Redesc ip ion, DNA ba coding, and new dis ibu ional eco ds o
Mo dellis ena mic ogemella a E misch, 1965 (Coleop e a, Mo dellidae)
Dá id Selneko ič1, Ján Kodada1, En ico Ruzzie 2,3
1 Depa men o Zoology,Facul yo Na u alSciences,ComeniusUni e si yinB a isla a,Ilko ičo a6,84215,B a isla a,Slo akia
2 Depa men o Science,Uni e si àRomaT e, ialeG.Ma coni446,00146Rome,I aly
3 Na ionalBiodi e si yFu u eCen e (NBFC),90133Pale mo,I aly
h ps://zoobank.o g/F8E5D1F2-DBBE-474F-87BB-E253A4BE253B
Co esponding au ho : Dá id Selneko ič ([email p o ec ed])
Academic edi o : Dmi y Telno ♦
Recei ed
25 Sep embe 2025 ♦
Accep ed
5 No embe 2025 ♦
Published
21 No embe 2025
Abs ac
Mo dellis ena mic ogemella a E misch, 1965, eco ded om Bulga ia, Cyp us, G eece, and Hunga y, is he e epo ed o he i s ime
om Tunisia. A de ailed edesc ip ion o he species, supplemen ed wi h pho og aphs and illus a ions o key diagnos ic cha ac e s, is
p o ided based on he examina ion o he holo ype and 25 addi ional specimens. Fu he mo e, six sequences o he DNA ba coding
agmen o he cy och ome c oxidase I (COI) gene gene a ed in his s udy ep esen he i s a ailable DNA da a o he species.
Key Wo ds
Cy och ome c oxidase, holo ype, phylogeny, umbling lowe bee les, Tunisia
In oduc ion
The auna o Tunisia is cu en ly known o comp ise 17
species o Mo dellidae (Ho ák 2020). In his s udy, we add
Mo dellis ena mic ogemella a E misch, 1965 o his lis .
The species was o iginally desc ibed based on wo speci-
mens collec ed in G eece, wi hou u he locali y de ails
(E misch 1965). Since i s desc ip ion, M. mic ogemella a
has been men ioned only ou imes in he li e a u e – in
he iden i ica ion keys o Cen al Eu opean Mo dellis ena
by E misch (1977) and Kaszab (1979), and in he dis i-
bu ional ca alogues o Ho ák (2008, 2020), whe e i was
epo ed o he i s ime om Bulga ia, Cyp us, and
Hunga y. The o iginal desc ip ion was b ie and limi ed o
male cha ac e s, which has p e en ed eliable sepa a ion
o M. mic ogemella a om he mo phologically simila
M. psammophila Peye imho , 1943, desc ibed om Al-
ge ia and known only om wo emale syn ypes. Mo e-
o e , no DNA sequences o M. mic ogemella a ha e been
a ailable o da e.
Du ing a collec ing ip o Tunisia in 2025, he i s au-
ho sampled 25 specimens iden i ied as M. mic ogemella a
h ough di ec compa ison wi h he male holo ype. This ma-
e ial allowed us o p o ide a de ailed edesc ip ion o he
species, including emale cha ac e s ha enable eliable sep-
a a ion om M. psammophila. The newly collec ed ma e ial
also yields he i s DNA ba codes o M. mic ogemella a,
which a e now publicly a ailable in he Ba code o Li e
Da a Sys em (BOLD) and GenBank.
Ma e ial and me hods
We examined he male holo ype o M. mic ogemella a and
25 addi ional specimens collec ed in Tunisia in 2025. Fo
compa a i e pu poses, we also s udied he p ima y ypes
o he ollowing species: M. alge iensis E misch, 1966
(SNSD); M. au eo omen osa E misch, 1977 (SNSD); M.
elb usicola E misch, 1969 (SNSD); M. gemella a Schil-
sky, 1899 (MNB); M. ma occana E misch, 1966 (SNSD);
M. mic ogemella a E misch, 1965 (SNSD); M. peloponne-
sensis Ba en, 1980 (NBCL); M. psammophila Peye im-
ho , 1943 (MNHN); M. py enaea E misch, 1966 (SNSD);
M. henana E misch, 1956 (SNSD); M. wankai E misch,
D sch. En omol. Z. 72 (2) 2025, 449–457|DOI 10.3897/dez.72.173155
Copy igh Dá id Selneko ič 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
Dá id Selneko ič e al.: Mo dellis ena mic ogemella a – edesc ip ion and new eco ds450
1966 (SNSD); and M. zol ani E misch, 1977 (HNHM).
The examined ma e ial is deposi ed in he ollowing p i a e
collec ions and ins i u ions: Dá id Selneko ič p i a e col-
lec ion, B a isla a, Slo akia (DSBS); Hunga ian Na u al
His o y Museum, Budapes , Hunga y (HNHM); Na u alis
Biodi e si y Cen e, Leiden, The Ne he lands (NBCL);
Museum ü Na u kunde de Humbold -Uni e si ä , Be -
lin, Ge many (NMB); Muséum na ional d’His oi e na-
u elle, Pa is, F ance (MNHN); Senckenbe g Na u his-
o ische Sammlungen, D esden, Ge many (SNSD).
All examined specimens, including ypes and
DNA- ouche specimens, we e soaked o se e al hou s
in 5% ace ic acid a oom empe a u e, dissec ed, and
moun ed on ca ds. Dissec ed body pa s used o d aw-
ings and pho og aphs we e clea ed in 5% KOH a oom
empe a u e o se e al days and hen moun ed on empo-
a y slides in glyce ol. A e examina ion, hese body pa s
we e moun ed on he ca d oge he wi h he espec i e
specimen using dime hyl hydan oin o maldehyde (En-
omop axis, Ba celona, Spain). Specimens we e s udied
unde a Leica M205 C s e eomic oscope (Leica, We zla ,
Ge many) wi h magni ica ion up o 160× and di used
LED ligh (6400 K). Habi us pho og aphs we e aken wi h
a Canon EOS 5D Ma k II came a (Canon, Tokyo, Japan)
a ached o a Zeiss Axio Zoom.V16 s e eoscope (Zeiss,
Obe kochen, Ge many), and pho og aphs o dissec ed
body pa s we e aken wi h a Zeiss Axio Image .M2 mi-
c oscope. Images we e s acked using Ze ene S acke .1.4
(h ps://ze enesys ems.com/cms/s acke ) and edi ed wi h
Adobe Pho oshop CC (h ps://www.adobe.com/p oduc s/
pho oshop.h ml). D awings o geni alia we e p epa ed
using a d awing ube a ached o a Leica DM1000 mi-
c oscope. Measu emen s we e aken wi h an ocula mi-
c ome e in he Leica M205 C s e eomic oscope. Values
a e gi en as anges, ollowed by he a i hme ic mean and
s anda d de ia ion in pa en heses. The measu ed cha ac-
e s a e abb e ia ed as ollows: EL – ely al leng h om
scu ella apex o ely al apices along su u e; EW – maxi-
mum ely al wid h; HL – head leng h om an e io clypeal
ma gin o occipi al ca ina along midline; HW – maximum
head wid h; LP L – maximum le pa ame e leng h; PL –
p ono al leng h along midline; PW – maximum p ono al
wid h; PygL – maximum pygidial leng h; RP L – maxi-
mum igh pa ame e leng h; TL – combina ion o head,
p ono al and ely al leng hs. Ocula index e e o a a io
o head wid h and minimum in e ocula dis ance.
DNA isola ion and ampli ica ion p ocedu es ollowed
Selneko ič e al. (2023). Ba coding agmen s o he COI
gene we e ampli ied by PCR using he s anda d p ime
pai LEP-F1 and LEP-R1 (Hebe e al. 2003). Nucleo ide
sequences we e immed and assembled in o con igs wi h
Ch omasP o .2.1.10 (Technelysium P y L d, Sou h B is-
bane, Queensland, Aus alia). The inal alignmen was
pe o med manually in Mesqui e .3.81 (Maddison and
Maddison 2023). The ini ial da ase consis ed o newly
gene a ed sequences o M. mic ogemella a oge he wi h
all sequences o Mo dellis ena om Eu ope and No h
A ica a ailable on BOLD. Unique haplo ypes we e il-
e ed using he FaBox DNA o haplo ype collapse ool
(Villesen 2007), and he collapsed da ase was used o
subsequen analyses. BOLD P ocess IDs and GenBank
accession numbe s o all sequences used in his s udy a e
p o ided in he Suppl. ma e ial 1.
Unco ec ed pai wise dis ances (p-dis ances) we e cal-
cula ed in MEGA .11 (Tamu a e al. 2021). All ambigu-
ous si es we e coded as missing p io o he compu a ion.
Maximum likelihood (ML) analysis was pe o med in
IQ-TREE (Nguyen e al. 2015). The bes - i subs i u ion
model (GTR+F+R4) was selec ed by he buil -in Mod-
elFinde using he Bayesian In o ma ion C i e ion. Node
suppo was assessed wi h 1000 ul a as boo s ap ep-
lica es and he SH-like aLRT es (Guindon e al. 2010),
also wi h 1000 eplica es. The esul ing ee was oo ed
and edi ed in In e ac i e T ee o Li e (iTOL) .6 (Le u-
nic and Bo k 2024). Tomoxia bucephala Cos a, 1854 and
Mo della aculea a Linnaeus, 1758 (Mo dellidae, Mo del-
lini) we e used as ou g oups.
Resul s
Species edesc ip ion
Mo dellis ena mic ogemella a E misch, 1965
Figs 1–3
Mo dellis ena mic ogemella a E misch, 1965: 265–267; E misch 1977:
163; Kaszab 1979: 40–41; Ho ák 2008: 99; Ho ák 2020: 96.
Ma e ial examined. Holo ype (designa ed in o iginal
desc ip ion) G eece • male; “G aecia”; SNSD [geni a-
lia dissec ed]. Pho og aph o he holo ype a ailable a
h ps://doi.o g/10.5281/zenodo.17435622.
O he ma e ial. Tunisia • 19 males, 6 emales; Bi Al
Hu ay; 34.908249°N, 9.179063°E; al . 435 m; 11 May
2025; D. Selneko ič leg.; ude al ege a ion, on Ane h-
um; DSBS, DSBS1079 o DSBS1083, DSBS1088.
Di e en ial diagnosis. Mo dellis ena mic ogemella a
was included in he M. gemella a mo phological species
g oup (E misch 1965) based on he ollowing ea u es:
an ennome es 5–10 dis inc ly longe and b oade han he
p eceding wo segmen s, and a me a ibia bea ing, in ad-
di ion o he apical c enidium, wo sho la e al c enidia
o ien ed app oxima ely pe pendicula o he do sal ibial
ma gin.
Wi hin he M. gemella a g oup, M. mic ogemella a
is u he cha ac e ised by i s en i ely black in egumen ,
including he me a ibial spu s; pale es i u e (yellowish
o b ownish); uni o mly colou ed ely al es i u e, wi h
only a na ow, da kened band along he su u e; and sho
an ennome es 5–10, each up o 1.25× longe han wide
(Fig. 2A). The ela i e leng h o an ennome es sepa a es
he species om M. gemella a (Po ugal, Spain), in which
an ennome es 5–10 a e abou 2.0× longe han wide.
Mo dellis ena mic ogemella a di e s om M. alge ien-
sis (Alge ia, I aly, Tunisia), M. au eo omen osa (Mo occo),
M. peloponnesensis (Cyp us, G eece, I aly, Tu key), and
M. py enaea (Spain) by he penul ima e abdominal e gi e
D sch. En omol. Z. 72 (2) 2025, 449–457
dez.penso .ne
451
being concealed by he ely a (Fig. 1) (in he o me species
i is usually exposed). In addi ion, he male s e ni e VIII
o M. mic ogemella a has b oadly ounded and indis inc
pos e o-la e al angles (Fig. 2H), whe eas in he o me spe-
cies hese angles a e p ominen (Selneko ič and Kodada
2022; Selneko ič e al. 2024). Each o hese species also
di e s om M. mic ogemella a in he shape o he pa am-
e es [see Selneko ič and Kodada (2022) o M. alge iensis;
Selneko ič e al. (2024) o M. peloponnesensis; E misch
(1966a) o M. py enaea; E misch (1966b) o M. au eo-
omen osa]. Mo dellis ena ma occana (Mo occo, Tunisia)
di e s om M. mic ogemella a in ha ing a wide body,
wi h sho e and wide ely a.
The mos simila species is M. psammophila, known
only om wo emale syn ypes om Alge ia (Peye im-
ho 1943). Females o M. mic ogemella a can be eliably
sepa a ed by he s uc u e o s e ni e VIII: in M. mic oge-
mella a he pos e io ma gin is p oduced and bea s long
ichoid sensilla, which a e nea ly as long as he s e ni e
wid h (Fig. 2I), while in M. psammophila he homologous
sensilla a e much sho e (Fig. 2J). Fu he mo e, he spic-
ulum en ale is longe and cla a e basally in M. mic oge-
mella a (ca. 0.7× as long as s e ni e along midline), bu
sho e and cylind ical in M. psammophila (ca. 0.4×).
The sympa ic M. pseudohi ipes E misch, 1965 may
also esemble M. mic ogemella a, bu i can be dis in-
guished by ha ing a leas h ee well-de eloped la e al
me a ibial c enidia and di e en ly shaped pa ame es
(Selneko ič and Kodada 2019).
Desc ip ion. Dimensions (mm; males n = 10, emales
n = 4): TL: ♂♂ 3.14–3.62 (3.40 ± 0.16), ♀♀ 3.14–3.41
(3.32 ± 0.12); HL: ♂♂ 0.61–0.69 (0.65 ± 0.02), ♀♀ 0.60–
0.66 (0.63 ± 0.03); HW: ♂♂ 0.65–0.76 (0.70 ± 0.03), ♀♀
0.61–0.67 (0.65 ± 0.03); PL: ♂♂ 0.78–0.91 (0.85 ± 0.04),
♀♀ 0.76–0.83 (0.81 ± 0.03); PW: ♂♂ 0.77–0.92 (0.86 ±
0.05); ♀♀ 0.78–0.86 (0.84 ± 0.04); EL: ♂♂ 1.76–2.05
(1.90 ± 0.11), ♀♀ 1.79–1.94 (1.89 ± 0.07); EW ♂♂ 0.85–
0.95 (0.91 ± 0.03), 0.82–0.95 (0.91 ± 0.06); PygL: ♂♂
1.01–1.14 (1.09 ± 0.04), ♀♀ 0.88–0.97 (0.94 ± 0.04); RP L:
0.25–0.30 (0.27 ± 0.01); LP L: 0.31–0.36 (0.34 ± 0.02).
Body elonga e, wedge-shaped (Fig. 1); wides a mid-
dle o p ono um in males and be o e midleng h o ely a
in emales. In egumen en i ely black. Ves i u e yellowish
wi h a weak eddish o pu plish lus e, sligh ly da kened
in a na ow band along he ely al su u e, and da k b own-
ish o black on he pos e io po ions o en i es 3 and 4,
he whole o en i e 5, and he pygidium.
Head mode a ely con ex, highes poin si ua ed be-
hind he middle in la e al iew; HW/HL a io: ♂♂ 1.03–
1.14 (1.07 ± 0.03), ♀♀ 1.02–1.06 (1.04 ± 0.02). An e io
clypeal ma gin s aigh . Occipi al ca ina e enly con ex.
Su ace inely mic o e icula e, wi h ound se i e ous
punc u es. Lab um ans e se, se ose, shallowly ema -
gina e a an e io ma gin. Eyes o al, abou 1.6× longe
han wide, inely ace ed, se ose; in e ace al se ae longe
han ace diame e ; ocula index: ♂♂ 1.44–1.60 (1.52 ±
0.05), ♀♀ 1.46–1.55 (1.49 ± 0.04). Pos ocula gena ex-
plana e, sligh ly na owe han ace diame e , no angu-
la e. An enna weakly se a e (Fig. 2A); scapus and pedi-
cel cylind ical, subequal in wid h, pedicel sligh ly longe ;
an ennome e 3 smalles , conical, sligh ly sho e and
na owe han ollowing; an ennome e 4 conical, abou
0.7× as long as ollowing and sligh ly na owe ; an enno-
me es 5–10 sub apezoidal, 1.20–1.25× longe han wide
in bo h sexes; an ennome e 11 o al, abou 2.0× longe
han wide. Galea mode a ely long, no p oduced (Fig. 2B,
C). Maxilla y palpome e 2 s ongly expanded wi h long
ichoid sensilla on en al su ace in males (Fig. 2B), no
expanded and wi hou ma kedly long se ae in emales
(Fig. 2C); e minal palpome e iangula , abou 2.2× lon-
ge han wide, wi h inne angle a midleng h in males,
slende e and wi h inne angle behind midleng h in e-
males. Labial e minal palpome e cylind ical. (Fig. 2D)
P ono um mode a ely con ex, wides a middle; PW/
PL a io: ♂♂ 0.99–1.07 (1.02 ± 0.03), ♀♀ 1.01–1.06
(1.04 ± 0.02); an e io ma gin weakly p oduced a mid-
dle; la e al ca ina conca e in la e al iew; pos e o-la e al
angles ec angula , na owly ounded in la e al iew. Su -
ace mic o e icula e, wi h asp-like se i e ous punc u es.
Scu ellum iangula . Me anepis e num sub apezoidal,
do sal ma gin conca e, en al ma gin s aigh , pos e i-
o ly abou as wide as meso ibia.
Ely a mode a ely con ex; EL/EW a io: ♂♂ 1.97–2.22
(2.10 ± 0.08), ♀♀ 2.03–2.18 (2.07 ± 0.07); la e al ma gins
ounded, e enly con e ging (Fig. 1); su ace s ongly mi-
c o e icula e, wi h e y dense asp-like se i e ous punc u es.
P o ochan in wi h ex ended sensillum only sligh -
ly longe and hicke han su ounding se ae, which a e
dis inc ly longe in males han emales (Fig. 2E, F). P o-
emu uni o mly se ose, wi hou conspicuous ex ended
sensilla; se ae dis inc ly longe in males han in emales
(Fig. 2E, F). P o ibia abou as long as p o a sus; in males
weakly expanded behind base and bea ing a dis inc
inge o ex ended se ae (Fig. 2E), in emales s aigh ,
wi hou ex ended se ae (Fig. 2F). P o a some es cylin-
d ical; in males wi h se e al hick ex ended se ae en-
o-la e ally. P o a some e 1 sligh ly longe han nex wo
combined; p o a some e 3 no expanded, apical ma gin
weakly conca e. Meso ochan in wi h dis inc ex ended
sensillum. Meso ibia abou 0.8× as long as meso a sus.
Me a ibia wi h wo sho la e al c enidia eaching o mid-
wid h o ibia, some imes wi h ace o a hi d c enidium
(Fig. 2G); apical spu s da k b own o black, inne spu
abou 1.6× as long as ou e one. Me a a some e 1 wi h
h ee c enidia; me a a some e 2 wi h wo (Fig. 2G).
Pygidium acumina e, no cons ic ed (Fig. 1); EL/PygL
a io: ♂♂ 1.65–1.87 (1.75 ± 0.07), ♀♀ 1.96–2.07 (2.02 ±
0.05). Ven i e 5 wi h apical ma gin con ex. S e ni e VIII
in males abou 1.5× longe han wide, wi h pos e io ma -
gin p oduced and bea ing long ichoid sensilla (Fig. 2H);
in emales pos e io ma gin p oduced, wi h ma kedly
long ichoid sensilla along la e al and pos e io ma gins,
being nea ly as long as s e ni e wid h (Fig. 2I); spiculum
en ale cla a e, abou 0.7× as long as s e ni e midline.
Phallobase long (Fig. 2K), 0.5× as long as EL; epime e
scle o ised, la e al ma gins weakly conca e. Le pa ame e
long (Fig. 3); do sal p ocess s ongly expanded, obliquely
unca e apically; median p ocess sho and ounded; en-
dez.penso .ne
Dá id Selneko ič e al.: Mo dellis ena mic ogemella a – edesc ip ion and new eco ds452
Figu e 1. Mo dellis ena mic ogemella a E misch, 1965 habi us. A. Male, DSBS1080; B. Female.
al p ocess sho and weakly cu ed do sad; i e o se -
en campani o m sensilla in longi udinal ow si ua ed nea
base o do sal p ocess. Righ pa ame e long (Fig. 3); do -
sal p ocess mode a ely expanded, ounded apically; en-
al p ocess sligh ly sho e han do sal one, shallowly bi-
u ca e apically. Penis long, weakly expanded be o e apex.
O iposi o wi h pa ap oc s sho (Fig. 2L); coxi e
abou 1.3× as long as pa ap oc , wi h h ee s ongly ex-
ended la e al sensilla; s ylus wi h wo apical ichoid sen-
silla; p oc ige wide, apex unca e.
Seconda y sexual dimo phism. Males a e sligh ly
mo e slende han emales (Fig. 1). The second maxilla y
palpome e is expanded and bea s s ongly ex ended se ae
in males (Fig. 2B) bu is unexpanded and lacks ex ended
se ae in emales (Fig. 2C); in addi ion, he male ou h
(apical) maxilla y palpome e is subs an ially wide han
ha o he emale. The p o ochan in and p o emu ca y
dis inc ly longe se ae in males (Fig. 2E) han in emales
(Fig. 2F). The p o ibia is expanded behind he base and
bea s a dis inc inge o ex ended se ae in males (Fig.
2E), while i is s aigh and wi hou ex ended se ae in e-
males (Fig. 2F). The pygidium is sho e in emales.
DNA sequences. Six pa ial sequences o he COI
gene a e a ailable on BOLD and GenBank. Fo accession
numbe s e e o Suppl. ma e ial 1.
Dis ibu ion. Bulga ia, Cyp us, G eece, Hunga y, Tu-
nisia (new eco d).
Resul s o DNA analyses
Six COI ba coding sequences o M. mic ogemella a we e
gene a ed in his s udy. All specimens o igina ed om a sin-
gle locali y in Bi Al Hu ay, Tunisia. The sequences ep-
esen ed wo haplo ypes sepa a ed by a single subs i u ion
(posi ion 340: C/T), co esponding o a 0.15% p-dis ance.
D sch. En omol. Z. 72 (2) 2025, 449–457
dez.penso .ne
453
Figu e 2. Mo dellis ena mic ogemella a E misch, 1965. A. An enna, male DSBS1079 (le ), emale ( igh ); B. Maxilla, male
DSBS1079; C. Maxilla, emale; D. Labium, male DSBS1079; E. Fo eleg, male DSBS1079; F. Fo eleg, emale; G. Hindleg, male
DSBS1079; H. S e ni e VIII, male DSBS1079; I. S e ni e VIII, emale; K. Phallobase, DSBS1079; L. O iposi o . Mo dellis ena
psammophila Peye imho , 1943; J. S e ni e VIII, emale, syn ype EC33082.
dez.penso .ne
Dá id Selneko ič e al.: Mo dellis ena mic ogemella a – edesc ip ion and new eco ds454
Figu e 3. Mo dellis ena mic ogemella a E misch, 1965 pa ame es. A. Holo ype; B. DSBS1081; C. DSBS1080; D. DSBS1079.
Scale ba : 0.1 mm.
D sch. En omol. Z. 72 (2) 2025, 449–457
dez.penso .ne
455
The nea es neighbou wi hin he da ase was M. hi ipes
Schilsky, 1895, wi h in e speci ic p-dis ances anging om
14% o 15.22%. Maximum likelihood analysis e ealed
ha M. mic ogemella a is no closely ela ed o membe s
o he M. gemella a species g oup (M. gemella a, M. pelo-
ponnesensis, M. py enaea), bu ins ead clus e s wi hin he
M. hi ipes species g oup (M. hi ipes, M. pseudohi ipes,
M. pu pu ascens A. Cos a, 1854) (Fig. 4). The clade con-
aining membe s o he M. hi ipes g oup, including M.
mic ogemella a, was eco e ed as monophyle ic wi h 98%
ul a as boo s ap suppo .
Figu e 4. Phylogene ic ee based on publicly a ailable sequences o Mo dellis ena om Eu ope and No h A ica. The ee was
in e ed om 259 unique haplo ypes o a 658 bp COI agmen using he maximum likelihood me hod in IQ-TREE. Node suppo
is indica ed by ul a as boo s ap and SH-aLRT alues. Each e minal clade co esponds o a unique Ba code Index Numbe (BIN)
assigned in he Ba code o Li e Da abase (BOLD).
BOLD:AAO0836 Mo dellis ena sp.
BOLD:AEH2398 Mo dellis ena sp.
BOLD:AEH2012 Mo dellis ena dalma ica
BOLD:AFD3884 Mo dellis ena pe oudi
BOLD:ACD0349 Mo dellis ena sp.
BOLD:AGB4114Mo dellis ena gemella a
BOLD:AEH2098 Mo dellis ena pla ypoda
BOLD:AGQ1255 Mo dellis ena in e sec a
BOLD:AFM896 Mo dellis ena sp.
BOLD:AEU6634 Mo dellis ena semi e uginea
BOLD:ADV4347 Mo dellis ena hume alis
BOLD:AFA5087 Mo dellis ena hume alis
BOLD:ACK3402 Mo dellis ena alsopa uli o mis
BOLD:ACG8926 Mo dellis ena s oeckleini
BOLD:ACG8844 Mo dellis ena sp.
BOLD:ACR4234 Pseudomo dellina klappe ichi
BOLD:AAO4468 Tomoxia bucephala
BOLD:AED3319 Mo della aculea a
BOLD:ABX9869 Mo dellis ena sp.
BOLD:ABU7177 Mo dellis ena sp.
BOLD:ACP2826 Mo dellis ena sp.
BOLD:ABX9922 Mo dellis ena sp.
BOLD:AAO4940 Mo dellis ena hu ingiaca
BOLD:ACG3504 Mo dellis ena sp.
BOLD:ACD1193 Mo dellis ena sp.
BOLD:ABX9786 Mo dellis ena s igipennis
BOLD:ABX9870 Mo dellis ena sp.
BOLD:AEV6360 Mo dellis ena con inis
BOLD:AEH2892 Mo dellis ena pu pu eonig ans
BOLD:AHB0195 Mo dellis ena mic ogemella a
BOLD:AED6443 Mo dellis ena pu pu ascens
BOLD:AED9695 Mo dellis ena hi ipes
BOLD:AED9694 Mo dellis ena hi ipes
BOLD:AFH1628 Mo dellis ena pseudohi ipes
BOLD:AEW6104 Mo dellis ena lindbe gi
BOLD:AED6814 Mo dellis ena minima
BOLD:ACP3789 Mo dellis ena langei
BOLD:AGB4113Mo dellis ena py enaea
BOLD:AFM319 Mo dellis ena peloponnesensis
BOLD:ACP5293 Mo dellis ena a sa a
BOLD:AEA3479 Mo dellis ena pseudo henana
BOLD:ACC3107 Mo dellis ena k aa zi
BOLD:AEH1356 Mo dellis ena epis e nalis
BOLD:AGG4349 Mo dellis ena pa aepis e nalis
BOLD:ADW7498 Mo dellis ena neuwaldeggiana
BOLD:ACE2081 Mo dellis ena hume alis
BOLD:AAP8305 Mo dellis ena hume alis
BOLD:ACA7847 Mo dellis ena a iega a
BOLD:AAX4356 Mo dellis ena sp.
BOLD:ABW9458 Mo dellis ena necopina a
BOLD:AEW4355 Mo dellis ena semi e uginea
BOLD:ACP3915 Mo dellis ena semi e uginea
BOLD:ADV3855, BOLD:ACG8927 Mo dellis ena e doesi
BOLD:ACO6512 Pseudomo dellina acu icollis
BOLD:AAO4939 Mo dellis ena inexpec a a
98.5 / 100
42.7 / 91
99.5 / 100
92.3 / 99
99.2 / 100
97.7 / 98
72.1 / 100
96 / 100
100 / 100
81.2 / 97
99.9 / 100
86.5 / 90
76.2 / 99
100 / 100
99.9 / 100
41.9 / 33
99.5 / 100
58.6 / 93
99.1 / 99
100 / 100
49.7 / 55
26.3 / 85
100 / 100
99.9 / 100
100 / 100
100 / 100
96.3 / 100
100 / 100
99.6 / 100
99.3 / 99
99.3 / 99
100 / 100
63.8 / 93
99.9 / 100
69.4 / 98
100 / 100
81.5 / 94
56.6 / 38
89.7 / 100
83.3 / 98
76.4 / 32
31.7 / 92
92.8 / 87
89.4 / 98
56.1 / 34
100 / 100
94.5 / 99
99.6 / 100
100 / 100
86.8 / 99
100 / 100
100 / 100
94.8 / 100
94.6 / 99
33.8 / 78
28.3 / 79
83.5 / 98
82.6 / 57
93.3 / 100
56.8 / 92
99.1 / 100
96.7 / 100
99.2 / 100
84.7 / 62
66.9 / 97
99.6 / 100
93.5 / 99
97.5 / 99
88.4 / 89
100 / 100
100 / 100
98.9 / 100
99.5 / 100
100 / 100
100 / 100
100 / 100
100 / 100
31.8 / 32
96.2 / 100
95.3 / 99
76.8 / 59
25.5 / 48
100 / 100
93.2 / 99
89 / 76
100 / 100
98.3 / 100
76.6 / 100
81.8 / 94
T
ee scale: 0.1
IQ-TREE: SH-aLRT suppo (%) / ul a as boo s ap suppo (%)
Ou g oup
Mo dellis ena mic ogemella a E misch, 1965
M. gemella a
clade
M. hi ipes
clade
dez.penso .ne
Dá id Selneko ič e al.: Mo dellis ena mic ogemella a – edesc ip ion and new eco ds456
Discussion
Since i s desc ip ion in 1965, no de ailed eco ds o Mo -
dellis ena mic ogemella a ha e been published, apa
om men ions om Bulga ia, Cyp us, and Hunga y by
Ho ák (2020), all wi hou u he de ails. E en he ype
locali y, “G aecia,” is ague and may e e ei he o main-
land G eece o o one o he many G eek islands in he
eas e n Medi e anean Sea. Al hough he i s au ho has
examined a subs an ial numbe o Mo dellidae specimens
om he Balkans, G eece, and Tunisia in he p i a e col-
lec ions and museums o e he pas yea s, no specimens
o M. mic ogemella a we e e e iden i ied. Du ing ield-
wo k in Tunisia in May 2025, eigh locali ies in he no h-
e n and cen al egions o he coun y we e sampled, bu
M. mic ogemella a was ound only a a single locali y
nea Bi Al Hu ay. Based on he sca ci y o a ailable e-
co ds, he species appea s o be localised and a e.
In his o iginal desc ip ion, E misch (1965) assigned
Mo dellis ena mic ogemella a o he M. gemella a spe-
cies g oup based on he p esence o wo sho la e al
c enidia on he me a ibia. In his iden i ica ion key o Cen-
al Eu opean Mo dellis ena (E misch 1977), howe e ,
M. mic ogemella a was e oneously cha ac e ised as ha -
ing an ennome es 5–10 ha a e 1.5–2.0 imes longe han
wide. The alue is inco ec , as bo h he holo ype and he
Tunisian specimens ha e an ennome es only 1.20–1.25
imes longe han wide. Ou obse a ion also ag ees wi h
he o iginal desc ip ion, in which E misch no ed ha he
an ennome es a e “only sligh ly longe han wide, in e-
male squa e.” The dis inc i e s uc u e o he male gen-
i alia – pa icula ly he le pa ame e, which has a long
basal po ion and a do sal p ocess s ongly cons ic ed
medially and ma kedly expanded and obliquely unca e
dis ally, and he igh pa ame e, whose en al p ocess is
shallowly bi u ca e apically – allowed s aigh o wa d
iden i ica ion o he newly collec ed ma e ial a e com-
pa ison wi h he M. mic ogemella a holo ype.
Resul s o he COI ba coding egion analyses sugges ha
Mo dellis ena mic ogemella a is no closely ela ed o he
species o ming he M. gemella a clade bu is ins ead nes ed
wi hin he M. hi ipes clade (Fig. 4). The close a ini y o M.
mic ogemella a o he M. hi ipes clade is u he suppo -
ed also by mo phological cha ac e s, such as he expanded
male second maxilla y palpome e bea ing s ongly ex ended
se ae, he shape o male and emale s e ni e VIII, and he
s uc u e o he pa ame es (Selneko ič and Kodada 2019).
Howe e , mo e comp ehensi e analyses based on la ge
po ions o he genome a e needed o es his hypo hesis.
The emales o Mo dellis ena mic ogemella a close-
ly esemble he wo emale syn ypes o M. psammophila
Peye imho , 1943, desc ibed om El Oued, Alge ia, and
no eco ded since. The ype locali y o M. psammophi-
la is si ua ed app oxima ely 275 km om Bi Al Hu ay.
Howe e , di e ences in he shape o emale s e ni e VIII
and in he leng h o i s sensilla con i m ha hese a e wo
dis inc species.
Acknowledgemen s
We would like o hank Be nd Jaege (MNB), Ola
Jäge (SNSD), An oine Man ille i (MNHN), Győző Szél
(HNHM), Osca Vo s (NBCL) o allowing us o s udy
he ype ma e ial held unde hei cu a ion. This esea ch
ecei ed inancial suppo om he Slo ak Resea ch and
De elopmen Agency (P ojec No. APVV-19-0076), as
well as SYNTHESYS+ P ojec h p://www.syn hesys.
in o/ which is inanced by Eu opean Communi y Resea ch
In as uc u e Ac ion unde he H2020 In eg a ing Ac i -
i ies P og amme (P ojec No. 823827). En ico Ruzzie
acknowledges he suppo o NBFC o he Uni e si y o
Roma T e, Depa men o Science, unded by he I alian
Minis y o Uni e si y and Resea ch, PNRR, “Missione
4 Componen e 2, ‘Dalla ice ca all’imp esa’, In es imen-
o 1.4”, P ojec CN00000033. Museum ü Na u kunde
(Be lin) wai ed he publica ion cos s o his a icle.
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Supplemen a y ma e ial 1
Sequences used in he analyses
Au ho s: Dá id Selneko ič, Ján Kodada, En ico Ruzzie
Da a ype: xlsx
Explana ion no e: Accession numbe s o sequences used
in he analyses, wi h espec i e Ba coding Index Num-
be and haplo ype numbe .
Copy igh no ice: This da ase is made a ailable unde
he Open Da abase License (h p://openda acommons.
o g/licenses/odbl/1.0). The Open Da abase License
(ODbL) is a license ag eemen in ended o allow us-
e s o eely sha e, modi y, and use his Da ase while
main aining his same eedom o o he s, p o ided
ha he o iginal sou ce and au ho (s) a e c edi ed.
Link: h ps://doi.o g/10.3897/dez.72.173155.suppl1
