A con ibu ion o he knowledge o he genus Thy eus Panze in he
Wes and Cen al Palaea c ic (Hymenop e a, Apidae), wi h wo new
species, axonomic upda es, hos ela ionships,
and a key o Eu opean species
Thomas Wood1, Vincen Lecle cq2, Ch is ian Schmid-Egge 3, Ch is ophe P az4,5
1 Na u alis Biodi e si y Cen e , Da winweg 2, 2333 CR, Leiden, Ne he lands
2 Obse a oi e des Abeilles, 10 D Rue Al ed Ma paux 21000 Dijon, F ance
3 Fische s aße 1, 10317, Be lin, Ge many
4 Uni e si y o Neucha el, Neuchâ el, Swi ze land
5 In o Fauna - Swiss Zoological Reco ds Cen e , Neuchâ el, Swi ze land
h ps://zoobank.o g/8DD12B45-AE34-45B8-A8FB-0C3A13478718
Co esponding au ho : Thomas Wood ([email p o ec ed])
Academic edi o : Dominique Zimme mann ♦
Recei ed
8 July 2025 ♦
Accep ed
10 Oc obe 2025 ♦
Published
4 No embe 2025
Abs ac
Thy eus Panze , 1806, is a mode a ely sized bee genus con aining abou 115 species wi h an Old Wo ld dis ibu ion ma ching ha
o hei p incipal hos s, bees o he genus Amegilla F iese, 1897. The Palaea c ic auna was e ised du ing he mid-20 h cen u y bu
has ecei ed ela i ely li le a en ion since hen. New collec ions, examina ion o museum specimens, and DNA ba coding ha e
imp o ed ou unde s anding o his genus in he Wes and Cen al Palaea c ic. The p e iously unknown sexes o T. pa henope Lie -
inck, 1968, T. pica on Lie inck, 1968, and T. p iesne i Lie inck, 1968, a e iden i ied and cla i ied. New hos da a a e p esen ed o
Thy eus hellenicus Lie inck, 1968, in G eece, whe e i pa asi ises An hopho a (Pa amegilla) supe ans Walke , 1871 = An hopho a
(Pa amegilla) incly a Walke , 1871 syn. no . (= An hopho a dubia E e smann, 1852 sensu auc o um). A lec o ype is designa ed o
Thy eus unca us (Pé ez, 1884), and i s ange and s a us a e discussed. New eco ds a e p esen ed o esol e he dis ibu ional anges
o con used o poo ly s udied species, along wi h new hos da a. Thy eus p ae alens (Kohl, 1905) is newly epo ed o Eu ope om
G eece. Specimens p e iously e e ed o as “T. pica on” in Cen al Asia do no belong o ha species and a e newly desc ibed as
Thy eus jansseni Wood, sp. no . (Ky gyzs an and A ghanis an). A u he new species in he scu ella is g oup is desc ibed, Thy eus
imp essus Wood, sp. no . (Ky gyzs an). A modi ied and upda ed iden i ica ion key o he 12 Eu opean membe s o he genus is p e-
sen ed, wi h he aim o s imula ing wo k on his challenging bee genus.
Key Wo ds
DNA ba coding, hos -pa asi e ela ionship, iden i ica ion key, museum collec ions, new species
In oduc ion
The genus Thy eus is he mos species- ich membe o
he obliga ely pa asi ic bee ibe Melec ini, wi h abou
115 species known ac oss he Old Wo ld, ep esen ing
app oxima ely 55% o he known species ichness (Mi-
chene 2007; Asche and Picke ing 2025). Melec ine
bees p esen nume ous axonomic issues, bo h in e ms
o how gene a a e de ined and ecognised (e.g. Lie inck
1972, 1983; Righ mye and Engel 2003; Michene 2007;
Onu e ko e al. 2021; O e al. 2024) and a he species
le el. Due o hei pa asi ic na u e, p edominan ly a ack-
ing bees o he sub amily An hopho inae (see Bosse
e al. 2019; O e al. 2024), melec ine species exhibi
D sch. En omol. Z. 72 (2) 2025, 259–302|DOI 10.3897/dez.72.164496
Copy igh Thomas Wood e al. This is an open access a icle dis ibu ed unde he e ms o he C ea i e Commons A ibu ion License (CC BY 4.0), which pe mi s un es ic ed use,
dis ibu ion, and ep oduc ion in any medium, p o ided he o iginal au ho and sou ce a e c edi ed.
dez.penso .ne
Thomas Wood e al.: A con ibu ion o Thy eus in he Palaea c ic, wi h wo new species260
la gely conse ed and homogeneous mo phology, en-
de ing species ecogni ion and delinea ion challenging.
As he la ges genus o Melec ini, his s a emen na u al-
ly applies o Thy eus. The genus was in a la gely chao ic
s a e un il he e isiona y wo ks o Lie inck (mainly 1959a,
1962, 1968), who deal wi h he Palaea c ic, Indo-Malay-
an, and Aus alasian aunas. Lie inck ne e add essed he
A o opical auna, and whils a e ision has been conduc ed
(Ea dley 1991), addi ional wo k is needed due o he long
his o y o s udy, ancien ype ma e ial, and complex na u e
o he genus. Wi hin he Palaea c ic auna, Lie inck’s (1968)
e ision esol ed many long- unning issues. Olde wo ks
such as Meye (1921), published be o e Lie inck’s e ision,
a e la gely unusable due o he high deg ee o axonomic
unce ain y, lack o ype e ision, axa desc ibed om ma-
e ial ha is now los , and gene ally muddled concep s; his
can be seen when examining his o ical specimens iden i ied
by Meye , hese o en being comple ely misiden i ied (TJW,
pe s. obs.). De Beaumon (1940) made a majo con ibu ion
by wo king on he Wes Eu opean auna and cla i ying he
inco ec applica ion o he name Thy eus scu ella is (Fa-
b icius, 1781), and hese imp o emen s we e cemen ed by
Lie inck’s mo e comp ehensi e wo k.
Since hen, he genus has ecei ed ela i ely li le ax-
onomic a en ion in he Palaea c ic (Ma iko skaya 1992;
Schwa z 1993), wi h compa a i ely mo e a en ion paid
o he A o opical auna (Ea dley 1991; S aka and Engel
2012; Alqa ni e al. 2014; Engel 2014). Howe e , ecen
wo k has demons a ed genuine ange expansions (Schmid-
Egge e al. 2022) and axonomic complexi y (Wood 2023)
in he auna o Eu ope and he Medi e anean basin. The e
has also been an oppo uni y o s udy he Wes and Cen al
Palaea c ic auna using DNA ba codes, which p o ided
bo h cla i y and in oduced new complexi y. The pu pose o
he p esen wo k is o p esen hese new da a on he genus
Thy eus in he Wes and Cen al Palaea c ic and o p oduce
an imp o ed iden i ica ion key o he Eu opean auna in o -
de o s imula e new wo k on his complex g oup.
Me hodology
Gene ics
Collec ed Wes and Cen al Palaea c ic Thy eus speci-
mens we e ba coded using di e en echniques. Fo mos
specimens (sequence p e ix WPATW), a single midleg
was emo ed om pinned specimens and sen o he Ca-
nadian Cen e o DNA Ba coding (CCDB) in Guelph,
Canada, o DNA ex ac ion and sequencing; specimens
we e sequenced ollowing s anda dised high- h oughpu
p o ocols (I ano a e al. 2006). BeeCox1F1/BeeCox1R2
p ime s (Bleido n and Henze 2021) we e used o a ge
he COI-5 egion. Sequences wi h he p e ix BCHYM
we e gene a ed ollowing he me hodology o Schmid
e al. (2015). Sequences wi h he p e ix HYMAA we e
gene a ed a he Uni e si y o Neuchâ el, Swi ze land,
ollowing he me hodology ou lined in P az e al. (2019),
using he o wa d p ime s LepF o UAE3 wi h he e e se
p ime LepF (see P az e al. 2019).
Phylogene ic ees we e supplemen ed wi h addi ional
published sequences (e.g. Schmid e al. 2015; Wood 2023;
Wood e al. 2024) ha we e downloaded om GenBank
(h ps://www.ncbi.nlm.nih.go /genbank/) and he Ba code
o Li e Da a Sys em. Also used we e newly gene a ed se-
quences om he Biodi e si y Genomics Eu ope p ojec ,
based on specimens om he Na u alis Biodi e si y Cen e
ha we e selec ed and cu a ed by TJW. Sequences a e a ail-
able a : h ps://po al.boldsys ems.o g/ esul ?que y=BGEN-
L[ eco dse code]. Taxa we e selec ed om wi hin he Wes
and Cen al Palaea c ic amewo k o his s udy (de ined be-
low), wi hou including any o he many addi ional a ailable
ex alimi al sequences o Thy eus. Sequences we e aligned
using MAFFT (Ka oh and S andley 2013). Aligned se-
quences we e analysed in Sea iew (Gouy e al. 2010) using
a maximum-likelihood analysis, which was un wi h 1,000
boo s aps. In a- and in e speci ic dis ances we e calcula ed
using MEGA-X (Kuma e al. 2018).
Mo phological and axonomic concep s
Mo phological e minology ollows Michene (2007),
excep o he ma ginal “zones” o he e ga, which a e
e e ed o as ma ginal “a eas”; an ennome es, which a e
numbe ed 1–12 ( emale) and 1–13 (male) and abb e ia -
ed as “A”; and he use o he wo d “ hina ia” o e e o
he imp essed pi s on he pos e io ace o he an ennal
segmen s (see Lie inck 1968). Me asomal e ga a e ab-
b e ia ed as “T”, and me asomal s e na as “S”. Fo one
diagnosis, he ela i e b ead h o he scu ellum is p e-
sen ed. This was measu ed as he dis ance be ween he
apical poin s o he axillae ( he b ead h, i.e. he maximum
wid h o he scu ellum) di ided by he dis ance be ween
he base o he scu ellum (s aigh junc ion wi h he scu-
um) and he pos e io scu ella poin s ( he leng h, i.e. he
maximum leng h o he scu ellum). Measu emen s we e
ob ained om pho og aphs, no using a calib a ed mic o-
scope, and hence a e p esen ed as a a io.
Unless explici ly s a ed, all specimens we e iden i ied
by TJW. An iden i ica ion key is p esen ed o Thy eus spe-
cies occu ing in Eu ope as de ined in Ghisbain and Rosa
e al. (2023), o acili a e hei iden i ica ion. Pu simply,
his includes Eu ope eas o he U al Moun ains, Eu ope-
an Russia sou h o bu no including he No h Caucasus,
Tu key in Eu ope (eas o he Bospo us), all Medi e anean
islands including Cyp us, and he Maca onesian islands
excluding he Cape Ve de islands. Whils he Wes and
Cen al Palaea c ic a e poo ly and inconsis en ly de ined
in he li e a u e, om he pe spec i e o he genus Thy eus,
hese wo a eas oge he a e conside ed o include No h
A ica, he A abian Peninsula, I an, and Cen al Asia (all
“-s an” coun ies, including A ghanis an and Pakis an).
Pho og aphs we e aken using an Olympus E-M1
Ma k II wi h a 60 mm mac o lens. Addi ional close-ups
we e aken wi h he addi ion o a Mi u oyo M Plan Apo
D sch. En omol. Z. 72 (2) 2025, 259–302
dez.penso .ne
261
10X in ini y-co ec ed objec i e lens in combina ion wi h
an Olympus M.Zuiko 2× elecon e e lens, a 10 mm
Kenko DG ex ension ube, and a Meike MK-P-AF3B
10 mm ex ension ube. Pho og aphs we e s acked using
Helicon Focus B (HeliconSo , Uk aine), and pla es we e
p epa ed in GNU Image Manipula ion P og am (GIMP)
2.10. Pos -p ocessing o some images was ca ied ou in
Pho oshop Elemen s (Adobe Sys ems, USA) o imp o e
ligh ing and highligh speci ic cha ac e s.
Resul s a e p esen ed in he ollowing o de : (1) gene -
ics, (2) desc ip ion o new species, (3) p oblema ic axa
iden i ied by gene ic s udy, (4) desc ip ion o missing
sexes, (5) new dis ibu ional da a and ange cla i ica ions,
and (6) new iden i ica ion key o Eu opean species. In
he summa y o dis ibu ions, new coun y eco ds a e
ma ked wi h an as e isk “*”. New da a on hos ela ion-
ships a e p esen ed o indi idual species whe e su icien
in o ma ion is a ailable o d aw con incing conclusions.
Abb e ia ions o specimen eposi o ies
CSE Pe sonal collec ion o Ch is ian Schmid-
Egge , Be lin, Ge many
EDC Pe sonal collec ion o E ic Du êne, Sain -
Hippoly e-du-Fo , F ance
KJCB Pe sonal collec ion o Kobe Janssen, Zon-
ho en, Belgium
LRC Pe sonal collec ion o F ancisco La Roche,
San C is óbal de La Laguna, Tene i e, Spain
MNCN Museo Nacional de Ciencias Na u ales, Ma-
d id, Spain
MNHN Muséum na ional d’His oi e na u elle, Pa is,
F ance
MSNG Museo Ci ico di S o ia Na u ale “Giacomo
Do ia”, Genoa, I aly
NHMD Na u al His o y Museum o Denma k, Co-
penhagen, Denma k
NHMUK Na u al His o y Museum, London, Uni ed
Kingdom
NMB Na u his o isches Museum Be n, Be n, Swi -
ze land
NHMW Na u his o isches Museum Wien, Vienna,
Aus ia
OÖLM Obe ös e eichisches Landesmuseum, Linz,
Aus ia
OUMNH Ox o d Uni e si y Museum o Na u al His o-
y, Ox o d, Uni ed Kingdom
RMNH Na u alis Biodi e si y Cen e , Leiden, he
Ne he lands
TJWC Pe sonal collec ion o Thomas J. Wood,
Leiden, he Ne he lands
VLC Pe sonal collec ion o Vincen Lecle cq, Di-
jon, F ance
WHLC Pe sonal collec ion o Wol -Ha ald Liebig,
Bad Muskau, Ge many
ZMHB Museum ü Na u kunde, Be lin, Ge many
ZSI Zoological Su ey o India, Kolka a, India
Resul s
Gene ics
Two main clades we e obse ed in he COI ba code ee
(Fig. 1). The i s had boo s ap suppo o 97 and con-
ained he mos commonly collec ed and obse ed Wes
and Cen al Palaea c ic Thy eus species, such as T. his i-
onicus (Illige , 1806) and T. amosus (Lepele ie , 1841).
The second g oup had boo s ap suppo o 87 and con-
ained T. a inis (Mo awi z, 1873), T. elegans (Mo aw-
i z, 1877) sensu la o, a newly desc ibed species Thy eus
imp essus Wood, sp. no ., and Thy eus scu ella is; his
clade is bes e e ed o as he scu ella is-g oup (sensu
Lie inck 1968). F om he RMNH collec ion, i was pos-
sible o gene a e a long 1,498 bp sequence om a speci-
men o T. scu ella is om U a in sou he n Eu opean Rus-
sia (BGENL2290-24), which was included in Lie inck’s
(1968: 46) e ision. Thy eus scu ella is was sepa a ed
om i s nea es neighbou , T. imp essus, by an a e age
gene ic dis ance o 8.05% ( ange 8.05–8.05%). The new
species T. imp essus was s ongly sepa a ed by 10.63%
( ange 10.33–10.96%) om T. elegans sensu la o and by
9.74% om T. a inis ( ange 9.57–9.88%). Fo T. elegans,
ou ba codes we e gene a ed. Specimens om Mo oc-
co, he Uni ed A ab Emi a es, and I an showed simi-
la ba codes, sepa a ed by an a e age o 1.12% ( ange
1.07–1.22%). Howe e , he specimen om sou h-eas e n
Kazakhs an (Shonzhy) was s ongly sepa a ed om hese
h ee sequences by an a e age o 10.75% ( ange 10.49–
10.96%). The issues associa ed wi h T. elegans sensu la o
a e discussed below.
Wi hin he clade o mo e commonly collec ed Thy eus
species (“O he species o Thy eus Panze ” in Lie -
inck 1968), esul s om T. pica on Lie inck, 1968 and
T. unca us (Pé ez, 1884) equi e in es iga ion. Thy eus
pica on sequences om Spain, I aly, Albania, and I an
o med a monophyle ic g oup wi h a e age in aspeci ic
a ia ion o 0.31% ( ange 0.00–0.76%), ecei ing boo -
s ap suppo o 100. This g oup di e ed om he single
T. a . pica on sequence om Ky gyzs an by an a e age
o 6.58% ( ange 6.40–6.69%), wi h he Ky gyz sequence
no alling immedia ely adjacen o he T. pica on clus e .
I is desc ibed below as Thy eus jansseni Wood, sp. no .
Fo T. unca us, he pic u e was complex. Th ee clades
we e obse ed (Fig. 1). The i s con ained specimens om
cen al Spain, which ecei ed boo s ap suppo o 97; his
can be e e ed o as he “Ibe ian lineage”. The second
con ained specimens om sou h-wes e n F ance (Aude)
and sou h-eas e n F ance (Bouches-du-Rhône), which e-
cei ed boo s ap suppo o 88; his can be e e ed o as
he “F ench lineage”. Finally, he hi d con ained speci-
mens om sou he n F ance (Ga d), Swi ze land, Ge many,
Poland, and Ky gyzs an and ecei ed boo s ap suppo o
96; his can be e e ed o as he “Eu o-Sibe ian lineage”.
Va ia ion wi hin lineages was low: an a e age o 0.21%
( ange 0.15–0.31%) in he Ibe ian lineage, 0.00% in he
F ench lineage, and 0.35% ( ange 0.00–0.83%) in he
dez.penso .ne
Thomas Wood e al.: A con ibu ion o Thy eus in he Palaea c ic, wi h wo new species262
Eu o-Sibe ian lineage, esul ing in he high obse ed boo -
s ap suppo s. Va ia ion be ween clades was 3.80% ( ange
3.50–4.16%) be ween he Ibe ian and Eu o-Sibe ian clades,
3.31% ( ange 3.12–3.48%) be ween he Ibe ian and F ench
clades, and 1.79% ( ange 1.53–2.13%) be ween he F ench
and Eu o-Sibe ian clades. All o he species in he g oup o
commonly collec ed Thy eus ecei ed boo s ap suppo
be ween 97 and 100 and we e well de ined gene ically.
Figu e 1. Phylogene ic ee (maximum likelihood) o Thy eus Panze , 1806 species, based on agmen s o he mi ochond ial COI
gene. Eupa lo skia obscu a (F iese, 1895) is used as an ou g oup. Numbe s adjacen o b anches ep esen boo s ap suppo ( al-
ues < 75% a e omi ed).
D sch. En omol. Z. 72 (2) 2025, 259–302
dez.penso .ne
263
New species desc ip ions
Thy eus jansseni Wood, sp. no .
h ps://zoobank.o g/37102F10-26D5-4F30-AAF3-64FAC7D3EC1A
Type ma e ials. Holo ype: Ky gyzs an • 1♂; Ki -
pichny; 42.4047°N, 77.8645°E; 11 Jul. 2019; K. Janssen
leg.; RMNH; RMNH.INS.1714348 (BOLD accession
numbe WPATW953-22).
Pa a ypes: a ghanis an • 2♂; Ghazni p o ince, Jag-
ho i dis ic ; 30 Jun. 1997; G.G.M. Schul en leg.; RMNH;
ZMA.INS.5144956–ZMA.INS.5144957; Ky gyzs an •
1♂; Na ynskaya, Dis . Dzhumgalsky, S Kyzyl-Oy; 1750–
1800 m a.s.l.; 5–6 Jul. 1996; H. Rausch leg.; OÖLM.
Diagnosis. Thy eus jansseni can be ecognised as a yp-
ical Thy eus (“O he species o Thy eus Panze ”) due o he
la ge la ened scu ellum which pos e io ly p ojec s o e
he me ano um and which is uni o mly la ened o e i s
en i e su ace and which is no medially sulca e (Fig. 2C),
combined wi h he geni al capsule wi h a la ge gonos y-
lus which is co e ed wi h long hai s (Fig. 3G). Due o
he whi e hai s on T1 which o m C-shaped pa ches (Fig.
3C; basally sligh ly incu ing owa ds he scu ellum), he
scu ellum wi h punc u es sepa a ed by 0.5–1 punc u e di-
ame e s and wi h polished shining in e spaces (Fig. 2C),
wi h S8 p oduced in o wo p ojec ions which a e apically
unca e and which cu e la e ally, hus appea ing o be
“gol -club”-shaped (Fig. 3E), and he geni al capsule wi h
a la ge apically unca e p ojec ion and basal do sally p o-
jec ing and acu ely poin ed sec ion o ien ed oughly 90°
ela i e o he la ge lobe (Fig. 3G), i can be con used
only wi h Thy eus pica on (Figs 2D, 3D, 3F, 3H).
Thy eus jansseni can be sepa a ed om T. pica on due
o he smalle body size o 10 mm (12–13 mm in T. pic-
a on), scu ellum 1.5 imes wide han long wi h he medi-
an no ch ba ely indica ed (Fig. 2C; in T. pica on wi h he
scu ellum 1.6 imes wide han long, and wi h he median
no ch deep and s ongly indica ed, Fig. 2D), an ennal seg-
men s iewed do sally wi h a s ong di ision be ween he
an e io ace which is co e ed wi h ine shining scales
and he dull pos e io ace (Fig. 2E; in T. pica on wi h a
weake di ision be ween he shinie an e io and dulle
pos e io aces o he an enna, Fig. 2F), pos e io aces o
he an ennal segmen s wi h e y weakly imp essed pai ed
hina ia, hose si ua ed mo e en ally e y di icul o see
(in T. pica on wi h he pos e io aces o he an ennal seg-
men s wi h s ongly imp essed pai ed hina ia, do sal and
en al hina ia equally isible), and pos e io basi a sus
wi h only a small whi e hai inge do sally (Fig. 3A; in T.
pica on wi h he hind basi a sus en i ely co e ed in ine
whi e hai s, hese becoming hicke do sally, Fig. 3B).
Desc ip ion. Female. Unknown, hough likely de-
sc ibed by Ma iko skaya (1992) as T. pica on auc o um
(see below).
Male. Body leng h: 10 mm (Fig. 2A). Head: Da k,
1.3 imes wide han long (Fig. 2B). Clypeus la ened,
densely punc a e, punc u es sepa a ed by ≤ 0.5 punc u e
diame e s, su ace be ween punc u es shining. Lab um
ounded ec angula , almos 2 imes longe han wide,
apex wi h ele a ed subapical ans e se ca ina o ming
ob usely poin ed oo h; lab um basally weakly p oduced
in o wo ube cles la e ally. Gena na owe han wid h o
compound eye; ocelloccipi al dis ance 2 imes diame e
o la e al ocellus. Hind ma gin o e ex wi h na ow ob-
scu e sligh ly aised ca ina-like im. Face be ween an en-
nal inse ions wi h aised longi udinal ca ina, educing in
heigh and becoming medial imp ession on ons an e io
o median ocellus. F ons punc a e, punc u es sepa a ed
by ≤ 0.5 punc u e diame e s, becoming spa se in a ea
an e io o median ocellus; e ex behind ocella ian-
gle densely punc a e, a eas adjacen o la e al ocelli im-
punc a e, smoo h and shining. Face wi h abundan whi e
pubescence, decumben on lowe hal o ace below an-
ennal inse ions. Gena en ally wi h da k hai s, do sal-
ly wi h sca e ed whi e hai s. An enna da k, A4–13 wi h
an e io aces ligh ened by p esence o g eyish scales;
pos e io aces o A4–13 wi h ine g anula mic o e ic-
ula ion, su ace dull and s ongly con as ing scales o
an e io aces (Fig. 2E). Pos e io aces o A4–13 wi h
small and supe icially imp essed pai ed hina ia placed
close o junc ion wi h p eceding segmen , en al hina ia
almos unde ec able. A3 0.8 imes leng h o A4.
Mesosoma: Scu um and scu ellum punc a e, punc u es
sepa a ed by 0.5–2 punc u e diame e s bu ypically by
1 punc u e diame e , su ace be ween punc u es smoo h
and shining. Scu ellum 1.5 imes wide han long, pos-
e io ma gin wa y, wi h small median no ch, pos e-
io ly wi h mode a e u o whi e hai p ojec ing om
en al su ace (Fig. 2C). Axilla lush wi h ou e ma gin
o scu ellum, scu ellum and axilla wi h da k hai s which
do no obscu e su ace. Mesepis e num densely punc a e
medially, punc u es con luen wi h sligh ly aised idg-
es, becoming spa se en ally, he e sepa a ed by 0.5–2
punc u e diame e s. Legs da k, wi h abundan whi e pu-
bescence on ou e ace o ibiae, co e ing en i e y o o e
and mid- ibiae and basal ½ o hind ibiae. Hind basi a sus
p edominan ly wi h da k hai s, do sal su ace wi h small
whi e hai inge (Fig. 3A). Fo ewing weakly in usca e.
Me asoma: Te ga da k, e gal discs punc a e wi h
hai -bea ing punc u es, punc u es p esen ing sho , pos-
e io ly p ojec ing black plumose hai s; punc u es sep-
a a ed by 0.5–1 punc u e diame e s (Fig. 3C). T1 wi h
L-shaped whi e hai pa ch la e ally, apically ex ending
u he owa ds cen e o e gum han basally, hai s p e-
dominan ly adp essed, some loose hai s ound basola e -
ally. T2 wi h almos ec angula pa ches o whi e hai s
la e ally, wi h na ow ex ension owa ds base o disc la -
e ally. T3–5 wi h ec angula pa ches o whi e adp essed
hai s, no o ming comple e bands. T7 wi h apical ma gin
essen ially s aigh , weakly wa y. S8 delica e, pos e io
p ojec ions wi h na ow s em be o e u ning 90° deg ees
o o m small la e ally p ojec ing discs (Fig. 3E). Geni-
al capsule compac , almos ounded, gonocoxa sligh ly
b oadened and unca e apically, in basal pa wi h do -
sally p ojec ing acu ely poin ed lobe o a ed 90° ela i e
o unca e sec ion (Fig. 3G). Basal and apical pa s o
gonos ylus co e ed wi h long b own hai s, hai s simple
apically, weakly plumose basally.
dez.penso .ne
Thomas Wood e al.: A con ibu ion o Thy eus in he Palaea c ic, wi h wo new species264
No es. Ma iko skaya (1992) desc ibed he emale o
“Thy eus pica on” om he Alai Moun ains in Ky gyzs an,
nea he illage o Дараут-Курган (= Da oo -Ko gon), wi h
12♀ and 7♂ specimens collec ed be ween 15 and 16 July
1986. Based on Ma iko skaya’s w i ing and illus a ions,
i is highly likely ha she was dealing wi h T. jansseni and
hence desc ibed he emale o his species. Un o una ely,
Ma iko skaya’s collec ion appea s o ha e been des oyed
(Pie e Rasmon , pe s. comm.), which does no allow us o
d aw conclusions wi h ce ain y. Mo eo e , Ma iko skaya
did no gi e he size o he emale o male specimens, bu
she did indica e ha he hos o he species was An hopho-
a (Mys acan hopho a) bo ealis Mo awi z, 1865. This spe-
cies, and indeed he subgenus o bees, is e y a e in Eu-
ope and is ound in empe a e o bo eal habi a s, which do
no o e lap a all wi h he obse ed dis ibu ion o T. pic-
a on in Eu ope and he Nea Eas , which is Medi e anean
in cha ac e . An hopho a bo ealis is ypically 10–11 mm in
leng h, which is simply oo small o hos T. pica on, which
has males 12–13 mm in leng h, and emales will o cou se
Figu e 2. Thy eus jansseni Wood, sp. no . male holo ype (RMNH). A. Habi us, la e al iew; B. Face, on al iew; C. Scu ellum, do -
sal iew; E. An enna, do sal iew. Thy eus pica on Lie inck, 1968 male (TJWC); D. Scu ellum, do sal iew; F. An enna, do sal iew.
D sch. En omol. Z. 72 (2) 2025, 259–302
dez.penso .ne
265
Figu e 3. Thy eus jansseni Wood, sp. no . male holo ype (RMNH). A. Hind basi a sus, la e al iew; C. Me asoma, do sal iew;
E. S8, do sal iew; G. Geni al capsule, do sal iew; Thy eus pica on Lie inck, 1968 male (TJWC); B. Hind basi a sus, la e al iew;
D. Me asoma, do sal iew; F. S8, en ola e al iew; H. Geni al capsule, do sal iew.
dez.penso .ne
Thomas Wood e al.: A con ibu ion o Thy eus in he Palaea c ic, wi h wo new species266
be sligh ly la ge on a e age. Thy eus pica on is likely
a acking la ge Amegilla (Amegilla) species (see below),
which would accoun o i s much la ge body size ela i e
o T. jansseni. An hopho a bo ealis and, possibly o he spe-
cies o Mys acan hopho a B ooks, 1988, he e o e seem o
be he likely hos (s) o T. jansseni.
E ymology. The name is o ecognise Kobe Janssen
(Belgium), who e y gene ously sha ed la ge numbe s o
bee specimens (including Melec ini) wi h he lead au ho
o a long pe iod o ime.
Dis ibu ion. Ky gyzs an and A ghanis an. Likely
p esen in o he Cen al Asian coun ies, gi en Lie inck’s
(1968: 92) pa a ypes o T. pica on om Uzbekis an (Dz-
huma) and Kazakhs an (Zha ken , Kazaly) (specimens
in S . Pe e sbu g, Ox o d, and P ague). Examina ion o
hese specimens is necessa y o clea ly delinea e he dis-
ibu ions o hese wo species. Sea ches in he OUMNH
(TJW, Augus 2025) could no loca e any “T. pica on”
specimens o egis a ion o such in he ype ca alogue.
Thy eus imp essus Wood, sp. no .
h ps://zoobank.o g8E37B113-D55C-4FB7-8874-2A9D628367DE
Type ma e ials. Holo ype: Ky gyzs an • 1♂; 45 km W
o Na yn; 41.4160°N, 75.3969°E; 20 Jul. 2019; K. Jans-
sen leg.; RMNH; RMNH.INS.1714347 (BOLD acces-
sion numbe WPATW950-22).
Pa a ype: Ky gyzs an • 1♂; 45 km W o Na yn; 20
Jul. 2019; 41.4160°N, 75.3969°E; K. Janssen leg.; TJWC
(BOLD accession numbe WPATW951-22).
Diagnosis. Thy eus imp essus can be placed in o he
scu ella is-g oup due o he geni al capsule, which is
simple, wi h he gonos ylus g ea ly educed in size and
poin ed- iangula wi hou an addi ional do sal p ojec ion
(Fig. 5E), and he gonos ylus wi h only a ew inconspic-
uous hai s. I can be sepa a ed om all o he membe s
o he g oup excep T. scu ella is by he s uc u e o he
scu ellum, which is medially imp essed and sulca e, wi h
he axillae sligh ly di e ging om he ou e ma gins o
he scu ellum and p oduced in o sligh bu dis inc poin s
(Fig. 4C), and due o he whi e hai pa ches a he base
and on he ma ginal a ea o T1, bo h widely sepa a ed
medially and la e ally by abundan black hai s, hus o m-
ing 4 dis inc pa ches in do sal iew (Fig. 4D).
The wo species a e simila , bu T. imp essus can
be ecognised due o he gonos ylus which is small
and acu ely poin ed, wi h he inne ma gin s aigh
(Fig. 5E; in T. scu ella is wi h he gonos ylus mo e
elonga e and apically blun o ounded, and he inne
ma gin wi h a dis inc subapical kink, Fig. 5F; see also
illus a ion in Lie inck 1968: 46), in on al iew wi h
A3 only sligh ly exceeding A4 in leng h (Fig. 5A; in
T. scu ella is wi h A3 clea ly exceeding A4 in leng h,
app oaching A4+5, Fig. 5B), and he pos e io ac-
es o he an ennal segmen s wi h s ongly p oduced
Figu e 4. Thy eus imp essus Wood, sp. no . male holo ype (RMNH). A. Habi us, la e al iew; B. Face, on al iew; C. Scu ellum,
do sal iew; D. Me asoma, do sal iew.
D sch. En omol. Z. 72 (2) 2025, 259–302
dez.penso .ne
267
indi idual hina ia (no pai ed), clea ly isible on he
pos e io ace o A13 (Fig. 5C; in T. scu ella is wi h he
pos e io aces o he an ennal segmen s wi h weakly
p oduced indi idual hina ia, ha on A13 so small and
weak as o no be immedia ely isible, Fig. 5D).
Desc ip ion. Female. Unknown.
Male. Body leng h: 9.5–10 mm (Fig. 4A). Head: Da k,
1.25 imes wide han long (Fig. 4B). Clypeus la ened,
densely punc a e, punc u es con luen , su ace dull wi h
excep ion o na ow shining ma gin. Lab um ounded
ec angula , 1.5 imes longe han wide, apex wi h ele a -
ed subapical ans e se ca ina o ming ob usely poin ed
oo h; lab um basally weakly p oduced in o wo ube cles
la e ally. Gena na owe han wid h o compound eye;
ocelloccipi al dis ance 1.5 imes diame e o la e al ocel-
lus. Hind ma gin o e ex wi h na ow obscu e sligh ly
aised ca ina-like im. Face be ween an ennal inse ions
wi h weak aised longi udinal ca ina, apidly educing in
heigh and becoming medial imp ession on ons an e io
o median ocellus. F ons punc a e, punc u es sepa a ed
by ≤ 0.5 punc u e diame e s, punc a ion becoming sligh -
ly weake on e ex, he e sepa a ed by up o 1 punc u e
diame e . Face wi h abundan whi e pubescence, decum-
ben on lowe hal o ace below an ennal inse ions.
Figu e 5. Thy eus imp essus Wood, sp. no . male holo ype (RMNH). A. A3–5, on al iew; C. A9–13, pos e io iew; E. Geni al
capsule, de ail o gonos ylus, do sola e al iew; Thy eus scu ella is (Fab icius, 1781) male (TJWC); B. A3–5, on al iew; D. A9–
13, pos e io iew; F. Geni al capsule, de ail o gonos ylus, do sal iew.
dez.penso .ne
Thomas Wood e al.: A con ibu ion o Thy eus in he Palaea c ic, wi h wo new species274
The p esence o whi e hai pa ches on he axillae
gi es T. pa henope a simila appea ance o T. hyalina-
us (Vachal, 1903) (Is ael, Egyp , Sudan, Chad*, E i ea,
Djibou i, Saudi A abia, Qa a *, Uni ed A ab Emi a es*,
Oman*, I an; Lie inck 1968), bu he wo species can be
sepa a ed by he p esence o he whi e hai spo s on he
scu ellum o T. pa henope (absen in T. hyalina us). The
wo species can also be sepa a ed due o he s uc u e o
he mesepis e num in i s en al hal ( he pa no co e ed
wi h pubescence), which has he punc u es dense and e y
weakly sepa a ed in T. pa henope (punc u es almos con-
luen ), whe eas T. hyalina us shows clea shining spaces
be ween he punc u es, hese spaces o en wide han he
diame e o a punc u e. These cha ac e s, in combina-
ion wi h he dis ibu ion, which is cu en ly es ic ed o
he sou he n hal o he A abian Peninsula and p obably
sou h-eas e n Egyp allows ecogni ion, pa icula ly in
combina ion wi h concu en ly ac i e males.
Desc ip ion. Female. Body leng h: 12 mm (Fig. 11A).
Head: Da k, 1.3 imes wide han long (Fig. 11B). Cly-
peus e y weakly domed, densely punc a e, punc u es
sepa a ed by ≤ 0.5 punc u e diame e s, in e spaces shin-
ing. Lab um ounded ec angula , 1.5 imes longe han
b oad, wi h longi udinal imp essed midline e mina ing
subapically, apex wi h iny sligh ly p ojec ing oo h; la-
b um basally s ongly p oduced in o wo ube cles la e -
ally, ou e hal o ube cules polished and shining, im-
punc a e. Gena much na owe han wid h o compound
eye; ocelloccipi al dis ance 2 imes diame e o la e al
ocellus. Face be ween an ennal inse ions wi h weakly
aised longi udinal ca ina, no s ongly ex ending do sal-
ly. F ons densely punc a e, punc u es sepa a ed by ≤ 0.5
punc u e diame e s, becoming weake la e ally adjacen
o la e al ocelli, he e wi h polished shining impunc a e
space subequal o diame e o la e al ocellus (Fig. 11C).
Head almos en i ely whi e-hai ed, wi h b ownish hai s
es ic ed o mandibles. An enna da k, measu ed along
en al su ace A3, mo e o less equalling A4.
Mesosoma: Scu um and scu ellum densely punc a e,
punc u es sepa a ed by ≤ 0.5 punc u e diame e s, up o 1
punc u e diame e on scu ellum, su ace be ween punc-
u es shiny. Scu um wi h con as ing black and whi e
pubescence, whi e pubescence along an e io ma gin,
la e al ma gins, medially wi h longi udinal line co e -
ing an e io ¾ o scu um, an e ola e ally wi h wo whi e
spo s, pos e ola e ally wi h wo hick pa ches along pos-
e io ma gin (Fig. 11D). Axilla wi h clea pa ch o whi e
plumose adp essed hai s. Scu ellum pos e o-la e ally wi h
wo pa ches o whi e plumose adp essed hai s. Scu ellum
p oduced in o wo pos e io ly p ojec ing poin s, be ween
hese wi h shallow ema gina ion, scu ellum i sel ela i e-
ly long, maximum wid h 1.5 imes longe han maximum
leng h; ema gina ion be ween pos e io poin s wi h hick
b oad u o whi e hai s eme ging om en al su ace
(Fig. 11E). Mesepis e num in do sal ½ co e ed wi h dense
pa ch o adp essed whi e hai s, en i ely obscu ing su ace;
in en al ½ wi h su ace lacking hai s, densely and egu-
la ly punc a e, punc u es almos con luen . Legs da k, ou -
e su ace o ibiae and a si co e ed wi h dense el -like
whi e hai s, mid and hind ibiae wi h sho da k spines p o-
jec ing h ough pubescence. Fo ewing almos en i ely hy-
aline, wi h only weak in usca ion adjacen o a ew eins.
Me asoma: Te ga da k, e gal discs densely punc-
a e wi h hai -bea ing punc u es, punc u es p esen ing
Figu e 10. Dis ibu ion o Thy eus unca us (Pé ez, 1884) ( ed ci cles) along wi h i s hos Te alonia den a a (Ge ma , 1839) (open
blue iangles). Dis ibu ional da a a e no comp ehensi e.
25N
30N
35N
40N
45N
50N
55N
60N
65N
0 20E 40E 60E 80E
Longi ude
La i ude
D sch. En omol. Z. 72 (2) 2025, 259–302
dez.penso .ne
275
sho , pos e io ly p ojec ing black plumose hai s,
punc u es sepa a ed by 0.5–2 punc u e diame e s (Fig.
11F). T1 la e ally wi h wide C-shaped pa ch o whi e
hai s, hai s along apical ma gin o e gum eaching
u he owa ds cen e o disc han hai s along basal
ma gin o e gum. T2 wi h L-shaped pa ch o whi e
hai s, T3–5 wi h ec angula whi e hai pa ches, hai
pa ches on all e ga b oadly sepa a ed medially, ne e
comple e. Pygidial pla e weakly con e ging apically,
la e al ma gins sligh ly aised, medially wi h s ongly
humped longi udinal bump, su ace dull.
Dis ibu ion. Egyp (p esumably sou he n Egyp ),
Sudan*, Saudi A abia*, Yemen, Uni ed A ab Emi a es*,
Oman* (Lie inck 1968).
O he ma e ial examined. (Thy eus hyalina us) chad
• 1♀; Tchad, Ndjamena; 30 Ma . 1978; G.G.M. Schul en
leg.; RMNH • 1♀; Tchad, Ndjamena; 22 Jun. – 6 Jul. 1978;
G.G.M. Schul en leg.; RMNH; oMan • 2♀; 20 km W Ba -
ka, Wadi Fa ; 14 Ma . 2022; C. Schmid-Egge leg.; CSE
• 6♂, 1♀; Dhau u , Thum ai 35 km N, KF dese a m;
21 Sep. 2023; C. Schmid-Egge & W-H. Liebig leg.; CSE/
TJWC/WHLC • 1♂; Dho a , 2 km W Taqah; 29 Aug. 2019;
Figu e 11. Thy eus pa henope Lie inck, 1968, emale. A. Habi us, la e al iew; B. Head, on al iew; C. Ve ex, do sal iew;
D. Scu um, do sal iew; E. Scu ellum, do sal iew; F. Me asoma, do sal iew.
dez.penso .ne
Thomas Wood e al.: A con ibu ion o Thy eus in he Palaea c ic, wi h wo new species276
C. Schmid-Egge leg.; CSE • 1♀; Dho a , 35 km N Thum-
ai , Najd; 3 Sep. 2019; C. Schmid-Egge leg.; CSE • 1♀;
SW o Su ; 3–5 Ma . 2017; M. Snižek leg.; OÖLM; Qa a
• 1♂; Al Shahaniyeh [Ash-Shahaniyah]; 9 May 1980; C.G.
Roche leg.; D.B. Bake de . 1993; OUMNH • 1♂; Al Sha-
haniyeh [Ash-Shahaniyah]; 23 May 1980; C.G. Roche
leg.; D.B. Bake de . 1993; OUMNH • 5♂; Al Shahaniyeh
[Ash-Shahaniyah]; 13 Jun. 1980; C.G. Roche leg.; D.B.
Bake de . 1993; OUMNH; Uni ed a ab eMi a es • 1♀;
Wadi Bih, dam; 29 No . 2009; A. an Ha en leg.; CSE.
Thy eus pica on Lie inck, 1968
Thy eus pica on Lie inck, 1968: 92, ♂ [C oa ia, RMNH, examined].
Ma e ial examined. albania • 1♂; Topo ë; 13 Jul. 2017;
K. Janssen leg.; KJCB; bUlga ia • 1♂; Melnik; 4 Jul.
2019; M. Halada leg.; OÖLM; c oa ia • 1♂; S ob eč (n .
Spli ); 13 Jun. 1962; C. . Heijningen leg.; M.A. Lie inck
de .; RMNH; RMNH.INS.1714262 (holo ype) • 1♂; Baš-
ka, Insel K k; 16 Aug. 1976; J. Hein ich leg.; OÖLM; cy-
p Us • 1♂; 20 km NNW Pa os, La a Beach; 20 Jun. 2013;
C. Schmid-Egge leg.; CSE • 1♂; Ak o i i; 26 Ap . 2018;
A. Va na a leg.; TJWC; ance • 1♂; Alpes Ma i imes,
1 km S Fon an; 13 Jul. 2009; C. Schmid-Egge leg.; CSE
• 1♂; Va , Le T ayas; O.W. Richa ds leg.; M.A. Lie -
inck de .; NHMUK; geo gia • 1♂; Tbilissi, Bo anical
Ga den; 26 Jul. 2021; V. Lecle cq leg.; E. Du êne de .,
VLC; g eece • 1♂; Chalkidiki, W o Niki i; 12–14 Jun.
2013; M. Snižek leg.; OÖLM • 1♂; Ka penission, 2000
[Ka penisi]; 20 Jul. 1976; K.M. Guicha d leg.; M.A.
Lie inck de .; NHMUK • 1♂; P e eza en .; 25 Jun. 1997;
K. Deneš leg.; OÖLM; i an • 1♂; Goles an, Kashida ;
1381 m a.s.l.; 9 Jul. 2018; W-H. Liebig leg.; WHLC •
1♂; Semnan p o ince, Shahmi zad, 13 km NW; 1836 m
a.s.l.; 7 Jul. 2022; C. Schmid-Egge leg.; CSE • 1♂; Te-
he an p o ince, Elbu z, Se kheh [So kheh Zamin]; 2266
m a.s.l.; 6 Jul. 2022; C. Schmid-Egge leg.; CSE • 1♀;
Teh an p o ., Elbu z, Fi uzkuh 1 km SW; 6 Jul. 2022;
C. Schmid-Egge leg.; CSE; i aly • 1♂; S-Ti ol, Me an,
Schloss-T au mansdo ; 21 Aug. 2013; T. Kop leg.; CSE
• 1♂; S-Ti ol, Me an, Schloss-T au mansdo ; 11 Aug.
2013; T. Kop leg.; CSE; lebanon • 1♂; No d Libanon,
Becha e [Bsha i]; 1400 m a.s.l.; 1–4 Jul. 1931; Ze ny
leg.; M.A. Lie inck de .; RMNH; RMNH.INS.1714263;
Mon eneg o • 1♂; Ulcinj; 29 Jun. 1969; Ho e leg.;
OÖLM; no h Macedonia • 1♂; Su odol, 18 km ENE
o Bi ola; 750–1000 m a.s.l.; 30 Jul. 1965; M.A. Lie -
inck de .; RMNH; RMNH.INS.1714264; oMania • 1♂;
Dob ogea [ emaining ex illegible]; 7 Jul. 1994; OÖLM;
slo aKia • 1♂; S u o o [Š ú o o]; 28 Jul. 1972; M. Ko-
cou ek leg.; OÖLM; spain • 1♂; Ca alonia, Palamos; 19
Jul. 1959; S.G. Bisho leg.; M.A. Lie inck de .; ZMHB
• 1♂; N. o Mad id, Rio Guada ama; 27 Jul. 1980; K.M.
Guicha d leg.; NHMUK • 1♂; Sego ia, B ie a, 5 km N,
Las Cañones de los ios Pi on y Viego; 16 Jul. 2021; T.J.
Wood leg.; TJWC • 1♂; Valencia; [unda ed]; Mo ode
leg.; MNCN • 1♂; Pon e ed a, Co ellas; [unda ed]; Va e-
la leg.; MNCN; U Key • 1♂; Bi lis, Nem u Dağı; 2300
m a.s.l.; 15 Aug. 1991; J. Halada leg.; OÖLM; UK aine
• 1♂; Odessa [Odesa] Oblas , K asnosilka en .; 23 Jun.
2024; A.V. Gon a enko leg.; E. Du êne de ., VLC.
Rema ks. I was possible o iden i y he emale o
T. pica on h ough DNA ba coding. Un o una ely, he
loca ed emale om I an (WPATW1024-23) appea s o
be comple ely iden ical o T. his ionicus (Panze , 1806).
I he e o e does no cu en ly seem possible o p o ide
a diagnosis o he emale sex, and so u u e eco ds o
T. his ionicus om wi hin he expanded dis ibu ional
ange ( ela i e o Lie inck 1968) o T. pica on mus be
conside ed in a b oad sense. Examina ion o addi ional
ma e ial has inc eased he dis ibu ional ange o T. pic-
a on, pa icula ly in sou h-eas e n Eu ope (Fig. 12). As
sugges ed by Gaspa e al. (2025), he hos is likely o be
one o bo h o wo o he la ge Amegilla sensu s ic o,
ei he Am. (Amegilla) ga ula (Rossi, 1790) o Am. (Ame-
gilla) och oleuca (Pé ez, 1879). Like hese wo possible
Figu e 12. Dis ibu ion o Thy eus pica on Lie inck, 1968 ( ed ci cles) along wi h i s wo suspec ed hos s, Amegilla ga ula (Rossi,
1790) (open blue iangles) and Amegilla och oleuca (Pé ez, 1879) ( illed yellow squa es). Dis ibu ional da a a e no comp ehensi e.
30N
35N
40N
45N
50N
10W 010E20E30E40E50
E6
0
E
Longi ude
La i ude
D sch. En omol. Z. 72 (2) 2025, 259–302
dez.penso .ne
277
hos s, T. pica on does no ex end in o No h A ica. Re-
co ds o Am. ga ula and Am. och oleuca a e p esen ed
in Suppl. ma e ial 1. The use o wo hos s is suspec ed
due o a eas whe e only one possible hos is known (wi h
a high deg ee o con idence) o be p esen , such as in
cen al Spain and no he n Po ugal (Am. ga ula absen )
and no he nmos I aly (Am. och oleuca absen ). A he
I alian si es su eyed in 2013, T. pica on was lying wi h
Am. ga ula, suppo ing his associa ion. Thy eus pica on
has also been egula ly collec ed wi h Am. ga ula in he
sou h o F ance (M. Aube , pe s. comm.).
Based on his hypo hesis, T. pica on should be com-
mon in Tu key; a s udy o addi ional Tu kish Thy eus
specimens o cla i y i s ange in his coun y is necessa y.
I is impo an o no e ha he e is a c yp ic species close
o Am. och oleuca which is cu en ly being desc ibed
om Lebanon, eas e n Tu key (Ağ ı, Hakkâ i), no h-
e n I an, and sou h-wes e n Tu kmenis an (P. Rasmon
& TJW, in p epa a ion); i is e y a e in collec ions and
is collec ed much mo e a ely han Am. och oleuca in
Tu key. I may se e as a hi d hos species o T. pic-
a on. Fo simplici y, hese eco ds a e he e olded in o
Am. och oleuca. In no he n I an, T. pica on, he c yp ic
“Am. och oleuca”, and Am. ga ula ha e all been caugh
in close p oximi y (Fig. 12). We epo T. pica on as new
o he auna o Uk aine, om whe e Am. och oleuca
has also been eco ded (see Suppl. ma e ial 1), and his
species is also known om he Odesa egion based on
specimens held in he Kansas collec ion and de e mined
by B ooks (KSEM1323579; Asche and Picke ing 2025),
p o iding a plausible hos . We also epo T. pica on as
new o Geo gia, a a collec ing locali y whe e se e al
Am. ga ula we e also cap u ed (see Suppl. ma e ial 1).
Dis ibu ion. Po ugal, Spain, F ance, I aly, Aus ia,
Slo akia*, Slo enia, C oa ia, Se bia, Romania, Uk aine*,
Mon eneg o*, Albania*, No h Macedonia, Bulga -
ia, G eece, Cyp us, Tu key, Geo gia*, Lebanon*, I an
(Lie inck 1968 pa im; Kuhlmann e al. 2014; Va na a e
al. 2020; Asche and Picke ing 2025; Gaspa e al. 2025;
Fig. 12). As discussed abo e, we suspec ha eco ds
om Cen al Asia will co espond o T. jansseni.
Thy eus p iesne i Lie inck, 1968
Thy eus p iesne i Lie inck, 1968: 124, ♂ [Egyp , ype loca ion unclea ].
Ma e ial examined. saUdi a abia • 1♂, 1♀; Abu A -
ish; 25 Ma . 1980; K.M. Guicha d leg.; D.B. Bake de .
1981; NHMUK.
No es. Lie inck (1968) desc ibed T. p iesne i om
a single male specimen collec ed om sou he n Egyp
(Gebel Elba). This specimen was ci ed as being in he
collec ion o He mann P iesne a “Linz”, bu i could no
be loca ed he e, and i s cu en deposi ion is he e o e
no immedia ely clea ; u he s udy is equi ed o ace
i . Fo una ely, he desc ip ion is well illus a ed, allow-
ing con iden ecogni ion. In he NHMUK collec ion,
wo specimens om Saudi A abia we e ound; as wi h
T. pa henope, hese specimens we e iden i ied by Bake
bu we e ne e published. The male clea ly ma ches he
c i e ia iden i ied by Lie inck, and due o co-occu ence
and close mo phology, he emale can now be ecognised
(Fig. 13). The emale can be diagnosed based on a com-
bina ion o cha ac e s, bu ca e mus be aken, as hey a e
qui e sub le. The clea es con usion is wi h Thy eus al-
libilis (Kohl, 1905), which was desc ibed om sou he n
Yemen (Fig. 14). Lie inck (1968) diagnosed his species
and desc ibed he unknown male; due o he simila i y o
he emales, his species is diagnosed las .
Diagnosis. As in T. pa henope, T. p iesne i has an
unb oken line o whi e hai s unning along he en i e
leng h o he la e al ma gins o he scu um (Fig. 13E), as
in T. amosus (Lepele ie , 1841) and T. amosellus Cock-
e ell, 1919. Thy eus pa henope is diagnosed abo e, dis-
playing whi e hai pa ches on he axillae and scu ellum,
whe eas hese a eas a e black-hai ed in T. p iesne i (Fig.
13E), which also shows he lowe hal o he mesepis e -
num wi h la ge shining in e spaces (Fig. 13D; wi h some
in e spaces eaching 3 punc u e diame e s), whe eas in
T. pa henope his a ea is densely punc a e, wi hou la ge
shining in e spaces. This shining and spa sely punc a e
mesepis e num also allows sepa a ion om T. amosus
and T. amosellus, and indeed i is s ange ha Lie inck
diagnosed T. p iesne i agains T. amosus when in he
cons uc ion o he geni al capsule and he shining low-
e hal o he mesepis e num i is much close o T. hy-
alina us (Vachal, 1903), wi h which i has an o e lap-
ping ange in Egyp and he A abian Peninsula. Thy eus
p iesne i can be di e en ia ed om he la e species due
o he axillae wi h black hai s (Fig. 13E; in T. hyalina-
us wi h whi e pa ches o hai s co e ing he axillae), he
L-shaped pa ches o hai s on he la e al pa s o T1 (Fig.
13F), hese being une en, wi h he apical pa p ojec ing
u he owa ds he cen e o he e gal disc han he basal
pa (in T. hyalina us wi h he C-shaped pa ches o hai s
mo e e en, wi h bo h he basal and apical pa s uni o mly
p ojec ing owa ds he cen e o he e gal disc), and he
pos e io ma gin o he scu ellum, which has pos e io ly
p ojec ing whi e hai s eme ging om below bu no si -
ing on he do sal side o he disc (Fig. 13E; in T. hyali-
na us wi h hai s eme ging bo h om below he pos e io
ma gin o he scu ellum and p esen on he do sal side
o he pos e io ma gin o he disc). This combina ion o
cha ac e s (con inuous line o whi e hai s on he la e al
ma gins o he scu um, lowe hal o he mesepis e num
spa sely punc a e wi h la ge shining in e spaces, and ax-
illae and scu ellum en i ely black-hai ed) should allow
ecogni ion o emales o T. p iesne i compa ed o o he
species in no h-eas e n A ica and he A abian Peninsula.
Fo diagnosis agains T. allibilis, he o e all pubes-
cence pa e n is highly simila , as well as he punc a ion
and mic osculp u e o he in egumen , such as in he
lowe pa o he mesepis e num wi h la ge shining in-
e spaces (Fig. 14D), he e ex domed in on al iew
(Fig. 14C), and he scu ellum o simila shape and wi h
dez.penso .ne
Thomas Wood e al.: A con ibu ion o Thy eus in he Palaea c ic, wi h wo new species278
simila punc u e densi y (Fig. 14E). The key di e ence is
in he s uc u e o he indi idual hai s; in T. allibilis, he
hai s a e la ened and s ongly plumose, wi h he b anch-
es sp eading la e ally, becoming almos scale-like (Fig.
14E, F). In T. p iesne i, he hai s a e plumose bu a e no
la ened, and he b anches do no sp ead la e ally, mean-
ing ha hey ha e a mo e quill-like appea ance (Fig. 13E,
F). This can be seen mos clea ly on he disc o T1. As no -
ed by Lie inck (1968: 96), he pubescence o T. allibilis
is also sligh ly in ed wi h blue, whe eas in T. p iesne i
he pubescence is pu e whi e. We main ain he species
concep s p esen ed by Lie inck, bu his pu a i e species
pai would bene i om molecula in es iga ion.
Desc ip ion. Female. Body leng h: 9 mm (Fig. 13A).
Head: Da k, 1.2 imes wide han long (Fig. 13B). Cly-
peus sligh ly ele a ed, mo e o less la ened ac oss disc,
densely punc a e, punc u es sepa a ed by ≤ 0.5 punc u e
diame e s, in e spaces shining. Lab um b oadly ounded
ec angula , la e al ma gins con e ging, apex hus na -
owe han base, lab um only sligh ly longe han basal
wid h. Lab um basally wi h wo sligh ly aised ube -
cules, medially wi h imp essed u ow, his e mina ing
Figu e 13. Thy eus p iesne i Lie inck, 1968, emale. A. Habi us, la e al iew; B. Head, on al iew; C. Lab um, en al iew;
D. Mesepis e num, en ola e al iew; E. Scu um, do sal iew; F. Me asoma, do sal iew.
D sch. En omol. Z. 72 (2) 2025, 259–302
dez.penso .ne
279
subapically a s ong ans e se ca inae, medially p o-
duced in o poin ed oo h (Fig. 13C). Gena much na owe
han wid h o compound eye, pos e io ly wi h s ongly
aised winged ca inae, na owing o be almos impe cep-
ible along pos e io ma gin o e ex; ocelloccipi al dis-
ance 1.5 imes diame e o la e al ocellus. Face be ween
an ennal inse ions wi h weakly aised longi udinal ca i-
na, no s ongly ex ending do sally. F ons densely punc-
a e, punc u es sepa a ed by 0.5–1 punc u e diame e s,
wi h sligh impunc a e space adjacen o la e al ocelli,
space subequal o diame e o la e al ocellus. Head almos
en i ely whi e-hai ed, wi h b ownish hai s es ic ed o
mandibles. An enna da k, measu ed along en al su ace
A3, sligh ly bu dis inc ly sho e han A4.
Mesosoma: Scu um and scu ellum densely punc a e,
punc u es somewha i egula , sepa a ed by < 0.5–1 punc-
u e diame e s, punc u es mos consis en ly dense on scu e-
llum, su ace shiny. Scu um wi h con as ing black and
whi e pubescence, abundan whi e pubescence along an-
e io ma gin, la e al ma gins, medially wi h longi udinal
line co e ing an e io ½ o segmen , an e io-la e ally wi h
wo whi e spo s, pos e o-la e ally wi h wo hick pa ches
Figu e 14. Thy eus allibilis (Kohl, 1905) holo ype emale. A. Label de ails; B. Habi us, la e al iew; C. Face, on al iew;
D. Mesepis e num, la e al iew; E. Scu um, do sola e al iew; F. Me asoma, do sal iew.
dez.penso .ne
Thomas Wood e al.: A con ibu ion o Thy eus in he Palaea c ic, wi h wo new species280
along pos e io ma gin (Fig. 13E). Axillae and scu el-
lum en i ely black hai ed on do sal su ace. Scu ellum
p oduced in o wo pos e io ly p ojec ing poin s, be ween
hese wi h shallow ema gina ion, scu ellum i sel ela i e-
ly long, maximum wid h 1.5 imes longe han maximum
leng h (measu ed om base o midpoin be ween pos e io
poin s); ema gina ion be ween pos e io poin s wi h hick
b oad u o whi e hai s eme ging om en al su ace.
Mesepis e num in do sal ½ co e ed wi h dense pa ch
o adp essed whi e hai s, en i ely obscu ing su ace; in
en al ½ wi h su ace lacking hai s, wi h spli punc a-
ion; in an e io po ion wi h su ace abundan ly punc a e,
punc u es sepa a ed by < 0.5–1, in pos e io po ion wi h
punc u es becoming e y spa se, punc u es sepa a ed by
1–4 punc u e diame e s, in e spaces smoo h and polished
(Fig. 13D). Legs da k, ou e su ace o ibiae co e ed
wi h dense el -like whi e hai s, mid and hind ibiae wi h
sho da k spines p ojec ing h ough pubescence. Fo e-
wing wi h a ied colou a ion, p edominan ly hyaline
wi h in usca ion adjacen o ein, wi hin subma ginal and
ma ginal cells, and on apical papilla e egion.
Me asoma: Te ga da k, e gal discs densely punc a e
wi h hai -bea ing punc u es, punc u es p esen ing sho ,
pos e io ly p ojec ing black plumose hai s, punc u es
sepa a ed by 0.5–2 punc u e diame e s, ex ending on o
majo i y o ma ginal a eas, wi h ma ginal a eas na owly
impunc a e along apical im (Fig. 13F). T1 la e ally wi h
la ge pa ch o whi e hai s, hese no ypically C-shaped,
mo e L-shaped wi h apical pa p ojec ing u he owa ds
cen e o disc han basal pa . T2 wi h b oad L-shaped
pa ch o whi e hai s, T3–5 wi h ec angula whi e hai
pa ches, hai pa ches on all e ga b oadly sepa a ed me-
dially, ne e comple e. Pygidial pla e long, iangula ,
na owly ounded, la e al ma gins sligh ly aised, su ace
la , i egula ly punc a e wi h punc u es o a iable sizes,
su ace e y obscu ely shining.
Dis ibu ion. Sou he n Egyp and Saudi A abia*
(Lie inck 1968).
O he ma e ial examined. (Thy eus allibilis (Kohl,
1905)) yeMen • 1♀; Makalla [Al Mukalla]; 1–30 Ap .
1899; O. Simony leg.; NHMW (holo ype).
New dis ibu ional da a and ange cla i ica ions
Thy eus hellenicus Lie inck, 1968
Thy eus hellenicus Lie inck, 1968: 71, ♀♂ [G eece, RMNH, exam-
ined] (Fig. 15A–D).
Ma e ial examined. g eece • 1♂; Pa nass[us]; 19 Jul.
1956; By inski-Salz leg.; M.A. Lie inck de .; RMNH;
RMNH.INS.1714335 (holo ype) • 1♂; Pa nass[us];
19 Jul. 1956; By inski-Salz leg.; M.A. Lie inck de .;
RMNH; RMNH.INS.1662710 (pa a ype) • 1♀; G ae-
cia, Peloponnesus, Al -Ko in h; 3 Jun. 1964; M. Schwa z
leg.; M.A. Lie inck de .; RMNH; RMNH.INS.1662711
(pa a ype) • 1♀; I. Kalymnos, Po hia; 2–4 Jun. 1935;
O. We s ein; NHMW • 1♂; Leg ena; 1 Jun. 1965; K.V.
K ombein leg.; M.A. Lie inck de .; RMNH; RMNH.
INS.1662712 (pa a ype) • 3♀; Amo gos, Agioi Sa an-
a; 1 m a.s.l.; 19 Jun. 2023; V. Lecle cq leg.; V. Lecle cq
de .; VLC/EDC • 1♂; Gy hion [Gy heio]; 3 Jun. 1937;
F. We ne leg.; NHMW (pa a ype); KazaKhs an • 2♂;
U alsk; M. Ba el leg.; NHMW (pa a ypes); spain • 1♂;
Mad id [no u he in o ma ion]; G. Me ce collec ion;
MNCN; MNCN_En 436609; (pa a ype); sy ia • 1♂; S.
Sy ia: Moun He mon; 30 Jul. 1945; G.H. Q. & M.E.F.
leg.; M.A. Lie inck de .; NHMUK (pa a ype); U Key •
1♂; Canki i, 15 kms Ilgaz-Canki i Rd.; 1400 m a.s.l.; 23
Jul. 1962; Guicha d & Ha ey leg.; M.A. Lie inck de .;
NHMUK (pa a ype) • 1♂; Canki i, 15 kms Ilgaz-Canki-
i Rd.; 1400 m a.s.l.; 23 Jul. 1962; Guicha d & Ha ey
leg.; M.A. Lie inck de .; RMNH; RMNH.INS.1714265
(pa a ype) • 1♂, 1♀; Konya, Sul an Dağla ı, 10 km S o
Çay; 1300 m a.s.l.; 18–25 Jul. 1980; H. . Oo scho leg.;
RMNH; ZMA.INS.5144951.
No es. Thy eus hellenicus is a poo ly known species
mos commonly encoun e ed in he Eas Medi e anean.
Lie inck (1968: 76) imp ecisely s a ed “ he Whole Med-
i e anean egion” when discussing he dis ibu ion, de-
spi e p esen ing specimens om only Spain, G eece, Tu -
key, Sy ia, and no h-wes e n Kazakhs an ( he sou he n
ip o he U al Moun ains). Discussion o he ange o his
species combined wi h i s hos is necessa y.
Thy eus hellenicus was desc ibed om Moun Pa nas-
sus in sou he n G eece, and mos specimen eco ds p e-
sen ed by Lie inck we e om his coun y. Specimens
om G eece, Tu key, and Sy ia (Moun He mon) a e
mo phologically consis en and ecognisable (see cha -
ac e s in he iden i ica ion key below). I was possible
o examine he pa a ype om Spain (MNCN) om he
collec ion o Rica do Ga cía Me ce (Fig. 16). I has been
dissec ed and is indeed T. hellenicus based on he size
and geni al capsule (Fig. 16D), as indica ed by Lie inck.
The p esence o his species in Spain emains pe plex-
ing, bu we sugges ha i may be plausible wi h some
ca ea s (see below conce ning hos use). The wo spec-
imens om U alsk (NHMW) a e mo phologically con-
sis en wi h T. hellenicus; hey we e no ac ually labelled
as T. hellenicus by Lie inck and bo e labels o “Thy eus
p ae alens de . M.A. Lie inck” wi h no da e, bu hey
a e clea ly he specimens o which he was e e ing in his
1968 publica ion. Pa a ype labels ha e been added. The
p esence o T. hellenicus in wes e n Kazakhs an is possi-
ble i b oad hos use o An hopho a bees o he subgenus
Pa amegilla F iese, 1897, is he li e-his o y choice o his
pa asi e (see below). Finally, we belie e ha Lie inck’s
commen s ha T. hellenicus could be p esen in Roma-
nia (based on he geni al illus a ions o T. his ionicus
p esen ed by Iuga 1958: 214) a e unjus i ied. The geni-
al capsule illus a ed by Iuga has he gonos ylus small,
slende , and spa sely hai ed, sugges ing a membe o he
scu ella is-g oup. We he e o e conside he p esence
o T. hellenicus in Romania o be implausible based on
he cu en ly a ailable e idence, including e ision o
Thy eus ma e ial om no he n G eece and Bulga ia.
D sch. En omol. Z. 72 (2) 2025, 259–302
dez.penso .ne
281
Figu e 15. Thy eus hellenicus Lie inck, 1968 holo ype male (RMNH). A. Label de ails; B. Habi us, la e al iew; C. Face, on al
iew; D. Me asoma, do sal iew; Pa a ype male (RMNH); E. Geni al capsule, do sal iew; F. Geni al capsule, la e al iew.
In addi ion, based on new obse a ions on he island
o Amo gos, we can demons a e ha a leas one o he
hos s o T. hellenicus is An hopho a (Pa amegilla) su-
pe ans Walke , 1871. On 19 June 2023, in a ba e, sandy
a ea o he p ese ed beach o Agioi Sa an a on he island
o Amo gos, G eece, one o us (VL) obse ed h ee e-
males o T. hellenicus lying in and ou o nes s loca ed
in he g ound, wi h simul aneous isi a ion o he same
nes s by An hopho a emales, speci ically An hopho a
(Pa amegilla) supe ans. Mo eo e , An. supe ans was he
only la ge an hopho ine bee o he same size as T. helle-
nicus p esen a he collec ing e en (and possibly e en
on he island o Amo gos) a his ime. Fu he suppo
o his associa ion is he pa a ype o T. hellenicus col-
lec ed by Max Schwa z a Co in h on 3 June 1964; he
also collec ed a long se ies o An. supe ans a he same
momen (specimens OÖLM/TJWC; see Suppl. ma e ial
1). The same can be said o he pa a ype om Gy heio
on 3 June 1937 (NHMW), which was also collec ed wi h
se e al An. supe ans (specimens NHMW).
The name o he An hopho a hos equi es explana-
ion, as his name has no p e iously been used o he
dez.penso .ne
Thomas Wood e al.: A con ibu ion o Thy eus in he Palaea c ic, wi h wo new species282
Eu opean auna. The p esence o a la ge Pa amegilla spe-
cies in sou he n and Aegean G eece has been known o a
long ime; Dou s (1869: 120) used he name An hopho a
dubia E e smann, 1852, gi ing a dis ibu ion o “Îles de
l’A chipel g ec, mon agnes de l’Ou al, p o ince d’O en-
bou g, d’ap ès E e sman”. The iden i y o An. dubia is
cu en ly being deal wi h elsewhe e (Rasmon e al., in
e iew), bu in sho , An. dubia was desc ibed om O en-
bu g in sou he n Eu opean Russia and comp ises a mixed
ype se ies (males and emales inco ec ly associa ed).
Nei he axon wi hin his ype se ies is conspeci ic wi h
he G eek bee, which is an Eas Medi e anean species.
Sea ches we e he e o e made o he app op ia e name o
apply o his la ge and conspicuous An hopho a species.
Examina ion o ype ma e ial has concluded ha he oldes
names ha can be applied a e ei he An hopho a (Pa ame-
gilla) incly a Walke , 1871, o An hopho a (Pa amegil-
la) supe ans Walke , 1871. These wo names come om
Walke ’s (1871) wo k on Hymenop e a collec ed a ound
he Red Sea in Egyp , A abia, and E i ea. F ancis Walk-
e (1809–1874) was a p oli ic wo ke who has been much
c i icised o his abundan and supe icial desc ip ions; he
lack o espec o he wo k o Walke by his con empo a ies
was clea a he ime (see he 1874 obi ua y ci ed in E en-
huis 2008), a sen imen echoed by Bake (1993: 300), who
commen ed on he case o An. incly a and An. supe ans.
Much o he ma e ial desc ibed in Walke (1871) has been
los (e.g. see Monks e al. 2024), bu some specimens o An.
incly a and An. supe ans we e p ese ed in he collec ion o
F ede ick Smi h and a e now held in he NHMUK (Bake
1993; see O he ma e ial examined, below).
Walke (1871: 58, no. 287) desc ibed An. incly a in he
emale sex om “Ra la, Wady Fe an, Moun Sinai”. This
co esponds o A a ali/I a ayle in E i ea, Wadi Fei an in
Sinai, and Moun Sinai in he sou he n pa o he Sinai
Peninsula. He hen (Walke 1871: 58, no. 288) desc ibed
An. supe ans in he “ emale” sex om Moun Sinai only.
In he NHMUK collec ion, he e a e 3♀ o An. incly a and
4♂ o An. supe ans. Al hough Walke w o e “ emale” o
An. supe ans, he desc ibed he head as “Head, excep ing
he e ex whi e and wi hou hai s”, sugges ing a male,
since he males ha e he clypeus, sup aclypeal a ea, and
lowe pa aocula a eas wi h he in egumen yellow-whi e
(Fig. 17C). I is he e o e conside ed ha An. supe ans
was desc ibed om he male sex. A dissec ed male was
labelled as “lec o ype” by Bake in 1979 and by B ooks
in 1983 (Fig. 17A), bu nei he o hese designa ions was
published. Al hough Bake (1993) w o e ha i was des-
igna ed as such, his was published in his PhD hesis,
which does no mee he ICZN (1999) c i e ia o a alid
Figu e 16. Thy eus hellenicus Lie inck, 1968 pa a ype male (MNCN). A. Label de ails; B. P o ile; C. Me asoma, do sal iew;
D. Geni al capsule, do sal iew.
D sch. En omol. Z. 72 (2) 2025, 259–302
dez.penso .ne
283
publica ion. Unde A icles 8 and 9, as he hesis was no a
o mally published wo k and only se en ha d copies we e
p oduced, and hese limi ed numbe s we e no ob ainable
ei he o ee o o a cha ge, i ails A icles 8.1.2 and
8.1.3. This specimen om he Con en Ga den on Moun
Sinai (Sain Ca he ine’s Monas e y; 28.55°N, 33.98°E) is
he e o e inally published he e as he lec o ype o An. su-
pe ans, by p esen designa ion. I is conspeci ic wi h he
ma e ial om G eece based on he hind basi a si, which
ha e a ounded bump apically (Fig. 17D; no oo h o
ee h), and based on he en o-la e al p ojec ion o he
gonocoxa, which is educed (Fig. 17F; lec o ype specimen
wi h he geni al capsule ixed in a capsule wi h a glyce -
ine-like subs ance). This educ ion in he la e al p ojec ion
o he gonos ylus allows sepa a ion om he Wes Medi-
e anean An hopho a (Pa amegilla) e uginea Lepele i-
e , 1841 (compa e Fig. 18E, F). Indeed, he geni al capsule
is so simila (Fig. 18C, D) ha Lie inck ea ed ma e ial
(RMNH) om he Eas Medi e anean as An. e uginea
ssp. dubia; hese specimens ha e been e-de e mined (see
Figu e 17. An hopho a (Pa amegilla) supe ans Walke , 1871 lec o ype male (NHMUK). A. Label de ails; B. Habi us, la e -
al iew; C. Head, on al iew; D. Hind leg, la e al iew; E. Me asoma, do sal iew; F. Geni al capsule, in e ed along he
ans e se plane, la e al iew.
dez.penso .ne
Thomas Wood e al.: A con ibu ion o Thy eus in he Palaea c ic, wi h wo new species290
Based on he newly examined specimens, we hy-
po hesise a possible hos h ough examina ion o
concu en ly ac i e and cap u ed An hopho inae. A
Bajanaul (2008) and a Lake Baikal (2011), T. scu e-
lla is was cap u ed wi h An hopho a (Mys acan ho-
pho a) me idionalis Fedchenko, 1875, by W-H. Lieb-
ig (specimens WHLC). Ac i e a mos cap u e si es in
Kazakhs an (2024) we e nume ous species o An ho-
pho a (Pa amegilla) F iese, 1897, hough hese seem
less likely o be hos s due o (i) hei la ge size ela i e
o T. scu ella is and (ii) hei he mophilic endencies,
wi h mos Pa amegilla species a ou ing wa m o ho
g asslands and no ex ending in o coole pa s o Sibe ia
whe e T. scu ella is is also p esen . Clea ly, addi ional
ocused s udy is equi ed, bu his ci cums an ial e i-
dence o ms he basis o a es able hypo hesis.
Lie inck (1968) ede ined he global ange as p edom-
inan ly Asian. In Eu ope, ou side o Eu opean Russia,
he e is a single eco d om C oa ia om 1886 (Lie -
inck 1968); o ou knowledge, he nex nea es eco ds
come om Tu key and Russia (As akhan). Lie inck’s
eco ds om Egyp , “Sy ia”, and A abia a e di icul o
unde s and o biogeog aphical easons. Mos eco ds o
T. scu ella is, including he o iginal e a ypica, come
Figu e 24. Thy eus p ae alens (Kohl, 1905) male holo ype (NHMW). A. Label de ails; B. Habi us, la e al iew; C. Head, on al
iew; D. Head, la e al iew; E. Scu um, do sola e al iew; F. Me asoma, do sal iew.
D sch. En omol. Z. 72 (2) 2025, 259–302
dez.penso .ne
291
om a eas wi h a s ongly con inen al clima e, includ-
ing cold win e s. Molecula cha ac e isa ion, along wi h
addi ional s udy o Nea Eas e n specimens, is necessa y
o mo e ully unde s and he sou he n ange limi o his
species as well as i s po en ial hos (s).
Dis ibu ion. ?Egyp , ?Sy ia sensu la o, ?A abia sensu
la o, C oa ia, Russia (Eu opean pa , U als, Wes e n Si-
be ia, Eas e n Sibe ia), Tu key, I an, Tu kmenis an, Ka-
zakhs an, A ghanis an, Mongolia, China (Gansu, Inne
Mongolia) (Lie inck 1959b, 1968).
Iden i ica ion key o Eu opean Thy eus species
The key o Lie inck (1968) con ains good in o ma ion in
la ge pa bu is challenging o use due o he b oad geo-
g aphic scope ( om No h A ica o China and he Ko ean
Peninsula), he use o oo many cha ac e s o a iable im-
po ance, and mos o all, he inclusion o males and emales
in he same key. This limi s he abili y o he use o swi -
ly ecognise species g oups, such as he scu ella is-g oup,
based on he male geni alia, esul ing in ela ed species be-
ing sca e ed ac oss widely sepa a ed pa s o he key.
The ollowing key aims o ex ac he mos salien in-
o ma ion om Lie inck (1968) wi hin a na owe geo-
g aphic scope, speci ically ha o Eu ope. Dis ibu ional
in o ma ion in he key e e s only o he Eu opean ange,
and he b oade wo k o Lie inck (1968) should be con-
sul ed o non-Eu opean dis ibu ions. Fo males, i is
s ongly ad ised o ex ac he geni al capsule (all species)
and o examine he lobes o he 8 h s e na (ce ain species).
Despi e he use o geni alia cha ac e s by de Beaumon
(1940), some p e ious keys (e.g. Ve gés 1967; Amie e
al. 2007) ha e no e e ed o he male geni al capsule
a all, which p esen s di icul ies o Thy eus gi en he
high le el o di icul y ende ed by a eliance on ex e nal
mo phology. Addi ional non-geni al cha ac e s a e gi -
en in he male key, bu hei use in isola ion may lead o
inco ec iden i ica ions.
In addi ion o he images p esen ed he e, good illus a-
ions and addi ional cha ac e s and pe spec i es can be ound
in de Beaumon (1940), Lie inck (1968), Schwa z (1993),
and Amie e al. (2007). Female specimens a e inhe en ly
mo e di icul o iden i y, and ab aded specimens a e o en
impossible; con iden ly de e mined o ba coded e e ence
specimens should be consul ed whene e possible.
1 Six isible e ga, 12 an ennome es; emales....................................................................................................... 2
– Se en isible e ga, 13 an ennome es; males ................................................................................................... 13
2 (1) Body p edominan ly co e ed in dense adp essed whi e pubescence ha obscu es he unde lying su ace, in do sal
iew app oxima ely 75% o e gal su aces co e ed in such pubescence (Fig. 25A). Scu ellum pos e io ly p oduced
in o wo e y long acu e poin s, hese o ming a deep U-shaped ema gina ion (Fig. 25B). Ou e ma gins o axillae
sligh ly de ached om ou e ma gin o scu ellum, o ming mode a ely acu e poin s. Found only on Cyp us ..........
............................................................................................................................................T. elegans (Mo awi z)
– Body wi h whi e pubescence, bu ne e co e ing a majo i y o he su ace, e ga always p edominan ly da k. Scu e-
llum pos e io ly p oduced in o blun o ounded poin s, o i p oduced in o acu e poin s hen wi h he median
ema gina ion much shallowe (Fig. 25D). Ou e ma gins o axillae ei he lush wi h ou e ma gin o scu ellum (e.g.
Fig. 25F) o o ming small sha p poin s (e.g. Fig. 25D). Found on Cyp us o no ................................................. 3
3 (2) T1 wi h ou dis inc and elonga e whi e hai pa ches, wo placed basally and wo apically, o ming widely in-
e up ed basal and apical hai bands; bands clea ly sepa a ed (in e up ed) by black hai s on la e al ma gins
(Fig. 25C, E) ...................................................................................................................................................... 4
– T1 wi h wo dis inc whi e hai pa ches placed la e ally (Fig. 27A, B), hese ei he C-shaped o L-shaped, widely
in e up ed medially in he cen e o he disc; always wi h con inuous whi e hai s on he la e al pa s o he disc
(unin e up ed).................................................................................................................................................. 5
4 (3) Pos e io ma gin o scu ellum dis inc ly imp essed medially, disc o scu ellum hus no la ened, dis inc ly bicon-
ex (Fig. 25D). Scu ellum pos e io ly p oduced in o sha p acu e poin s. Ou e ma gins o axillae sligh ly de ached
om ou e ma gin o scu ellum, o ming small sha p poin s. Ra e in Eu ope, known as a wes as C oa ia, p oba-
bly pe sis ing mos ly in Eu opean Russia .......................................................................... T. scu ella is (Fab icius)
– Scu ellum e enly la ened, pos e io ma gin o scu ellum no dep essed medially (Fig. 25F). Scu ellum pos e i-
o ly p oduced in o blun 90° poin s. Ou e ma gin o axillae lush wi h ou e ma gin o scu ellum, no p oduced
in o a poin . Widesp ead ac oss sou he n Eu ope om Po ugal o Bulga ia, G eece, and Cyp us, no h o he
Pannonian basin ..................................................................................................................... T. a inis (Mo awi z)
5 (3) Scu um la e ally wi h a con inuous band o sho whi e hai s unning up o he junc ion wi h he axillae (Fig. 26A)
Small, always < 10 mm in leng h ........................................................................................T. amosus (Lepele ie )
– Scu um la e ally ne e wi h an unb oken band o whi e hai s, always wi h some black hai s (Fig. 26B). La ge ,
usually > 10 mm in leng h, hough some small specimens o T. unca us can be 9 mm in leng h ....................... 6
6 (5) Found on he island o G an Cana ia (Cana y Islands) ....................................................................................... 71
– Found elsewhe e in Eu ope, including o he Cana ian islands ............................................................................. 8
1 Due o he es ic ed geog aphic dis ibu ion, T. hohmanni is keyed ou he e; mos wo ke s will ne e encoun e his species.
dez.penso .ne
Thomas Wood e al.: A con ibu ion o Thy eus in he Palaea c ic, wi h wo new species292
7 (6) Face wi h abundan sho black pubescence below he an ennal inse ions. T1, when iewed do sally wi h na ow
L-shaped whi e hai pa ches la e ally, hese ba ely ex ending owa ds he cen e o he disc a i s base. Scu ellum
densely punc a e, punc u es sepa a ed by ≤ 0.5 punc u e diame e s ..................................... T. hohmanni Schwa z
– Face en i ely whi e-hai ed. T1, when iewed do sally wi h b oad C-shaped whi e hai pa ches la e ally, hese ex-
ending inwa ds owa ds he cen e o he disc almos as much a he base o he disc as on he ma ginal a ea
(Fig. 27A). Scu ellum mo e spa sely punc a e wi h punc u es sepa a ed by 0.5–1 punc u e diame e s, in e spaces
polished and shining ...............................................................................................T. his ionicus (Illige ) (pa im)
8 (6) Found in G eece wi h he ollowing combina ion o cha ac e s: scu ellum sho and b oad, medially wi h a na ow
bu deep sulca e imp ession, su ace densely punc a e, punc u es sepa a ed by ≤ 0.5 punc u e diame e s (Fig.
23C). Ve ex behind ocella iangle densely punc a e, punc u es sepa a ed by 0.5 punc u e diame e s, s ongly
con as ing a eas adjacen o la e al ocelli, which a e polished, shining, and almos impunc a e (Fig. 23E). T1
iewed do sally wi h small iangula hai pa ches, hese ba ely ex ending owa ds he base o he e gum (Fig.
23D). An enna wi h hina ia elonga e lanceola e, longi udinally-o ien a ed, and placed on he en al su ace o he
an enna. Face below an ennal inse ions wi h abundan da k pubescence (Fig. 23B) .............. T. p ae alens (Kohl)2
– Dis ibu ion o combina ion o cha ac e s no as abo e ..................................................................................... 93
9 (8) Face below an ennal inse ions usually wi h abundan black hai s (Fig. 26C), bu some specimens o T. o ba us
om sou he n Spain (Sie a Ne ada) p esen aces wi h abundan whi e pubescence and only a ew sca e ed black
hai s. Fo T. o ba us, i in doub , hen la e al whi e hai pa ch o T3 in e up ed (Fig. 26D), dis inc ly in e up ed by
black hai s and hus o ming wo dis inc ly sepa a ed whi e hai pa ches ......................................................... 10
– Face below an ennal inse ions always whi e-hai ed (Fig. 26E), ei he en i ely whi e-hai ed o a mos wi h only a
ew sca e ed black hai s. La e al whi e hai pa ch o T3 always en i e, ne e b oken in o wo dis inc whi e hai
pa ches ........................................................................................................................................................... 11
10 (9) La e al whi e hai pa ch o T3 in e up ed, dis inc ly in e up ed by black hai s and hus o ming wo dis inc whi e
hai pa ches (Fig. 26D). Usually a li le smalle , ange 8–10 mm. Associa ed wi h An hopho a quad imacula a. The
mos no he ly Thy eus species, ex ending om he moun ains o sou he n Eu ope in o he Low Coun ies and
no he n Ge many, as well as eas o he Balkan Peninsula .................................................. T. o ba us (Lepele ie )
– La e al whi e hai pa ch o T3 always en i e, ne e b oken in o wo dis inc whi e hai pa ches (Fig. 26F). Usually a
li le la ge , ange 10–12 mm. Associa ed wi h Echium-specialised An hopho a (Pa amegilla) species. Res ic ed o
he Wes Medi e anean no h o sou he n Swi ze land ..................................................... T. hi us (de Beaumon )
11 (9) T1, when iewed do sally wi h b oad C-shaped whi e hai pa ches la e ally, hese ex ending inwa ds owa ds he
cen e o he disc almos as much a he base o he disc as on he ma ginal a ea (Fig. 27A). Scu ellum mode a ely
densely punc a e, punc u es sepa a ed by 0.5–1 punc u e diame e s, in e spaces polished and shining (Fig. 27C)
...............................................T. his ionicus (Illige ) (pa im) and pica on Lie inck ( emales cu en ly insepa able)
– T1 when iewed do sally wi h na ow L-shaped whi e hai pa ches la e ally, hese ex ending inwa ds owa ds he
cen e o he disc only on he ma ginal a ea (Figs 7F, 9F, 27B). Scu ellum e y densely punc a e, punc u es sepa-
a ed by ≤ 0.5 punc u e diame e s, in e spaces weakly shining o dull (Figs 7E, 9E, 27D) ................................. 12
12 (11) Smalle species, body leng h ange 9–12 mm. Rhina ia absen . Scu ellum sho and b oad, su ace almos la ,
pos e io angles blun and poo ly indica ed (Figs 7E, 9E). Associa ed wi h Te alonia den a a. Widesp ead ac oss
con inen al Eu ope ..................................................................................................................T. unca us (Pé ez)
– La ge species, body leng h a ound 15 mm. Wi hou hina ia, bu wi h a se ies o low longi udinal idges. Scu ellum
longe , pos e io angles mo e s ongly p oduced (Fig. 27D). Associa ed wi h An hopho a supe ans. P edominan ly in
G eece, including he islands, bu possibly also sou he n Spain ........................................... T. hellenicus Lie inck
13 (1) Geni al capsule simple, wi h gonos ylus b oadly iangula o p oduced in o an acu e poin ; ne e wi h an addi ional
do sal p ojec ion (Figs 2F, 28A, B). Gonos ylus almos hai less, ne e wi h long hai s. T1 wi h hai s p esen basally
and apically, o ming ei he a comple e basal and nea ly comple e apical bands (Fig. 29A) o wi h bands in e up ed
medially and sepa a ed la e ally, hus appea ing o o m ou dis inc hai pa ches (Fig. 29B) (scu ella is-g oup) ...14
– Geni al capsule complex, wi h gonos ylus p esen ing a ounded o unca e apex (Figs 3H, 28C–H); gonos ylus
o en wi h an addi ional basodo sal p ojec ion oughly a a 90° angle o he la ges lobe o he gonos ylus (Figs
3H, 28C–E). Gonos ylus wi h long hai s, hese o en abundan and co e ing and obscu ing he su ace (Figs 3H,
28C–H). T1 wi h hai s a iable, bu ne e o ming a comple e basal band and always comple e la e ally, hus ne e
o ming ou dis inc hai pa ches .................................................................................................................... 16
2 Thy eus p ae alens is keyed ou he e h ough a combina ion o cha ac e s; whils i could po en ially be included in he ollowing
couple based on he da k acial pubescence, i emains unclea i his is a obus cha ac e , and so a cau ious app oach is aken he e.
3 Sepa a ion o he ollowing six species depends hea ily on pubescence cha ac e s, which a e necessa ily dependen on specimen
condi ion. No a emp should be made o y and iden i y s ongly ab aded specimens, and associa ion wi h concu en ly ac i e
males should be made whe e possible.
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14 (13) Body abundan ly whi e-hai ed, pa icula ly hickly co e ing he ace (including ons), mesepis e num, and an e io
½ o scu um, obscu ing he unde lying su ace. T1 wi h hai s o ming a comple e basal band and almos comple e
apical band, hese bands connec ed la e ally (Fig. 29A). T2 wi h b oad apical hai band co e ing ½ he leng h o
he e gum, only na owly in e up ed medially. Found only on Cyp us ...................................T. elegans (Mo awi z)
– Body wi hou abundan whi e pubescence, only mode a ely hai y, whi e hai s o ace, mesepis e num, and
scu um no obscu ing unde lying in egumen . T1 wi h ou dis inc and elonga e whi e hai pa ches, wo
placed basally and wo apically, o ming widely in e up ed basal and apical hai bands (Fig. 29B). Found on
Cyp us o no ......................................................................................................................................... 15
15 (14) Pos e io ma gin o scu ellum dis inc ly imp essed medially, disc o scu ellum hus no la , dis inc ly bicon ex (as
in Fig. 4C). Scu ellum pos e io ly p oduced in o sha p acu e poin s, medially wi h loose hin whi ish hai s on do sal
su ace, hese no obscu ing he unde lying su ace. Ou e ma gin o axillae sligh ly de ached om ou e ma gin o
scu ellum, p oduced in o a sub le bu dis inc poin (as in Fig. 4C). Ra e in Eu ope, known as a wes as C oa ia,
p obably pe sis ing mos ly in Eu opean Russia ................................................................. T. scu ella is (Fab icius)
– Scu ellum e enly la , pos e io ma gin o scu ellum no dep essed medially. Scu ellum pos e io ly p oduced in o
blun ~90° poin s, medially wi h dis inc pa ch o adp essed hick whi e hai s which obscu es he unde lying su -
ace (Fig. 29C; may be ab aded in old specimens). Ou e ma gin o axillae lush wi h ou e ma gin o scu ellum, no
p oduced in o a poin . Widesp ead ac oss sou he n Eu ope om Po ugal o Bulga ia, G eece, and Cyp us, no h
o he Pannonian basin ........................................................................................................... T. a inis (Mo awi z)
16 (13) In la e al iew, gonos ylus dis inc ly bi u ca e, wi h a la ge apically p ojec ing and unca e lobe and basal de ached
and do sally p ojec ing acu ely poin ed lobe o ien ed oughly 90° ela i e o he unca e lobe (Fig. 3H, 28C–E) 17
– In la e al iew, gonos ylus wi h only a single apically p ojec ing lobe, ne e wi h a dis inc de ached do sally p o-
jec ing lobe (Fig. 28F–H) .................................................................................................................................. 20
17 (16) Apical p ojec ions o S8 slende wi h dis inc 90° bend, hus appea ing dis inc ly “L-shaped” (Fig. 3F) .................
............................................................................................................................................... T. pica on Lie inck
– Apical p ojec ions o S8 pa allel-sided, apexes unca e o ounded (Fig. 29D) ................................................. 18
18 (17) Face wi h abundan black hai s on he ace below he an ennal inse ions. In do sal iew, la e al ma gins o T1 wi h
whi e hai s loose, no adp essed, la e ally e ec and p ojec ing hus con as ing he whi e hai s on he ma ginal
a ea o T1 which a e adp essed, obscu ing he unde lying su ace (Fig. 29E). Res ic ed o he Wes Medi e anean
no h o sou he n Swi ze land .......................................................................................... T. hi us (de Beaumon )
– Face en i ely whi e-hai ed. In do sal iew, la e al ma gins o T1 wi h whi e hai s adp essed, no di e en ia ed om
adp essed whi e hai s on he ma ginal a ea o T1 (Fig. 29F). Widesp ead h oughou Eu ope ........................... 19
19 (18) Geni al capsule wi h do sal p ojec ion s ongly p oduced, s anding dis inc ly away om ou e ma gin o gonocoxa,
wi h long hai s, leng h o hese hai s a leas wice he b ead h o he do sal p ojec ion (Fig. 28D). Main lobe o
gonos ylus wi h abundan long hai s; in la e al iew, hai s p ojec ing beyond he apex o he gonos ylus by a dis-
ance equal o he maximum b ead h o he hind basi a sus (Fig. 30A) .................................T. his ionicus (Illige )
– Geni al capsule wi h do sal p ojec ion weakly p oduced, no s ongly isola ed om ou e ma gin o gonocoxa, wi h
sho hai s, hese hai s equalling o only sligh ly exceeding b ea h o he do sal p ojec ion (Fig. 28E). Main lobe
o gonos ylus wi h abundan bu sho hai s; in la e al iew, hai s only sligh ly p ojec ing by much less han he
maximum b ead h o he hind basi a sus (Fig. 30B) ................................................................T. unca us (Pé ez)
20 (16) Scu ellum sho and b oad, medially dis inc ly imp essed and na owly sulca e (Fig. 24E). T1 iewed do sally
wi h small apicola e al iangula hai pa ches, no ex ending owa ds base o e gum (Fig. 24F). Ve ex behind
ocella iangle densely punc a e, punc u es sepa a ed by 0.5 punc u e diame e s, s ongly con as ing a eas
adjacen o la e al ocelli, which a e polished, shining, and almos impunc a e (Fig. 23F). Found only in G eece ..
......................................................................................................................................... T. p ae alens (Kohl)
– Scu ellum la , wi hou such a median sulca e imp ession. T1 iewed do sally wi h hai pa ches na ow o
b oad, bu always ex ending o he base o he e gum. Ve ex wi hou such s ongly con as ing punc a ion. Dis-
ibu ion a iable ............................................................................................................................................. 21
21 (20) Face below he an ennal inse ions usually wi h abundan black hai s (c . Fig. 26C). In well-p ese ed specimens,
T3 la e ally wi h whi e band in e up ed, hus o ming wo dis inc whi e hai pa ches (Fig. 30C). Hind ibia en-
ally p oduced in o a small bu dis inc sha ply poin ed oo h o e lying ou e ibial spu (di icul o see wi hou
p ac ice). Geni alia as in Fig. 28F. The mos no he ly Thy eus species, ex ending om he moun ains o sou he n
Eu ope in o he Low Coun ies and no he n Ge many o a ound 52°N .................................T. o ba us (Lepele ie )
– Face below he an ennal inse ions ne e wi h abundan black hai s (Figs 15C, 16B). T3 la e ally wi h a single
unb oken whi e hai band (Figs 15B, 16B, 30D). Hind ibia wi hou such a en ally p ojec ing oo h. Geni alia as in
Figs 15E and 30E–F. No ex ending no h o 49°N ............................................................................................ 22
22 (21) Geni al capsule wi h gonos ylus long, in la e al iew gonos ylus elonga e iangula , s ongly p ojec ing away om
he gonocoxa by a dis ance exceeding he maximum b ead h o a hind basi a sus (Fig. 15F). In do sal iew, he
dez.penso .ne
Thomas Wood e al.: A con ibu ion o Thy eus in he Palaea c ic, wi h wo new species294
long gonos yli gi e he geni al capsule a homboidal appea ance (Fig. 15E). La ge, body leng h a ound 15 mm.
Res ic ed o G eece, including he islands ........................................................................... T. hellenicus Lie inck
– Geni al capsule mo e compac , gonos ylus sho , in la e al iew gonos ylus ba ely p ojec ing beyond he gonocox-
ae. In do sal iew, geni al capsule o oid o ounded (Fig. 28G, H). Ei he la ge bodied (12–13 mm), in which case
ound only on he Cana y Islands, o i ound in con inen al Eu ope, hen smalle , < 10 mm in leng h............... 23
23 (22) T7 wi h apical ma gin ema gina e, ema gina ion ounded ec angula , b oade han deep (Fig. 30E). Smalle , body
leng h < 10 mm. Widesp ead ac oss con inen al Eu ope, bu no p esen on he Cana y Islands ...........................
..........................................................................................................................................T. amosus (Lepele ie )
– T7 wi h apical ma gin ema gina e, ema gina ion e enly ounded (Fig. 30F). La ge, body leng h 12–13 mm. Res ic -
ed o G an Cana ia (Cana y Islands) .....................................................................................T. hohmanni Schwa z
Figu e 25. Thy eus elegans (Mo awi z, 1877) emale. A. Me asoma, do sal iew; B. Scu ellum, do sal iew. Thy eus scu ella is
(Fab icius, 1781) emale; C. T1, do sal iew; D. Scu ellum, do sal iew. Thy eus a inis (Mo awi z, 1873) emale; E. T1, do sal
iew; F. Scu ellum, do sal iew.
D sch. En omol. Z. 72 (2) 2025, 259–302
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Discussion
The esul s p esen ed he e and he axonomic concep s
e ined h ough he inclusion o COI DNA ba coding
o e all suppo he axonomic amewo k o Lie inck
(1968) o he Wes and Cen al Palaea c ic species
s udied o da e. The small numbe o changes necessa y
ha e ela ed o mino e o s in dis ibu ions esul ing
om misiden i ied o c yp ic species, and he inclusion
o gene ic da a and concu en s udy wi h hei p edom-
inan ly an hopho ine hos s has allowed ange limi s o
be cha ac e ised wi h g ea e con idence.
Al hough mos opical species o Thy eus a e likely
o exclusi ely a ack Amegilla due o he absence o
Figu e 26. Thy eus amosus (Lepele ie , 1841) emale. A. Scu um, do sola e al iew. Thy eus his ionicus (Illige , 1806) emale;
B. Scu um, do sola e al iew; E. Head, on o-la e al iew. Thy eus hi us (Beaumon , 1940) emale; C. Head, on o-la e al iew,
F. Me asoma, la e al iew. Thy eus o ba us (Lepele ie , 1841) emale; D. Me asoma, la e al iew.
dez.penso .ne
Thomas Wood e al.: A con ibu ion o Thy eus in he Palaea c ic, wi h wo new species296
he genus An hopho a in e y humid a eas ( o exam-
ple, in mos o Sou h Eas Asia and all o Aus alasia;
B ooks 1988), he pa e n in he Palaea c ic is much
mo e mixed. Species known o suspec ed o a ack
An hopho a include T. hellenicus, T. hi us, and T. o -
ba us; species known o suspec ed o a ack Amegil-
la include T. his ionicus, T. pica on, and T. amosus,
whils T. unca us a acks bees o a di e en sub amily
(Euce inae). Thy eus elegans sensu la o is suspec ed
o a ack Amegilla (Mic amegilla) B ooks, 1988 based
on co-occu ence (TJW, unpublished da a), and hy-
po heses a e p esen ed he e o associa ions be ween
T. jansseni and T. scu ella is wi h An hopho a (Mys a-
can hopho a) species. E en in he absence o a mode n
phylogeny o he genus Thy eus, i is di icul o see
how his would ansla e in o a pa e n o conse ed
hos use ela i e o he phylogene ic s uc u e seen in
apid bees (Bosse e al. 2019).
Finally, whils gene ic concep s a e cu en ly un-
de e ision in melec ine bees due o he gene a ion o
powe ul new gene ic da a and mo phological analyses
(Onu e ko e al. 2021; O e al. 2024), he p esen
wo k se es o es a e he need o obus ly es spe-
cies-le el concep s. I also emphasises he impo ance
o balancing molecula esul s wi h mo phological c i-
e ia. Al hough gene ic a ia ion in he COI ba code
is much highe in T. unca us han in o he Thy eus
species s udied he e, i was no possible o ind mo -
phological c i e ia consis en wi h ba code clus e ing
o allow o axon delinea ion; hence, a conse a i e
and b oad species concep was main ained. Whils he
high ba code di e ences sugges a pa e n o his o ical
isola ion, pe haps in an Ibe ian e ugium o Ibe ian
specimens, eliance pu ely on gene ic dis ance would
p oduce a challenging sys em o mo phological iden-
i ica ion, as well as po en ially lead o nomencla u al
ins abili y depending on which mi ochond ial lineage
he lec o ype is assigned o. This app oach is consis-
en wi h o he ecen ea men s (e.g. P az and Benon
2023) and will hope ully main ain he undamen ally
obus species concep s o Lie inck (1968) o cu en
and u u e species moni o ing.
Figu e 27. Thy eus his ionicus (Illige , 1806) emale; A. T1, do sal iew; C. Scu ellum, do sal iew. Thy eus hellenicus Lie inck,
1968, emale; B. T1, do sal iew; D. Scu ellum, do sal iew.
D sch. En omol. Z. 72 (2) 2025, 259–302
dez.penso .ne
297
Figu e 28. Thy eus elegans (Mo awi z, 1877) male. A. Geni al capsule, do sal iew. Thy eus a inis (Mo awi z, 1873) male; B. Gen-
i al capsule, do sal iew. Thy eus hi us (Beaumon , 1940) male; C. Geni al capsule, do sal iew. Thy eus his ionicus (Illige , 1806)
male; D. Geni al capsule, do sal iew. Thy eus unca us (Pé ez, 1884) male; E. Geni al capsule, do sal iew. Thy eus o ba us
(Lepele ie , 1841) male; F. Geni al capsule, do sal iew. Thy eus hohmanni Schwa z, 1993 male; G. Geni al capsule, do sal iew.
Thy eus amosus (Lepele ie , 1841) male; H. Geni al capsule, do sal iew.
dez.penso .ne
Thomas Wood e al.: A con ibu ion o Thy eus in he Palaea c ic, wi h wo new species298
Figu e 29. Thy eus elegans (Mo awi z, 1877) male. A. T1, do sal iew. Thy eus a inis (Mo awi z, 1873) male; B. T1, do sal iew.
C. Scu ellum, do sal iew. Thy eus hi us (Beaumon , 1940) male; D. S8, do sal iew; E. La e al pa o T1, do sal iew. Thy eus
his ionicus (Illige , 1806) male; F. La e al pa o T1, do sal iew.
D sch. En omol. Z. 72 (2) 2025, 259–302
dez.penso .ne
299
Figu e 30. Thy eus his ionicus (Illige , 1806) male. A. Geni al capsule, la e al iew. Thy eus unca us (Pé ez, 1884) male; B. Gen-
i al capsule, la e al iew. Thy eus o ba us (Lepele ie , 1841) male; C. Me asoma, la e al iew. Thy eus amosus (Lepele ie , 1841)
male; D. Me asoma, la e al iew; E. T7, do sal iew. Thy eus hohmanni Schwa z, 1993 male; F. T7, do sal iew.
