly), Y.M. Ma usik and
K.G. Mikhailo (Russia), and M. Cha zaki (G eece) who kindly sen us he ma e ial deposi ed in hei
collec ions. We a e pa icula ly g a e ul o R. Jocqué (Belgium), A. Hen a d and an anonymous e iewe
o cons uc i e commen s on an ea lie e sion o he manusc ip . We a e also g a e ul o S. Ko enko,
E. Lízna o á, O. Michálek, D. O iz, L. Sen enská, and V. Šol ýso á o hei kind help in he ield; and
o M. Řezáč, F. Šťáhla ský, J. Dolanský, E. Ga ish, Y. Lubin, G. Le y, and A. Exne o á o p o iding
hei ma e ial. Figu es om Denis (1937, 1962) a e ep oduced wi h he pe mission o he publishe s.
The s udy was suppo ed by he Czech Science Founda ion g an no. 22-20229S.
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