Re ision o he genus Lepidop ia Kie e (Hymenop e a,
Diap iidae, Diap iinae) o he wo ld auna
Vasilisa G. Chemy e a1,2, Seonwoo Yoon3, Deok-Seo Ku4
1A.A. Bo issiak Paleon ological Ins i u e, Russian Academy o Sciences, P o soyuznaya 123, Moscow, 117647,
Russia 2Zoological Ins i u e, Russian Academy o Sciences, 1 Uni e si e skaya Emb., S . Pe e sbu g, 199034,
Russia 3Ulsan Ag icul u al Technology Cen e , Ulsan, Republic o Ko ea 4The Science Museum o Na u al
Enemies, Geochang 50147, Republic o Ko ea
Co esponding au ho : Vasilisa G. Chemy e a ([email p o ec ed])
Academic edi o : Elijah Talamas|Recei ed 31 Augus 2025|Accep ed 4 No embe 2025|Published 18 No embe 2025
h ps://zoobank.o g/D1292408-DD83-4D78-A557-F3A63128ACB4
Ci a ion: Chemy e a VG, Yoon S, Ku D-S (2025) Re ision o he genus Lepidop ia Kie e (Hymenop e a, Diap iidae,
Diap iinae) o he wo ld auna. Jou nal o Hymenop e a Resea ch 98: 1107–1126. h ps://doi.o g/10.3897/jh .98.169802
Abs ac
All known species o he diap iid genus Lepidop ia Kie e , 1911 a e e ised, keyed and illus a ed. A new
species, L. bica ina a Chemy e a & Ku, sp. no . is desc ibed om Sou h Ko ea. The diagnosis o he
genus Lepidop ia, as well as ha o all desc ibed species, has been cla i ied. New da a on he dis ibu ion
o he species L. masne i No on, 1994 and L. pedes is Kie e , 1911 a e p o ided. The associa ion o
L. masne i wi h he nes s o Solenopsis japonica Wheele , 1928 is epo ed and he biology o Lepidop ia
species is discussed. Taxonomic s a us o he mys e ious L. abe ans B ues, 1916 is conside ed and a new
combina ion, B uesop ia abe ans (B ues), comb. no ., sugges ed.
Keywo ds
B uesop ia, Diap iini, my mecophilic wasps, new eco ds, new species, symphiles, Solenopsis, axonomy
In oduc ion
Lepidop ia Kie e , 1911 comp ises a ew species o iny wasps belonging o he my -
mecophilic diap iids (Hugge and Masne 1983). The genus cu en ly includes he
ollowing ou species: L. abe ans B ues, 1916, eco ded in Ma yland, USA; L. lloydi
(Fe iè e, 1935), eco ded in Tanzania; L. masne i No on, 1994, eco ded in Japan
JHR 98: 1107–1126 (2025)
doi: 10.3897/jh .98.169802
h ps://jh .penso .ne
Copy igh Vasilisa G. Chemy e a 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.
RESEARCH ARTICLE
V. G. Chemy e a e al. / Jou nal o Hymenop e a Resea ch 98: 1107–1126 (2025)
1108
and Sou h Ko ea; and L. pedes is Kie e , 1911, eco ded in Eu ope. Acco ding o
Masne and Muesebeck (1968), he L. abe ans is doub ully placed in his genus and
may belong o he Nea c ic genus B uesop ia Wing, 1951 (Hugge and Masne 1983;
Masne and Ga cía 2002); howe e , he easons o he doub s and he mo phological
ea u es o his species we e no discussed. The speci ic cha ac e s o he Lepidop ia spe-
cies, such as educed eye size and numbe o palpal segmen s, educed pigmen a ion
(especially in he emale an enna, which has a yellow cla a), he seconda y ap e ism,
abundan se a ion, and an enla ged pe iole (simila o many an s), sugges ha hey all
ha e my mecophilic habi s (No on 1994). Lepidop ia abe ans and L. pedes is we e
ea ed om he pupa ia o achinid lies, which we e ound inside adul sca abaeid bee-
les (B ues 1916; Hugge and Masne 1983). I has also been con i med ha bo h spe-
cies a e symphilic in he nes s o an s o he genus Solenopsis Wes wood, 1840 (B ues
1916; Fe iè e 1927; Hölldoble 1928; Wing 1951; Lachaud 1981b; Lachaud and
Passe a 1982; Masne and Ga cía 2002). The in e ac ion wi h an s has been s udied in
de ail o only one species, L. pedes is (Hölldoble 1928; Lachaud 1980, 1981a, b, c,
1982; Lachaud and Passe a 1982).
Acco ding hese s udies L. pedes is is a highly adap ed my mecophilous diap iid
wasp ha li es in close associa ion wi h he an Solenopsis ugax (La eille, 1798). The
wasp in eg a es in o an colonies p ima ily by mimicking o sha ing ac ile solici a ion
signals and by p oducing cu icula sec e ions ha induce in ense g ooming o licking
om an wo ke s. Adul s (especially emales) ob ain ood by ophallaxis om an s in-
side nes s wi hou agg ession (Lachaud 1982). Mo phological specializa ions like clus-
e s o ac i e sec e o y cells opening in o dense u s o hai s on he pe iole and an e io
pa o S2 ( hese sec e ions a e associa ed wi h he egions an s lick mos in ensi ely)
unde lie chemical in eg a ion o L. pedes is in he an colony (Lachaud 1981a). The
o e leg apping pa e ns and an ennal con ac s a e simila o hose used by an s and
unde lie he beha io al in eg a ion o L. pedes is (Lachaud 1981c, 1982). The success
o he wasp’s in eg a ion is mos clea ly demons a ed by he ac ha wo ke s o S.
ugax may egu gi a e ood o newly in oduced L. pedes is wi hin ~10 minu es and
show e y li le agg ession (Lachaud 1981b).
A de ailed phenological s udy o L. pedes is was ca ied ou by Lachaud and Pas-
se a (1982). Al hough he au ho s o his s udy we e unable o es ablish he p ima y
hos o his species, hey eached se e al signi ican conclusions. Fi s ly, L. pedes is
is undoub edly my mecophilic; i uses Solenopsis ugax nes s o p o ec ion, eeding
pu poses and hibe na ion in win e . Secondly, L. pedes is does no use S. ugax as a
hos and does no ep oduce in i s nes . Thi dly, L. pedes is specimens spend mos o
hei li e cycle ou side he nes ( om ea ly sp ing o la e au umn), whe e hey ma e,
pa asi ize he p ima y hos and de elop in o adul s, and hen, when win e a i es,
hey e u n o he S. ugax nes s. Finally, L. pedes is is s ic ly associa ed wi h S. ugax
nes s, since i is no ound in he nes s o o he an species pa asi ized by he les obio ic
S. ugax and was e en elimina ed by wo o hese species (Plagiolepis pygmaea (La eille,
1798) and Te amo ium semilae e And é, 1883) when expe imen ally in oduced in
hei nes (Lachaud and Passe a 1982). Only la e he main hos o L. pedes is was e-
Re ision o Lepidop ia (Hymenop e a, Diap iidae) 1109
po ed as achinid lies, which pa asi ise sca abaeid bee les ha a e no associa ed wi h
an nes s (Hugge and Masne 1983).
Un o una ely, we do no know much abou he li e his o ies o o he Lepidop ia
species. The A ican species Lepidop ia lloydi was ea ed om pupa ia o he Glossina
Wiedemann, 1830 (Fe iè e 1935; No on 2014), whe eas he biology o L. masne i
emains unknown (No on 1994). These wo species ha e mo phology ha sugges
my mecophily, howe e , hei connec ion wi h an s has no ye been documen ed.
This wo k aims o e iew he mo phology and biology o all desc ibed species
o he genus Lepidop ia, cla i y he diagnos ic cha ac e is ics o his genus and some
closely ela ed gene a, and desc ibe a new species o Lepidop ia om he sou h-eas e n
Palaea c ic egion.
Ma e ial and me hods
Ma e ial obse ed in his s udy o igina es om he se e al collec ions: MCSN − he
Museo Ci ico di S o ia Na u ale “Giacomo Do ia”, Genoa, I aly; NHML − he Na u-
al His o y Museum, London, UK; NIBR − he Na ional Ins i u e o Biological Re-
sou ces, Incheon, Republic o Ko ea; SMNE − he Science Museum o Na u al En-
emies, Geochang, Republic o Ko ea; NMNH − he Na ional Museum o Na u al
His o y, Smi hsonian Ins i u ion, Washing on, USA; ZISP − he Zoological Ins i u e
o he Russian Academy o Sciences, S Pe e sbu g, Russia; UATC – Ulsan Ag icul u al
Technology Cen e , Republic o Ko ea.
Mos o images we e aken wi h a Nikon s e eo mic oscope SMZ25 equipped wi h
objec i e lens Plan Apo 1.6× and digi al came a DS-10. Some images we e aken wi h a
came a Olympus E-M5 Ma k II and objec i e lens M.Zuiko 60 mm 1:2.8 combined wi h
a Raynox MSN-505 (Fig. 7A, C). Final images we e s acked using Helicon Focus 7.6.
The mo phological e minology, abb e ia ions and measu emen s ollow Masne
and Ga cía (2002), Yode (2004) and he Hymenop e a Ana omy On ology (Yode e
al. 2010). New egional and coun y eco ds a e indica ed wi h an as e isk (*).
Taxonomy
Genus B uesop ia Wing, 1951
No e. A de ailed diagnosis o his genus has been gi en by Masne and Ga cía (2002).
The e o e, he e we gi e only a sho compa a i e diagnosis o his genus, emphasizing
he diagnos ic di e ences be ween i and he genus Lepidop ia.
Compa a i e diagnosis. Female an enna 11–12-me ous [only 12-me ous in Lep-
idop ia], wi h ab up 3-me ous cla a [nonab up o 4-me ous cla a in Lepidop ia];
pos e io ma gin o p opodeum wi h deep semici cula exca a ion, median ca ina o
p opodeum udimen a y, pos e ola e al co ne o p opodeum angula and s ongly
V. G. Chemy e a e al. / Jou nal o Hymenop e a Resea ch 98: 1107–1126 (2025)
1110
p ojec ing [same in Lepidop ia o wi h median ca ina, wi hou exca a ion o pos e io
ma gin and wi hou pos e ola e al p ojec ion]; pe iole s ongly modi ied, ema kably
highe han long in la e al iew, wi h inge -like p ojec ion p oduced do sally abo e
an e io ma gin o T2 (Fig. 1C, D) [same in Lepidop ia (Fig. 9E) o as long as high in
la e al iew and no p oduce do sally abo e T2 (Fig. 2C, D)]; base o S2 ba e, smoo h,
la : no con ex medially and no g oo ed la e ally, wi h deep semici cula exca a ion
o an e io ma gin (Fig. 1A) [densely se ose, wi h la e al g oo es and con ex medially,
exca a ion o an e io ma gin no deep in Lepidop ia (Fig. 1B)].
Rema ks. This genus comp ises only wo desc ibed species om he New Wo ld
(Wing 1951; Johnson 1992). Masne and Ga cía (2002) epo ed an undesc ibed spe-
cies associa ed wi h Solenopsis sp. in A izona. An unde e mined species o B uesop ia
(possibly B. abe ans) was ound by D Alexande L. Wild in a colony o Solenopsis
moles a (Say, 1836) a Konza P ai ie in Kansas, USA (see Fig. 1E, F). The biodi e si y
and mo phological a iabili y o he h ee closely ela ed gene a – Lepidop ia, B ueso-
p ia, and Solenopsia Wasmann, 1899 – ha e no ye been su icien ly s udied and u u e
disco e ies may necessi a e he synonymisa ion o hese gene a (Masne and Ga cía
2002). Howe e , he diagnos ic di e ences be ween Lepidop ia and B uesop ia, as de-
sc ibed abo e, a e p esen ly clea enough o main ain hese axa as alid. Fu he mo e,
hese gene a ha e di e en dis ibu ions: all Lepidop ia species a e ound in he Palae-
a c ic egion, whe eas all B uesop ia species a e ound in he Nea c ic egion.
B uesop ia abe ans (B ues, 1916), comb. no .
Fig. 1C, D
Lepidop ia abe ans B ues, 1916: 126.
Lepidop ia abe ans: Masne and Muesebeck 1968: 10.
?B uesop ia abe ans: Masne and Ga cía 2002: 73.
Ma e ial examined. Holo ype, • emale: USNM no.20474, Hage s own, Md., [Wash-
ing on, Ma yland, USA] Aug. 15, 1913, ex pupa ium o ?C yp omeigenia heu is
(Walke ) in an adul o Phyllophaga in e sa (Ho n); s o ed in NMNH.
The emale o B. abe ans can easily be dis inguished om o he desc ibed B ueso-
p ia species by i s 12-me ous an ennae, whe eas males o all known B uesop ia spe-
cies ha e no been desc ibed o illus a ed in de ail (Wing 1951). Some de ails abou
B uesop ia male mo phology a e known only om he diagnosis o his genus gi en by
Masne and Ga cía (2002).
Biology. B uesop ia abe ans was ea ed om he pupa ium o a achinid ly (pos-
sibly C yp omeigenia heu is (Walke , 1849)), which was ound inside an adul June
bee le, Phyllophaga in e sa (Ho n, 1887). The associa ion o his species wi h Solenopsis
sp. was epo ed by Masne and Ga cía (2002), bu wi hou indica ion o he da e o
loca ion o he obse a ion.
Dis ibu ion. Reco ded only om he USA.
Re ision o Lepidop ia (Hymenop e a, Diap iidae) 1111
Figu e 1. B uesop ia ame icana (A), Lepidop ia masne i (B), B. abe ans, comb. no ., emale, slide o
holo ype (C, D) and B uesop ia sp.(E, F): A, B me asoma, en al iew C base o me asoma D habi us,
do sola e al iew E, F emale o B uesop ia sp. in he colony o hie an Solenopsis moles a (Say), image
Alex Wild, used by pe mission.
Rema ks. Masne and Muesebeck (1968) epo ed on he ype o Lepidop ia abe -
ans B ues and epo ed ha his species was doub ully placed in his genus. La e ,
Hugge and Masne (1983) addi ionally no ed ha i may belong o he Nea c ic ge-
nus B uesop ia Wing. This species was no men ioned again un il 2002, when Masne
and Ga cía w o e abou i as a membe o he genus B uesop ia, despi e ha no new
combina ion had been sugges ed o jus i ied. Howe e , e-examina ion o he ype spec-
V. G. Chemy e a e al. / Jou nal o Hymenop e a Resea ch 98: 1107–1126 (2025)
1112
imen o his species and some o he a ailable B uesop ia specimens, as well as all known
Lepidop ia species, led us o he same conclusion: Lepidop ia abe ans B ues should be
conside ed a membe o he genus B uesop ia, as i sha es all he cha ac e s ypical o
B uesop ia species, as ou lined in he compa a i e diagnosis o his genus abo e.
Genus Lepidop ia Kie e , 1911
Type species. Lepidop ia pedes is Kie e , by mono ypy.
Genus diagnosis. Small sized insec s (1.0–1.8 mm) wi h yellowish o almos black
body, co e ed wi h sho , e ec , e enly dis ibu ed se ae (usually emales mo e pale
and mo e densely se ose han males). Head subglobula in do sal and la e al iews,
sub iangula o globula in on al iew; eyes o emale small, i s la ges diame e
sho e han leng h o mala space; palpal o mula 4–2. Male an enna 14-me ous wi h
only A4 o A3−A4 modi ied; emale an ennae 12-me ous, wi h mo e o less ab up
4-me ous cla a o cla a indis inc ; A1 abou as long as ou ollowing an ennome es
combined; A4−A8 om subquad a e o ans e se. P opleu on, mesopleu on en al-
ly, me apleu on, la e al side o p opodeum and pe iole co e ed wi h dense se a ion
o oamy s uc u es; mesoscu um ans e se, wi hou no auli, almos la bu sligh ly
con ex an e o-medially and dep essed pos e o-medially; mesoscu ellum usually wi h
single ans e se an e io scu ella pi o i subdi ided in o wo pi s; p opodeum wi h
mo e o less dis inc median ca ina ha is no ele a ed an e io ly o ca ina absen ; plica
and pos e io plical p ocess sligh ly o mode a ely p ojec ing. Ala e, o e wing as long
as body o sligh ly sho e , wi h single cos al ein de eloped in i s basal hi d. All legs
wi h cylind ical a si, a some es abou as long as wide. Pe iole in la e al iew humped
do sally, ele a ed abo e le el o base o T2 and no b oadened pos e io ly; base o T2
dis inc ly wide han pe iole, smoo h and la ; base o S2 densely se ose, wi h wo
g oo e-like in agina ions la e ally and median con exi y be ween hem; an e io ma -
gin o S2 wi hou deep exca a ion; gas e dep essed and o oid.
Lepidop ia bica ina a sp. no .
h ps://zoobank.o g/F9D56D76-04B2-4E9E-9D8E-EA21D8292FA0
Figs 2, 3
Type ma e ial. Holo ype, • emale: Sou h Ko ea, Jeju-do (JJ), Jeju-si, Sanduk- o,
593-40, nea Temple Gwaneum es a ea (Li es ock Resea ch Ins i u e), MT, 1–24.
IX.2022, Deokseo Ku & Jeongjun Ahn legs (NIBR). Pa a ypes: same locali y as holo-
ype, 1–16.VII.2022, 16–31.VII.2022, 1♀ and 2♂ (ZISP).
E ymology. The name de i es om La in “bi-” and “ca ina” and e e s o he well-
de eloped wo ca inae on male A3 and A4.
Diagnosis. Head globula in on al iew (Fig. 2E); emale an ennae s ou , wi h
A9−A11 ans e se, A9 abou as long as, and as wide as A10 (Fig. 2F); male A3 wi h ca-
ina and wi hou ema gina ion, male A4 sligh ly enla ged, sligh ly longe and sligh ly
Re ision o Lepidop ia (Hymenop e a, Diap iidae) 1113
Figu e 2. Lepidop ia bica ina a sp. no ., emale. A habi us, la e al iew B head, mesosoma and base o
me asoma, do sal iew C me asoma, la e al iew D head and mesosoma, la e al iew E ace F an enna,
la e al iew. Scale ba s: 0.5 mm (A); 0.3 mm (B–E).
wide han A3, wi h shallow ema gina ion and memb anous, pale ca ina (Fig. 3D, E);
mesoscu ellum wi h single ans e se scu ella pi (Figs 2B, 3B); p opodeum densely
se ose do sally, wi h median ca ina; plica ha dly indica ed unde se ae (Figs 2B, D,
V. G. Chemy e a e al. / Jou nal o Hymenop e a Resea ch 98: 1107–1126 (2025)
1114
3B); pe iole weakly swollen do sally and p ojec ing upwa ds, do sally co e ed wi h
oamy s uc u e in emale and long pale se ae in male (Figs 2B, D, 3B).
Desc ip ion. Female, holo ype. Body leng h 1.5 mm, an ennae leng h 0.85 mm;
o e wing leng h 1.4 mm. Body da k b own excep pe iole and apex o me asoma
b own; egulae, an enna and legs b ownish yellow.
Head ounded, smoo h, in do sal iew as long as wide, and 1.15 imes as high
as long in la e al iew, co e ed wi h sho , e ec se ae; pos genal cushion dense; ocelli
small, inconspicuous, a anged in an almos equila e al iangle; LOL 2.5 imes as long
as diame e o on al oculus; OOL la ge, 3 imes as long as POL; compound eye small
wi h ewe han 50 omma idia; ace smoo h, epis omal sulcus indis inc ; mala space
la ge, 1.1 imes as long as heigh o eye; en o ial pi small and dis inc ; mala dis ance
abou as long as sho es dis ance be ween eye in on al iew; mandibles biden a e,
wi h apices sligh ly o e lapping.
An enna 12-me ous, co e ed wi h sho , semidecumben o e ec se ae; A1 sligh ly
comp essed and cu ed, abou 3.7 imes as long as i s g ea es wid h, smoo h, shining,
and ba e en ola e ally, co iaceous and se ose o he wise; in do sal iew A2 cylind ical
and 1.8 imes as long as wide, in la e al iew i ape ing owa ds base; A3 sligh ly na -
owe han he A2, 1.5 imes as long as wide; A4 and A5 abou as wide as A3, subequal
o each o he and abou 1.2 imes as wide as long; A6−A8 dis inc ly ans e se and be-
coming g adually wide ; A9−A12 o ming ab up 4-segmen ed cla a, i s se a ion dense
and sligh ly sho e han on o he an ennome es; A10 and All subequal, sligh ly ans-
e se; A9 ans e se and sligh ly wide han A12; A12 o oid, 1.2 imes as long as wide.
Mesosoma abou as wide as head; p ono al colla weakly de eloped (Fig. 2B, D);
p ono al shoulde s p ominen , c ossed wi h sha ply p ojec ing p ono al sup ahume al
sulcus (Fig. 2D); la e al side o p ono um smoo h, wi h sho sca e ed and e ec se a-
ion; p opleu on wi h se ae and oamy s uc u es on en al 0.3 and smoo h and ba e
do sally; mesoscu um weakly con ex an e io ly and la e ally, and sligh ly dep essed
pos e omedially, wi hou no auli and co e ed wi h sho , sca e ed e ec se ae; mes-
oscu ellum wi h ans e se and eni o m an e io scu ella pi , pi smoo h and ba e
inside; mesoscu ella disc wi h e y weak median ca ina and elonga e ugulae an e i-
o ly, co e ed wi h se e al e ec se ae la e ally; mesopleu on smoo h, la e ally co e ed
wi h sca e ed, sho , e ec se ae; mesopleu on en ally wi h sho ecumben se ae
and oamy s uc u es an e io o mid-coxa; me ano um en i ely se ose, sculp u ed
wi h sca e ed se ige ous punc u es; me ascu ellum sho , wi h low median ca ina and
weakly p ojec ing la e al ca ina; en al side o me apleu on wi h oamy s uc u e; p o-
podeum and me apleu on en i ely densely se ose, co e ed wi h sho , adp essed whi e
se ae; me apleu on and la e al side o p opodeum wi hou isible ca inae; in do sal
iew pos e io ma gin o p opodeum deeply a cua e: p opodeum sho medially and
signi ican ly longe la e ally; la e al and median ca inae o p opodeum almos pa al-
lel, na ow and hidden unde se a ion; nuchal a ea smoo h and ba e. Legs mode a ely
obus , co e ed wi h sho , e ec se ae simila o hose o body; o e ibia wi h apical
spine, i s leng h abou hal o apical wid h o ibia; all emo a wi h sho basal s alk;
hind emu 2.3 imes as long as i s g ea es wid h. Fo e wing 0.9 imes as long as body,
wi h single cos al ein eaching 0.3 o o e wing leng h.
Re ision o Lepidop ia (Hymenop e a, Diap iidae) 1115
Figu e 3. Lepidop ia bica ina a sp. no ., male. A habi us, la e al iew B mesosoma and me asoma, do sal
iew C an enna, la e al iew D, E base o an enna in do sal (D) and la e al (E) iews. Scale ba s: 0.5 mm
(A); 0.3 mm (B, C).
Me asoma. Pe iole cylind ical in do sal iew, as wide as long, densely co e ed wi h
oamy s uc u e on do sal su ace; in la e al iew, medial po ion sligh ly swollen up-
wa ds; en al side o pe iole co e ed wi h sho ecumben whi e se a ion, no p ojec -
ing en ally; gas e abou 0.9 imes as long as head and mesosoma combined, mo e
han 1.7 imes as long as i s g ea es wid h, do so en ally comp essed; T2 do sally
wi h sho , e ec , e enly dis ibu ed se ae, base o T2 la , wide han pe iole; T3−T5
smoo h and sca cely se ose; base o S2 densely se ose, wi h deep la e al g oo e and
median con exi y; basal co ne s o S2 ex ending o wa ds ei he side o pe iole beyond
base o la ge e gi e; o iposi o sho ly exse ed.
Male. Body leng h 1.4 mm. Di e ing om emale as ollows: colou o body
black, mandibles and egula b own, an ennae da k b own wi h A2 yellowish, legs yel-
lowish b own wi h coxae, emo al and ibial clubs da kened; body less densely se ose
han in emale, especially me asoma, oamy s uc u es de eloped only on p opleu on
V. G. Chemy e a e al. / Jou nal o Hymenop e a Resea ch 98: 1107–1126 (2025)
1122
Figu e 9. Lepidop ia pedes is, male. A ace and base o an ennae B head, mesosoma and base o me a-
soma, do sal iew C habi us, do sal iew D lagellum, do sal iew E pe iole, la e al iew. Scale ba s:
0.5 mm (C); 0.3 mm (A, B, D).
Rema ks. Hugge and Masne (1983) epo ed abou “an in e es ing case o si-
mul aneous pa asi ism by Lepidop ia pedes is Kie e and T ichop ia inquilina Kie e
on a achinid pupa ium in an adul may bee le, Melolon ha melolon ha Linnaeus is
known o us om Yugosla ia (specimens in Na . Museum, P aha)”. The male speci-
men om Bosnia and He zego ina men ioned abo e mos likely belongs o his g oup
o specimens, and we a e p o iding i s label da a o he i s ime he e. D Jan Macek
kindly p esen ed i o us many yea s ago and i is now s o ed in ZISP.
Re ision o Lepidop ia (Hymenop e a, Diap iidae) 1123
Key o he all species o Lepidop ia Kie e
1 Female an ennae slende wi h non-ab up cla a (Fig. 4D); male A1 swollen
medially, A3 and A4 modi ied wi h deep exca a ion (Fig. 4A, B); p ono-
um wi h dense p ono al cushion; bo de s o an e io scu ella pi indis inc ,
smoo hed (Fig. 4A, C) .................................................... L. lloydi (Fe iè e)
– Female an ennae s ou o /and wi h mo e o less dis inc 4-segmen ed cla a
(Figs 1A, 2F, 5E, 8A); male A1 cylind ical, A3 no modi ied o , a leas , wi h-
ou exca a ion; p ono al cushion educed (Figs 2D, 5C); bo de s o an e io
scu ella pi dis inc (Figs 2B, 3B, 5D, 6B, 9B) ........................................... 2
2 Pe iole s ongly swollen do sally and p ojec ing upwa ds (Fig. 9E); p ono al
shoulde s no p ojec ing in do sal iew, wi hou p ono al sup ahume al sul-
cus (Figs 8A, 9B); an e io scu ella pi subdi ided in o wo pi s, wi h aised
a ea in he middle (Figs 8A, C, 9B); head sub iangula in on al iew (Figs
8A, C, 9B); emale A9−A11 as long as wide (Fig. 8A); male A4 s ongly
enla ged, dis inc ly longe and wide han A3, wi h deep ema gina ion and
s ong, pigmen ed ca ina (Fig. 9A, D)............................ L. pedes is Kie e
– Pe iole weakly swollen do sally and p ojec ing upwa ds (Figs 2D, 3A, 5F, 6A);
p ono al shoulde s dis inc ly p ojec ing in do sal iew, wi h sha p p ono al
sup ahume al sulcus (Figs 2D, 5C); an e io scu ella pi no subdi ided in o
wo pi s (Figs 2B, 3B, 5D, 6B); head globula in on al iew (Figs 2E, 5A);
emale A9−A11 ans e se (Figs 2F, 5E); male A4 sligh ly enla ged, sligh ly
longe and almos no wide han A3, wi h mode a ely shallow ema gina ion
and memb anous, pale ca ina (Figs 3D, E, 6D, E) ......................................3
3 P opodeum e y inely se ose do sally (Figs 5D, 6B); emale pe iole pa ly
ba e o se ose do sally (Fig. 5D); emale an ennae wi h A9 dis inc ly sho e
and na owe han A10 (Fig. 5E); male A3 no modi ied (Fig. 6D, E) ..........
.......................................................................................L. masne i No on
– P opodeum densely se ose do sally (Fig. 2B); emale pe iole co e ed wi h
oamy s uc u e do sally (Fig. 2B); emale an ennae wi h A9 abou as long as
A10 and as wide as A10 (Fig. 2F); male A3 wi h ca ina (Fig. 3D, E) .............
..................................................................................L. bica ina a sp. no .
Discussion
The gene ic diagnoses o Lepidop ia gi en by Kie e (1911, 1916) we e based on a sin-
gle species, Lepidop ia pedes is. Subsequen ly, Hugge and Masne (1983) ca ied ou
a compa a i e analysis o my mecophilous species o he Hola c ic. Al hough hey did
no cla i y he gene ic diagnosis, many diagnos ic ea u es we e illus a ed and used in
hei key. The desc ip ion o he genus by Hugge and Masne (1983) is no en i ely
unambiguous. Acco ding o he key p o ided, L. pedes is has a p opodeum ha is “no
V. G. Chemy e a e al. / Jou nal o Hymenop e a Resea ch 98: 1107–1126 (2025)
1124
exca a ed pos e io ly, ... wi h dis inc median keel”, which does no align wi h he mo -
phology o he holo ype (Fig. 8A, B) and Fig. 6 in Hugge and Masne (1983). In ou
opinion, he la e illus a ed a Lepidop ia sp. wi h a pos e io ly exca a ed p opodeum,
whe eas he holo ype o L. pedes is does no ha e a median ca ina (keel). Howe e , his
disc epancy is no cu en ly signi ican , as high a ia ion in hese wo ea u es has been
epo ed in subsequen ly desc ibed species o he genus (No on 1994; cu en s udy).
The genus Lepidop ia is mos closely ela ed o he Nea c ic genus B uesop ia ( he
con used classi ica ion o L. abe ans may con i m his), and o he Palaea c ic genus
Solenopsia (Masne and Ga cía 2002). The di e ences be ween Lepidop ia and B ueso-
p ia a e gi en in he compa a i e diagnosis abo e. Two desc ibed species o Solenopsia
can be dis inguished om Lepidop ia and B uesop ia by ha ing 11-me ous an enna
wi h a 2-me ous cla a in he emales. They a e also cha ac e ised by he p esence o
a dis inc median p opodeal ca ina which is sligh ly ele a ed an e io ly and o ms a
oo h-like, sub iangula s uc u e. O he impo an diagnos ic ea u es o Solenopsia
(mos o which a e men ioned in he compa a i e diagnosis gi en o B uesop ia and
Lepidop ia abo e) a e simila o hose o species Lepidop ia o B uesop ia. Cu en ly,
we belie e ha i is no possible o easonably discuss he synonymy o hese gene a, as
addi ional mo phological, molecula , and biological da a on hese wasps a e equi ed.
I mus be conside ed ha all h ee gene a may ha e a isen independen ly h ough
con e gen e olu ion. Howe e , we would like o no e ha he li e cycles o B. abe ans
and L. pedes is mos likely a e ema kably simila .
Pho os aken by D Alexande Wild in Kansas (Fig. 1E, F) con i m once again he
connec ion be ween B uesop ia species and he an Solenopsis moles a. The B uesop ia
in hese pho os ei he belongs o he B. abe ans species, as i has 12-me ous an ennae
( wo o he desc ibed species o his genus ha e 11-me ous an ennae) and 3-segmen ed
cla a wi h sligh ly elonga ed segmen s ha a e nea ly equal in wid h and abou wice
as wide as A4–A8 (cla ame es a e ans e se and much wide in B. ame icana (B ues,
1936) and B. see e si Wing, 1951), o i belongs o an undesc ibed species o his ge-
nus ha is mo phologically close o B. abe ans. The e o e, i is likely ha all cu en ly
known species o B uesop ia a e associa ed wi h S. moles a (Wing 1951).
Un o una ely, he de ailed biology o he o he species B uesop ia, Lepidop ia and
Solenopsia is no well unde s ood, bu all species o hese h ee gene a, i known, a e associ-
a ed wi h he genus Solenopsis (B ues 1916; Lachaud 1980, 1981b, c, 1982; Lachaud and
Passe a 1982; Masne and Ga cía 2002; cu en s udy). A he same ime, we can s a e wi h
g ea con idence ha he ac ual species di e si y o hese my mecophilous wasps is signi i-
can ly g ea e han is cu en ly ecognized. Thus, i canno be discoun ed ha addi ional
disco e ies will elimina e he enuous bounda ies be ween hese my mecophilous gene a.
Acknowledgmen s
We a e e y hank ul o M Joseph Monks (NHML) and D Eka e ina Tselikh (ZISP) o
p o iding c ucial in o ma ion and images o he ype ma e ial o Lepidop ia lloydi. We also
wish o hank D . Ma Bu ing on (NMNH) o he nice pho os o B. abe ans slide. Ou
Re ision o Lepidop ia (Hymenop e a, Diap iidae) 1125
app ecia ion goes o D . Ma ia Ta ano (MCSN) o p o iding he oppo uni y o examine
he ypes o L. pedes is. We hank D . Alexande Wild (The Uni e si y o Texas a Aus in,
USA) o pe mission o use he pho os o B uesop ia sp. Special hanks go o D Jean-Paul
Lachaud (El Colegio de la F on e a Su , Che umal, Mexico) o eading he i s d a o
he manusc ip and o his use ul sugges ions and addi ional li e a u e, which enabled
us o signi ican ly imp o e his a icle. We would also like o hank D . Elijah Talamas
(Flo ida Depa men o Ag icul u e and Consume Se ices, USA) o his aluable com-
men s on he i s d a o he manusc ip and his comp ehensi e suppo . This wo k was
suppo ed by a g an om he Na ional Ins i u e o Biological Resou ces (NIBR), unded
by he Minis y o En i onmen (MOE) o he Republic o Ko ea (NIBR202502202)
and pa ially suppo ed by Russian S a e Resea ch P ojec (No. 125012901042-9).
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