521
T iassic io acyphids shed ligh on he ena ion o c own
Thysanop e a and hei s em ela i es (Condylogna ha:
Holo hysanop e a)
Ma hieu Bode au1,*, Ma eo Mon agna2,*, Fabio Magnani3, Giulia Magoga2, Alexey Shmako 4,
Michael S. Engel5,6, Pa icia Nel1, And é Nel1
1 Ins i u de Sys éma ique, É olu ion, Biodi e si é (ISYEB) Muséum na ional d’His oi e na u elle, CNRS, So bonne Uni e si é, EPHE-PSL,
Uni e si é des An illes, CP50, 57 ue Cu ie , 75005 Pa is, F ance
2 Depa men o Ag icul u al Sciences, Uni e si y o Naples Fede ico II, Piazza Ca lo di Bo bone 1, 80055 Po ici, I aly
3 Museo can onale di s o ia na u ale, Via C. Ca aneo 4, 6900 Lugano, Swi ze land
4 Labo a o y, Palaeon ological Ins i u e, Russian Academy o Sciences, P o soyuznaya s ., 123, Moscow 117997, Russia
5 Di ision o In e eb a e Zoology, Ame ican Museum o Na u al His o y, 200 Cen al Pa k Wes , New Yo k, 10024 New Yo k, USA
6 Pe e i Museum Founda ion, Baumschulweg 13, 6045 Meggen, Swi ze land
* These au ho s con ibu ed equally.
h ps://zoobank.o g/CD5A5819-21A1-4256-A2B2-4007305EABCE
Co esponding au ho s: Ma hieu Bode au ([email p o ec ed]), Ma eo Mon agna ([email p o ec ed]), And e Nel ([email p o ec ed])
Recei ed 08 June 2025
Accep ed 12 Sep embe 2025
Published 10 Oc obe 2025
Academic Edi o Mónica Soló zano-K aeme
Ci a ion: Bode au M, Mon agna M, Magnani F, Magoga G, Shmako A, Engel AS, Nel P, Nel A (2025) T iassic io acyphids shed ligh on he
ena ion o c own Thysanop e a and hei s em ela i es (Condylogna ha: Holo hysanop e a). A h opod Sys ema ics & Phylogeny 83: 521–530.
h ps://doi.o g/10.3897/asp.83.e161465
Abs ac
We epo he disco e y o he i s T iassic ‘shielded h ips’ o he amily Io acyphidae, ep esen ed by wo species: T iassocypha
p ima gen. e sp. no . and T. secunda sp. no . The species a e desc ibed and illus a ed, and compa isons a e made wi h o he
s em-Thysanop e a lineages. The unique wing ena ion o Io acyphidae and ela ed amilies is ein e p e ed in he ligh o hese new
ossils. The ena ion pa e n suppo s he diagnosis o he amily, as well as o highe clades wi hin he b oade lineage ha includes
bo h h ips and hei ex inc ela i es (in he o al g oup Holo hysanop e a, o me ly Th ipida), and con ibu es o cla i ying he
sys ema ics o h ips.
Keywo ds
Ace ca ia, classi ica ion, ‘Lophioneu ida’, Middle T iassic, Mon e San Gio gio, wing ena ion
1. In oduc ion
Ex an h ips (Thysanop e a) a e minu e insec s wi h ex-
cep ionally na owed wings, wi h educed ena ion and
inged ma gins. The mou hpa s a e asymme ical and,
like Hemip e a, o m a sucking appa a us o eed on plan
luids o ungal spo es (Heming 1993). Al hough mode n
h ips a e easily cha ac e ized and dis inc i e, he e is a
ich ossil eco d o s em g oups (‘p o o h ips’) o ming a
g ade leading o c own-Thysanop e a and ex ending back
o he Paleozoic. Mos o hese s em g oups a e di e se
du ing he Ea ly Mesozoic ( om he T iassic h ough
he Ea ly C e aceous), wi h mo e mode n-appea ing
h ips domina ing om he Ea ly C e aceous o he la e
A h opod Sys ema ics & Phylogeny 83, 2025, 521–530 | DOI 10.3897/asp.83.e161465
Copy igh Ma hieu Bode au 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.
Bode au M e al.: T iassic io acyphids
522
Cenozoic, and la gely p ese ed as inclusions in ambe
(PBDB: h ps://paleobiodb.o g). The s em-g oup lineag-
es a e, howe e , c ucial o unde s anding he cha ac e
ans o ma ions ha p og essi ely ga e ise o he sui e o
synapomo phies we oday unde s and as dis inguishing
mode n h ips om hei many ela i es (Nel e al. 2014).
One o he mo e di e se g oups among s em-Thysan-
op e a is he amily Lophioneu idae Tillya d, 1921 (he e-
in classi ied as comp ising wo dis inc amilies, below).
These ossils, p ese ed la gely as comp essions bu also
including some inclusions in mid-C e aceous ambe , a e
i al o elucida ing ea ly ela ionships and homologies
wi hin Condylogna ha. He e we epo he disco e y o
a lineage o s em-Thysanop e a om he Middle T iassic
o Swi ze land, p e iously known only om mid-C e a-
ceous ambe o no he n Myanma . These insec s signi i-
can ly expand ou knowledge o he “Lophioneu ida”,
ex ending hei dis ibu ion in o he T iassic o Eu ope,
and allowing us o p o ide a ein e p e a ion o ena-
ional homologies ac oss li ing and ossil ela i es o he
Thysanop e a.
2. Ma e ial and me hods
Mon e San Gio gio’s Middle T iassic ca bona e succes-
sion is a UNESCO Wo ld He i age si e (Swi ze land-I -
aly) amous o i s well-p ese ed ossil ishes and ma-
ine ep iles (e.g., Rieppel 2019). In ecen yea s, Mon e
San Gio gio has gained signi ican impo ance o ossil
insec s, e ealing an excep ionally di e se and well-p e-
se ed assemblage (Mon agna e al. 2024a, 2024b).
The unique sedimen a y condi ions we e he esul o a
long- e m ansg ession and dysoxic o anoxic bo om
wa e s (e.g., Be nasconi 1994; S ocka e al. 2013). Fos-
sils we e collec ed du ing he exca a ion led by he Mu-
seo can onale di s o ia na u ale (MCSN) in 2020 a he
Val Ma a VM 12 si e (DMS geog aphical coo dina es:
45°53’26’’N; 8°56’49’’E; alley o he Gaggiolo Ri e ,
Fig. 1A), wes o Me ide illage (Mon e San Gio gio,
Swi ze land) (Mon agna e al. 2024a). The ossil-bea -
ing s a a, which p ese e he insec auna, belong o he
uppe “Kalkschie e zone” (Senn 1924), da ed o he la e
Ladinian (~ 239 Ma; S ocka e al. 2012) (Fig. 1B). These
s a a consis o al e na ing hin-bedded lamina ed lime-
s ones and ma ly limes ones, wi h subo dina e ma ls ones
and calca eous ma ls ones, e lec ing a shallow lagoon
en i onmen wi h seasonal salini y a ia ions (Tin o i
1990). Du ing he Ladinian, he egion expe ienced mon-
soonal ci cula ion (P e o e al. 2010), leading o seasonal
a ia ions and densi y s a i ica ion in he hype saline la-
goon (Fu e 1995).
Figu e 1. Loca ion o Mon e San Gio gio (I aly-Swi ze land) UNESCO Wo ld He i age and s a ig aphic sec ion o he Middle
T iassic sedimen s. A Map wi h Mon e San Gio gio, showing he ca bona e Anisian–Ladinian sequence, he s a indica es he loca-
ion o he Val Ma a VM12 exca a ion si e. B S a ig aphic column o he Mon e San Gio gio, he black a ow indica es he VM12
s a a ha yielded he he e desc ibed specimens. Commissione scien i ica ansnazionale (a e S ocka e al. 2012).
A h opod Sys ema ics & Phylogeny 83, 2025, 521–530 523
The ossils we e p epa ed using a s eel needle unde a
s e eomic oscope, and high-quali y images we e cap u ed
using an Olympus SZX 12 mic oscope coupled wi h an
Olympus UC50 came a and KEYENCE VHX-X1F dig-
i al mic oscope wi h VHX-7020 came a wi h VH-Z20T
zoom lens. Pos -p ocessing o images was pe o med us-
ing Adobe Pho oshop CS2. Vena ion e minology ollows
Nel e al. (2012) o he Ace ca ia, modi ied by Schubnel
e al. (2020) o conside he p esence o a pos cubi al ein
(see also al e na i e hypo heses in in e p e a ion p esen -
ed in he Discussion, below). Vein names be ween b ack-
e s, e.g., ‘CuA’ o ‘M’, a e hose p oposed by Cumming
e al. (2024); hose wi hou b acke s a e newly accep ed
he ein. Abb e ia ions o eins and c oss eins a e:
A anal ein; C cos a; Cu cubi us; CuA cubi us an e io ;
CuP, cubi us pos e io ; cua-cup specialized c oss ein
be ween CuA and CuP; M median ein; MA media an-
e io ; MP media pos e io ; PCu pos cubi us; R adius;
RA adius an e io ; RP adius pos e io ; ScP subcos a
pos e io .
3. Resul s
Sys ema ic palaeon ology
Coho Ace ca ia Bö ne , 1904
3.1. Supe o de Holo hysanop e a Engel,
Mon agna, Bode au, P. Nel, & A. Nel
no .
(Ca boni e ous Wes phalo h ipidesidae P. Nel, Aza , P okop, Roques,
Hodebe , & A. Nel, 2012 o c own-Thysanop e a)
(= “Th ipida Fallén, 1814” sensu P. Nel e al. 2014)
Diagnosis. Le mandibula s yle , igh absen ; pie cing
lacinial s yle s; p e a sus wi h e e sible a olium; p e a sal
claws educed in adul s; wings wi h inge o long se ae
and CuA simple.
Included clades. Wes phalo h ipidesidae, Lophioneu-
op e a (spec al and shielded h ips), Be a hysanop e a
Engel (Zo opsocidae Tillya d, 1935, be a h ips), Thysa-
nop e a ( ue h ips).
3.2. Clade Lophioneu op e a Engel, Mon-
agna, Bode au, P. Nel, & A. Nel no .
(= “Lophioneu ida Tillya d, 1921” sensu auc o um)
Rema ks. We ejec he name “Lophioneu ida” as ap-
plied o his clade on he g ounds ha i and o he names
like i abo e he amily g oup, ep esen an a emp o
impose a ogue nomencla u al sys em by way o changes
o he P inciple o Coo dina ion ela i e o he ICZN
amily-g oup se ies. In b ie , he e is no such hing as
“Lophioneu ida Tillya d, 1921” as names abo e he am-
ily g oup a e no coo dina e wi h he amily-g oup se ies
o names. Tillya d (1921) es ablished a amily-g oup
name, Lophioneu idae, based on he ype genus Lophio-
neu a Tillya d, 1921 and he e o e his name is coo di-
na e h oughou he ICZN- egula ed amily g oup (ICZN
1999), bu no u he h ough names ou side o he am-
ily g oup. E o s o ei he ypi y and/o make names
abo e he le el o he amily g oup coo dina e wi h hose
o amilies (e.g., Rohdendo 1977; Rasni syn 1982)
ha e ne e been adop ed no sanc ioned by he ICZN
(1999). Acco dingly, such e o s o make hem pa o
he na u al coo dina ion and ypi ica ion o amily-g oup
names a e ope a ing ou side o he bounds o he ICZN
(1999). We he e o e ejec he use o “Lophioneu ida”
and o he ogue a emp s o ci cum en he ICZN and
o ce ypi ied coo dina ion o amily-g oup names wi h
hose clea ly ou side o he ICZN- egula ed sys em o
names. The same p oblem applies o he amily-g oup
name Th ipidae Fallén, 1814 as ex ended o a supe o de
as “ Th i p ida”.
Diagnosis. Fo ewing wi h CuA e-eme ging o hogonally
om R+M+CuA.
Included amilies. Lophioneu idae Tillya d, 1921 (spec-
al h ips) and Io acyphidae Shche bako , Bashkue , &
Shmako , 2024 (shielded h ips).
3.3. Family Io acyphidae Shche bako ,
Bashkue , & Shmako , 2024, s a .
no .
Io acyphinae Shche bako , Bashkue , & Shmako , 2024: 416. Type ge-
nus: Io acypha Shche bako , Bashkue , & Shmako , 2024.
Diagnosis. These insec s a e well cha ac e ized by he
synapomo phic p esence in he o ewing o la ge median
cells and s ong angles in he cou ses o R and M, all ab-
sen in he Lophioneu idae (sensu Nel e al. 2012, 2014)
and Mound h ipidae Nel, Aza , & Nel, 2007. These h ee
amilies ha e a CuA e-eme ging i s om R+M+CuA
bu CuA is no o hogonal o R+M in he Mound h ipidae
e sus o hogonal in Io acyphidae and Lophioneu idae
(see below).
Included gene a. Bu macypha Zhe ikhin, 2000 (mid-
C e aceous Kachin ambe ); Io acypha Shche bako ,
Bashkue , & Shmako , 2024 (mid-C e aceous Kachin
ambe ); Re ip e a Cumming, Engel, Lian, & Uli zka,
2024 (mid-C e aceous Kachin ambe ); T iassocypha
gen. no . (Middle T iassic, Mon e San Gio gio, Swi ze -
land).
Bode au M e al.: T iassic io acyphids
524
3.4. T iassocyphinae sub am. no .
h ps://zoobank.o g/BC3F6D67-3CC8-41E0-BC21-
1E75C7D7B857
Type genus. T iassocypha gen. no .
Diagnosis. Two closed cells be ween R and M and a
closed cell be ween he b anches o M.
3.5. T iassocypha gen. no .
h ps://zoobank.o g/F19D8697-6BE5-473B-A616-09FB13D-
9CC0F
Type species. T iassocypha p ima sp. no .
E ymology. The name is an a i icial combina ion o T i-
assic, e e ing o pe iod om which he ossils o igina e,
and he Ancien G eek adjec i e κῡφός (kūphós, mean-
ing, “ben o wa d” o “cu ed”; eminine κῡφή). The
gende o he name is eminine.
Diagnosis. Fo ewing ScP e-eme ging om R as a sho ,
s aigh einle pe pendicula o C; a closed cell be ween
MA and MP; wo longi udinal cells be ween R and M/
MA; CuP+PCu well sepa a ed om A.
Included species. T iassocypha p ima sp. no . and T. se-
cunda sp. no ., bo h om he T iassic o Mon e San
Gio gio.
3.6. T iassocypha p ima sp. no .
h ps://zoobank.o g/393685B4-0B19-4461-93FA-
78D75D123F11
Figu es 2A, B, D, 3A
Ma e ial. Holo ype, MCSN 8715 (a body wi h wo o e-
wings and one hind wing, legs, and one an enna appa -
en ly comple e), s o ed in Museo can onale di s o ia na u-
ale, Lugano, Swi ze land.
Locali y and ho izon. Uppe Kalkschie e zone membe
o Me ide Limes one ( ansi ion in e al o A chelaus and
Regoledanus Ammonoid Zones, uppe Ladinian), Middle
T iassic; VM12 si e (DMS geog aphical coo dina es:
45°53’26’’N; 8°56’49’’E), Val Ma a nea Me ide, Mon e
San Gio gio, Swi ze land.
E ymology. The speci ic epi he is he La in adjec i e
p īmus, meaning, “ i s ” o “ea ly”, and e e s o his as
he i s desc ibed species o he genus.
Diagnosis. Fo ewing cha ac e s only: ma-mp longe han
-ma; dis al-mos cell be ween R and MA h ee imes as
long as wide.
Desc ip ion. Body 1.3 mm long, p esence o long mac-
ose ae on he ho ax and some wing eins. Head: head
appa en ly o hogna hous o p ogna hous, wi h a ela-
i ely long buccal cone; compound eyes ounded wi h
diame e 0.1 mm; an enna elonga e, wi h lagellome es
elonga e, dis al mos lagellome es sho e han o h-
e lagellome es, scape la ge, pedicel appa en ly small.
Tho ax: ho ax 0.5 mm long; all legs p ese ed and o
simila shapes, appa en ly only wo a some es, basal one
ligh b own and dis al one da k b own. Abdomen: abdo-
men elonga e and b oad, 0.7 mm long, 0.55 mm wide;
ce ci absen . Fo ewing ena ion: o ewing 1.6 mm long,
0.6 mm wide, pea -shaped, ounded apically, na ow
basally; a common s em ScP+R+M+CuA om which
CuA sepa a es 0.2 mm om wing base, pe pendicula o
ScP+R+M and sho be o e cua-cup e mina es in o ein;
cua-cup 0.07 mm long, s aigh , aligned wi h dis al pa
o CuA; CuA simple and cu ed dis ally, ending a wing
ma gin 0.5 mm om wing base; CuP+PCu s aigh , well
sepa a ed om A; A sho , weakly cu ed; ScP e-eme g-
ing om R 0.8 mm om wing base, pe pendicula o C
and R, s aigh , 0.12 mm long; M sepa a ing om R 0.1
mm dis al o base o CuA; s em o M cu ed, 0.25 mm
long, sepa a ing in o MA and MP and delimi ing a b oad
elonga e closed cell, 0.5 mm long, 0.2 mm wide; bo h
MA and MP wi h a s ong angle dis ally and ending on
pos e io ma gin a igh angle; a long ma-mp pe pendic-
ula o MA and MP; wo elonga e, b oad cells be ween R
and MA, sepa a ed by -ma pe pendicula o R and MA,
basal-mos cell 0.35 mm long, 0.07 mm wide; dis al-mos
cell 0.6 mm long, 0.2 mm wide; R o ming an angle dis al
o ScP and s aigh dis ally; sepa a ing in o RA and RP
0.45 mm dis al o ScP; RA s aigh , sho , a he oblique,
nea ly o hogonal wi h RP; RP o ming a s ong angle
and eaching wing apex; s aigh , elonga e p-ma, 0.1 mm
long.
3.7. T iassocypha secunda sp. no .
h ps://zoobank.o g/A658AE57-4368-406E-9706-
25E54BA181E2
Figu es 2C, E, 3B
Ma e ial. Holo ype MCSN 8689 (a body wi h wo o e-
wings and agmen s o legs), s o ed in Museo can onale
di s o ia na u ale, Lugano, Swi ze land.
Locali y and ho izon. Uppe Kalkschie e zone membe
o Me ide Limes one ( ansi ion in e al o A chelaus and
Regoledanus Ammonoid Zones, uppe Ladinian), Middle
T iassic; VM12 si e (DMS geog aphical coo dina es:
45°53’26’’N; 8°56’49’’E), Val Ma a nea Me ide, Mon e
San Gio gio, Swi ze land.
E ymology. The speci ic epi he is he La in nume al se-
cundus, meaning “second”, and e e s o his as he sec-
ond desc ibed species o he genus.
A h opod Sys ema ics & Phylogeny 83, 2025, 521–530 525
Figu e 2. Habi us and o ewing de ails o T iassocypha gen. no . (Lophioneu op e a: Io acyphidae) species. A, B, D T iassocypha p i-
ma sp. no ., holo ype MCSN 8715 imp in (A, D) and coun e imp in (B). C, E T iassocypha secunda sp. no . holo ype MCSN 8689.
Figu e 3. Fo ewing econs uc ions o T iassocypha gen. no .
(Lophioneu op e a: Io acyphidae) species. A T iassocypha p i-
ma sp. no ., holo ype MCSN 8715. B T iassocypha secunda sp.
no . holo ype MCSN 8689. The small ed a ow indica es he
cua-cup c oss ein. Abb e ia ions: A = anal ein; Cu = cubi us;
CuA = cubi us an e io ; CuP = cubi us pos e io ; M = media
ein; MA = media an e io ; MP = media pos e io ; PCu = pos -
cubi us; R = adius; RA = adius an e io ; RP = adius pos e io ;
ScP = subcos a pos e io .
Bode au M e al.: T iassic io acyphids
526
Diagnosis. Fo ewing cha ac e s only: ma-mp as long as
-ma; dis al-mos cell be ween R and MA wo imes as
long as wide.
Desc ip ion. Body 1.4 mm long, p esence o long mac-
ose ae on he head, ho ax, and some wing eins. Head:
head appa en ly o hogna hous o p ogna hous, wi h
a ela i ely long buccal cone; compound eyes ounded
wi h diame e 0.1 mm; [an enna no p ese ed]. Tho ax:
0.37 mm long, 0.4 mm high; legs pa ly p ese ed. Ab-
domen: b oad, 0.9 mm long, 0.7 mm wide; ce ci absen .
Fo ewing ena ion: o ewing 1.7 mm long, 0.8 mm
wide, pea -shaped, ounded apically, na ow basally; a
common s em ScP+R+M+CuA; CuA, cua-cup, and CuP
no isible; ScP e-eme ging om R 0.8 mm om wing
base, pe pendicula o C and R, s aigh , 0.15 mm long;
M sepa a ing om R 0.6 mm dis al o base o CuA; s em
o M cu ed, 0.5 mm long, sepa a ing in o MA and MP
and delimi ing a b oad elonga e closed cell, 0.4 mm long,
0.19 mm wide; bo h MA and MP wi h a s ong angle
dis ally and ending on pos e io ma gin a igh angle; a
long ma-mp pe pendicula o MA and MP; wo elonga e,
b oad cells be ween R and MA, sepa a ed by -ma pe -
pendicula o R and MA, basal-mos cell 0.4 mm long,
0.15 mm wide; dis al-mos cell 0.46 mm long, 0.23 mm
wide; R o ming an angle dis al o ScP and s aigh dis al-
ly, sepa a ing in o RA and RP 0.5 mm dis al o ScP; RA
s aigh , sho , a he oblique, nea ly o hogonal wi h RP;
RP o ming a s ong angle, eaching wing apex; s aigh ,
elonga e p-ma, 0.12 mm long.
4. Discussion
4.1. Un angling Lophioneu op e a
sys ema ics and wing ena ion
homologies
Since bo h ossils exhibi ema kably simila o ewing
ena ion, i is mos app op ia e o place hem wi hin he
same genus. They di e in he leng hs o he cell be ween
ScP and RA, ha o he dis al cell be ween R and MA,
and o he cell be ween b anches o M. Collec i ely, hei
ena ion is qui e pa icula , wi h la ge polygonal cells and
main eins o ming p onounced angles. This kind o e-
na ion is encoun e ed in some o he Ace ca ia, such as
Sphae opsocidae (Psocodea) o some Hemip e a (e.g.,
Coleo hyncha, He e op e a: Schizop e idae). Rega d-
less, he wo new ossils di e g ea ly om Hemip e a in
he simple CuA, a synapomo phy o he Holo hysanop-
e a, e sus wi h an a eola pos ica p esen in Hemip e a
and he Te as igmop e a (= ‘Pe mopsocida’, Pe mopso-
cop e a) (Huang e al. 2016). The Sphae opsocidae ha e
an en i ely di e en pa e n o ena ion (see Smi he s
1990: ig. 121; G imaldi and Engel 2006), excep o he
ea ly C e aceous Asphae opsoci es neli Aza , Engel, &
G imaldi, 2010. The la e genus sha es wi h he new os-
sils eins ScP and RA eeme ging as sho einle s om
ScP+R, a simple CuA, and h ee la ge polygonal cells
medially in he o ewing, bu no homologous, wi h wo
be ween R and M; and he hi d one be ween M and CuA
(Aza e al. 2010: Fig. 2) a he han be ween MA and MP
in he wo new ossils.
The o ewing ena ions o he wo new ossils a e
mos simila o hose o he Io acyphidae (o iginally p o-
posed as a sub amily bu he ein acco ded amilial ank),
a small clade cu en ly placed in he holo hysanop e an
lineage as ela ed o Lophioneu idae, and which comp is-
es he gene a Io acypha, Bu macypha, and Re ip e a, all
om mid-C e aceous Kachin ambe . Zhe ikhin (2000),
Cumming e al. (2024), Shche bako e al. (2024), and
Shche bako and Shmako (2024) did no jus i y he a -
ibu ion o he Io acyphidae o Holo hysanop e a and i s
subclade Lophioneu op e a. Nel e al. (2012) discussed
he posi ion o Bu macypha and sugges ed ha i could
belong o he Holo hysanop e a based on he simple o e-
wing CuA bu also ha i is di icul o de e mine he ex-
ac na u e o he ena ion and body s uc u es in he a ail-
able ma e ial o ha genus. Io acypha and Re ip e a ha e
elonga e mou hpa s wi hou any isible mandibles, un-
like Psocodea and Te as igmop e a, sugges ing ha hese
a e in e nalized. Addi ionally, he dis al pa o he lab um
o Re ip e a b ennae Cumming, Engel, Lian, & Uli zka,
2024 has a lab al pad pa ly enclosed in he pai o elon-
ga e pa aglossae o he labium (Cumming e al. 2024: ig.
4D). This complex s uc u e p obably guided he s yle s
o he laciniae and mandible(s), in he same manne as in
ex an h ips (Chis olm and Lewis 1984: Fig. 1D,F; Hun -
e and Ullman 1989: ig. 1B). The p esence o a lab al pad
is also a es ed in he lophioneu op e an Mound h ipidae
Mound h ips bea i icus P. Nel, Aza , & A. Nel, 2007 (P.
Nel e al. 2025), and ep esen s p obably a synapomo phy
o he Holo hysanop e a. Un o una ely, he new ossils
do no show clea ly he head s uc u es, bu bo h seem o
ha e a buccal cone (Fig. 2), a lab al pad, and no s ong
mandibles ha would be isible la e ally, and collec i ely
hese cha ac e s a es sugges a posi ion in Holo hysan-
op e a.
Cumming e al. (2024: ig. 3) in e p e ed he o ewing
ena ion o Io acyphidae as ollows: CuA and M a e ba-
sally used wi h R and dis ally eeme ge om i , and he e
is a specialized cua-cup be ween CuP and R+M+CuA, o
M+CuA, o CuA (bo h synapomo phies o he Ace ca ia,
sensu A. Nel e al. 2012). Based on a compa ison o he
cu en ossils, Io acyphidae, Lophioneu idae, and o he
holo hysanop e ans we o e a di e en in e p e a ion o
some de ails as pe ain he in e p e a ions o Cumming e
al. (2024), Shche bako e al. (2024), and Shche bako
and Shmako (2024) ega ding CuA, CuP, and 1A.
I we ollow he wing ena ion pa e n o he Neop e a
as p oposed by Schubnel e al. (2020), a supplemen a-
y longi udinal main ein is p esen be ween he cubi al
and he anal eins, iz. he pos cubi al ein PCu. In he
Ace ca ia, his ein is especially impo an as i is pa ly
o o ally used wi h CuP. In he pho og aph o o ewing
base o Io acypha zhe ikhini Shche bako , Bashkue , &
Shmako , 2024 ( igu ed in Cumming e al. 2024: igs
1D, 2A, 3D) and in pho og aphs o he same species in
A h opod Sys ema ics & Phylogeny 83, 2025, 521–530 527
Shche bako e al (2024: igs 1–3), a s aigh , whi e (de-
scle o ized?), and oblique s uc u e is clea ly isible in
he a ea be ween ‘CuP’ and ‘1A’. This s uc u e me ges a
he ex eme wing base wi h he ein e med ‘1A’ by hese
au ho s. This ‘1A’ eme ges om a cu ed basi enale bul-
la ha has he ypical shape o ha o he ein PCu. The
ein in e p e ed as he basal pa o ‘CuP’ anishes in he
ex eme basal pa o he a ea be ween R+M+CuA and
he ‘s aigh , whi e (descle o ized?), and oblique s uc-
u e’. Gi en ha i does no ex end mo e p oximally han
he bulla, we he e o e do no belie e ha his is CuP.
Acco dingly, we ein e p e he ena ion o I. zhe ikhini
as ollows: he e is a basal usion o R wi h M and wi h
CuA in a common s em; CuP is independen o he s em
R+M+CuA and co esponds o he ‘s aigh , whi e (des-
cle o ized?), and oblique s uc u e’; he ein in e p e ed
as ‘1A’ is PCu and is basally ouching CuP, as in many
Neop e a, and especially Ace ca ia; he anal ein is e-
duced o he basal pa o he pos e io ma gin o he
wing, no e ha in Neop e a he ex eme base o he anal
ein is well sepa a ed om he base o Cu and o PCu;
cua-cup is he ein in e p e ed as he basal pa o ‘CuP’
by Cumming e al. (2024); CuA eme ges i s and sepa-
a ely om R+M. CuA eme ging i s and o hogonal o
R+M is in e p e ed as a pu a i e synapomo phy o he
clade Lophioneu op e a (‘Lophioneu ida’ sensu P. Nel e
al. 2014: ig. 11), his CuA is simple; he ein in e p e ed
as ‘CuA’ by Cumming e al (2024: ig. 2B) is likely M, i
is no pe pendicula o he adial ein bu ins ead oblique.
I s in e p e a ion as ‘CuA’ is no cong uen wi h an a -
ibu ion o hese insec s as lophioneu op e ans; he ein
in e p e ed as ‘M’ in Cumming e al. (2024: ig. 2B) is RP
and he ein in e p e ed as ‘MP’ is MA, while he ein
in e p e ed as ‘MP+CuA’ is likely MP. I is qui e unusual
ha a ein ‘MP’ would ha e a e e sed cou se, di ec ed
owa ds he wing base; bu i in e p e ed as MA, hen i s
di ec ion is mo e ‘usual’.
The ac ha he cha ac e ‘CuA eme ges i s and
sepa a ely om R+M’ is a pu a i e synapomo phy o he
clade Lophioneu op e a is no ob ious and could be con-
adic ed by i s p esence in he Te as igmop e a, sis e
g oup o he Holo hysanop e a + Hemip e a (Huang e
al. 2016). This would sugges ha i is a plesiomo phy
o much mo e likely independen ly de i ed and simply a
con e gence. The ela i e posi ions o he e-eme gences
o CuA and M a e highly a iable wi hin he Hemip e a:
basal o, a he same poin as, o dis al o M+CuA. The
cha ac e a leas aci ly suppo s Lophioneu op e a de-
spi e i s mo e homoplas ic occu ence in a mo e global
phylogene ic dis ibu ion.
Based on ou ein e p e a ion (see Fig. 4), in R. b en-
nae he ein ‘CuP’ sensu Cumming e al. (2024: ig. 2C)
should be CuA me ging dis ally wi h CuP+PCu; he e is a
sho common s em M+CuA sepa a ing om R, and om
which CuA eme ges pe pendicula ly; ‘CuA’ would be M,
while ‘M’ should be a c oss ein be ween MA and RP;
‘Rs+M’ should be RP, ‘MP+CuA’ is MP, and ‘MP’ would
be MA. Likewise, in Bu macypha longico nis Zhe ikhin,
2000, he e would be a longe common s em o M+C-
uA han in Re ip e a, ‘Cu’ should be M+CuA, sepa a ing
in o CuA (‘CuP’), pe pendicula o M (‘CuA’), his CuA
me ges dis ally wi h CuP+PCu+1A (pos e io ma gin o
wing), M sepa a es in o MA (‘MP’) and MP (‘MP+CuA’),
‘M’ is a c oss ein be ween MA and RP; ‘Rs+M’ would
be RP. Fo ewing ScP is used basally wi h R+M+CuA
in Re ip e a, Io acypha, and T iassocypha and e-eme g-
es dis ally as a cu ed ein in he wo o me and as a
s aigh einle in he la e . In Bu macypha, he ein in-
e p e ed as ‘RA’ is in exac ly a he same posi ion as in
T iassocypha, suppo ing he hypo hesis ha i is also he
e-eme gence o ScP and ha ScP basally is used wi h
R+M+CuA a wing base (and no sepa a ed om i ).
Ou cu en ein e p e a ion o he ena ion o hese
insec s is cong uen wi h he p esence o a CuA ha is
simple (synapomo phy o Holo hysanop e a), and pe -
pendicula o R+M o o M (pu a i e synapomo phy o
he Lophioneu op e a, bu see below), and wi h he p es-
ence o a PCu (a cha ac e o he wing ena ion g ound-
Figu e 4. Kachin ambe Io acyphidae, ein e p e a ion o o e-
wing ena ion. A Io acypha zhe ikhini Shche bako , Bashkue ,
& Shmako , 2024. B Bu macypha longico nis Zhe ikhin, 2000;
C Re ip e a b ennae Cumming, Engel, Lian, & Uli zka, 2024
( ed awn a e Cumming e al. 2024). Abb e ia ions as in Fig. 3.
Bode au M e al.: T iassic io acyphids
528
plan o Neop e a). The educ ion o CuA o a sho ein
in Io acypha and Bu macypha is also cong uen wi h he
gene al endency o he educ ion o his ein in he Holo-
hysanop e a, CuA pe haps being o ally absen in ex an
Thysanop e a (bu see discussion, below, on he absence
o CuA in c own-Thysanop e a). Rega dless, ou in e -
p e a ion o he ena ion suppo s an a ibu ion o hese
clades o he Holo hysanop e a and o he Lophioneu op-
e a. The new ossils di e om Io acypha, Bu macypha,
and Re ip e a in he p esence o wo closed cells be ween
R and M and a closed cell be ween he b anches o M,
e sus only one cell be ween R and M and no closed cell
be ween he b anches o M (Fig. 4).
4.2. P esence o absence o an inde-
penden e-eme gence o CuA in
c own-Thysanop e a
Nel e al. (2012) in e p e ed he ena ion o he ex an
h ips as ha ing no CuA e-eme ging om R+M+CuA
due o he exceedingly na ow wing and educed ena-
ion. Ne e heless, ano he in e p e a ion is possible. In
he o ewing o ex an Aeolo h ipidae, he h ips amily
wi h he mos comple e ena ion and a wingspan sui able
o s udying basal ena ion s uc u es, CuP is used ba-
sally wi h PCu in a la ge basi enale bulla, common in
all neop e an lineages (Schubnel e al. 2020), and dis ally
sepa a es om i in a longi udinal ein ‘1’ ha makes a
s ong angle and p oceeds o he pos e io ma gin as a
sho einle (Fig. 5). A ha poin , he e is a ein ha
con inues he basal pa o CuP and ends in o a ein ‘2’
ha pos e io ly eme ges om R+M+CuA (Fig. 5). The
ein ‘1’ has he same pa e n o o namen a ion as ein
‘2’, as equen ly happens o he cua-cup o Hemip e a.
One in e p e a ion is ha ‘1’ is cua-cup, and hen, ein ‘2’
is M+CuA in which cua-cup e mina es. Unde his in e -
p e a ion, CuA would me ge wi h M, and he i s sho
einle be ween M+CuA and he pos e io wing ma gin
could be CuA ( s. M2 as p oposed by P. Nel e al. 2012).
Acco ding o his, M would be simple in c own-Thysan-
op e a and, ollowing his hypo hesis, he synapomo phy
o c own-Thysanop e a would be ha M is simple s CuA
absen as p e iously p oposed by P. Nel e al. (2012). In
he ep esen a i es o s em-g oup Thysanop e a, M is
o ked and CuA emains p esen . This in e p e a ion is
ce ainly consis en wi h all obse a ions.
The hypo hesis abou a e-eme gence o CuA om
M+CuA is en a i e because i is no possible o dis in-
guish he shape and colo be ween he wo dis al b anches
o M+CuA. In Th ipidae he e is no sho ein be ween
M+CuA and he pos e io wing ma gin, hus CuA (o M2)
is absen as an independen ein. Fo now, we en a i ely
conside he hypo hesis iden i ying a cua-cup ein and
a e-eme gence o M+CuA om R+M+CuA as s onge
gi en he o namen a ion o he eins and he posi ion
and na u e o cua-cup as a c oss ein be ween CuP and
CuA (o M+CuA) mo e widely in he Ace ca ia. Bu we
do no e ha he wo in e p e a ions o a e-eme gence
o CuA s. M2 a e challenging o dis inguish amongs ,
and u u e wo k is needed in a b oade sampling o ossil
Holo hysanop e a o p ope ly elucida e he si ua ion o
c own-Thysanop e a. Fu he mo e, new compa a i e an-
a omical s udies based on basi enal bullae and mic o o-
mog aphy da a (Schubnel e al. 2023) may allow a mo e
accu a e iden i ica ion o he main eins in Thysanop e a,
he eby helping o cla i y whe he he e-eme gence o
CuA has occu ed.
Figu e 5. Aeolo h ips i a us Haliday, 1836 (Thysanop e a: Aeolo h ipidae). A Gene al habi us (scale ba = 1 mm), B o ewing
de ails (scale ba = 250 μm).
A h opod Sys ema ics & Phylogeny 83, 2025, 521–530 529
5. Decla a ions
Au ho s’ con ibu ions. Concep ualiza ion: MB, MM, MSE, PN, AN.
Da a Cu a ion: MM, FM, GM. Fo mal analysis: MB, MM, FM, GM,
AS, MSE, PN, AN. In es iga ion: MB, MM, MSE, PN, AN. Supe i-
sion: MM, MSE, AN. W i ing—o iginal d a : MB, MM, MSE, AN.
W i ing— e iew and edi ing: MB, MM, FM, GM, AS, MSE, PN, AN.
Compe ing in e es s. The au ho s decla e no compe ing in e es s.
E hical aspec s. Fossils we e collec ed in Swi ze land in acco dance
wi h na ional legal equi emen s and e hical s anda ds.
Pe missions. All ossils we e collec ed unde exca a ion pe mi s g an -
ed o he Museo can onale di s o ia na u ale o Lugano, in acco dance
wi h Swiss legal equi emen s.
Funding. Fieldwo k and ossil p epa a ion ha e been g an ed by he
Dipa imen o del e i o io del Can one Ticino and he Swiss Fede al
O ice o he En i onmen .
6. Acknowledgemen s
We acknowledge D . Rudol S ocka (Museo can onale di s o ia na-
u ale) o his help ulness and aluable ad ice on li e a u e ela ed o
Mon e San Gio gio, o loca ing si e VM 12, and o di ec ing he pa-
laeon ological exca a ion campaign a his si e in 2020. We also ac-
knowledge D . F édé ic Legend e (MNHN) o g an ing access o he
ex an Thysanop e a collec ion a MNHN, as well as An oine Man ille i
(MNHN) and Ch is ophe Ri ie (MNHN) o p o iding access o En-
omology Collec ions’ pho og aphic acili ies. MB con ibu ed o his
wo k du ing his Ph.D. wi hou ecei ing any dedica ed unding.
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