P o ube a o no aki gen. no ., sp. no ., a new an h oleucosoma id millipede
E-mail: d agan.an [email protected]. s
P o ube a o no aki gen. no ., sp. no ., a new
an h oleucosoma id millipede (Diplopoda: Cho deuma ida:
An h oleucosoma idae) om he Caucasus
D agan ANTIĆ, Slobodan MAKAROV
Uni e si y o Belg ade—Facul y o Biology, Ins i u e o Zoology, S uden ski T g 16, 11000 Belg ade, Se bia
Biologia Se bica, 2025, 47: 54-62 DOI: 10.5281/zenodo.17679416
Summa y. The p esen s udy desc ibes a new monospeci ic cho deuma idan genus, P o ube a o gen. no ., and i s
ype species, P o ube a o no aki gen. no ., sp. no ., om Machakhela Na ional Pa k in Adja a, Geo gia. This new mil-
lipede is cha ac e ized by a p o oundly modi ied head, wi h a e y la ge an e io bulge, as well as dis inc i e a si on
leg-pai 6, and unique gonopods in male. These unique ea u es make he axon easily dis inguishable, no only om
he Caucasian An h oleucosoma idae, bu also om all o he membe s o he amily. The new genus and species a e
desc ibed and diagnosed in de ail, and illus a ions o he male habi us, legs and gonopods a e p o ided. The a ini ies
o he new genus wi h o he membe s o he amily An h oleucosoma idae and head modi ica ions in gene al a e also
b ie ly discussed.
Keywo ds: Adja a, biodi e si y, Geo gia, head modi ica ions, Lesse Caucasus, axonomy.
Accep ed: 13 No embe 2025 / Published online: 17 No embe 2025
INTRODUCTION
The cho deuma idan amily An h oleucosoma idae is
among he mos he e ogeneous membe s o he o de ; he
amily is clea ly non-monophyle ic, and cu en ly includes
92 species in 36 gene a (An ić and Maka o 2016; An ić
2017; An ić e al. 2020, 2023, 2025; Še ić e al. 2022). The
dis ibu ion o cho deuma idan amily An h oleucosoma i-
dae is almos exclusi ely con ined o he Wes e n Palaea c ic,
wi h he main cen e s o di e si ica ion loca ed in he Balkan
Peninsula and he Caucasus.
Mo e han hal o he desc ibed axa wi hin he amily
a e known om he Caucasus eco egion. In e es ingly, his
le el o di e si y emained la gely hidden un il ecen ly, as
only se en species we e known p io o 2016 (An ić and Ma-
ka o 2016). Howe e , o e he pas decade alone, as many
as 45 new species and 15 new gene a o an h oleucosoma ids
ha e been desc ibed. Today, wi h 20 gene a and 52 species,
An h oleucosoma idae ep esen s one o he mos di e se mil-
lipede g oups in he egion (An ić and Maka o 2016, 2022;
An ić e al. 2018, 2023, 2025).
In he p esen s udy, we desc ibe a ema kable new an-
h oleucosoma id genus om he Caucasus, which is cha ac-
e ized by a s ongly modi ied head in he male.
MATERIAL AND METHODS
A single male specimen (holo ype) is p ese ed in 70%
e hanol and deposi ed in he collec ion o he Na u his o-
isches Museum Wien, Aus ia (NHMW). The specimen was
examined using Nikon SMZ745T, Nikon SMZ1270, and Ca l
Zeiss S emi 2000-C binocula s e eo mic oscopes. All axo-
nomically impo an s uc u es we e dissec ed and moun ed
in glyce in as empo a y mic oscope p epa a ions, and ob-
P o ube a o no aki gen. no ., sp. no ., a new an h oleucosoma id millipede
se ed wi h a Ca l Zeiss Axioscope 40 mic oscope. Habi-
us pho og aphs we e aken using a Nikon DS-Fi2 came a
equipped wi h a Nikon DS-L3 came a con olle a ached o
a Nikon SMZ1270 binocula s e eo mic oscope. Pho og aphs
o he legs we e also eco ded wi h a Nikon DS-Fi2 came a
equipped wi h a Nikon DS-L3 came a con olle a ached
o a Nikon SMZ1270 binocula s e eo mic oscope; as well
as wi h a Canon Powe Sho A80 digi al came a connec ed
o a Ca l Zeiss Axioscope 40 mic oscope. All images we e
s acked using Ze ene S acke P o essional 1.04. Line d aw-
ings o he gonopods we e made using acing pape placed
on a compu e moni o displaying images o hese s uc u es
aken wi h a Canon Powe Sho A80 digi al came a connec ed
o a Ca l Zeiss Axioscope 40 mic oscope. The inal images
we e p ocessed and assembled in Adobe Pho oshop CS6.
The e minology used in he desc ip ion ollows An ić
e al. (2025), wi h some modi ica ions.
Abb e ia ions used o deno e pa icula s uc u es o
he gonopods a e explained di ec ly in he desc ip ion and
igu e legend.
RESULTS
Taxonomy
Class Diplopoda de Blain ille in Ge ais, 1844
O de Cho deuma ida Pocock, 1894
Family An h oleucosoma idae Ve hoe , 1899
Genus P o ube a o An ić & Maka o , gen. no .
Diagnosis. The new genus di e s om all o he mem-
be s o he amily An h oleucosoma idae by he p esence in
he male o a e y la ge an e io bulge on he head, dis inc-
i e a si o leg-pai 6 wi h la ge p oximal do sal bulges co -
e ed by small nipples, and uniquely shaped an e io gono-
pods wi h a well-de eloped synangiocoxi e. In addi ion, he
pos e io gonopods lack elopodi es, while he angiocoxi es
a e in he o m o a ew lagelli o m p ocesses.
Name. The genus name P o ube a o is de i ed om he
La in e b p o ube a e (“ o swell, bulge ou ”), e e ing no
only o he la ge head bulge, he mos dis inc i e ea u e o
he genus, bu also o he dis al bulges o he a si o leg-pai
6. The name is ea ed as a masculine noun.
Desc ip ion. Adul male wi h 31 body segmen s (in-
cluding collum and elson). Body whi ish wi h poo ly de-
eloped la e al keels and a he sho and ichoid mac o-
chae e. Head s ongly modi ied, wi h a la ge an e io bulge.
Ocula ield wi h 4+5 poo ly pigmen ed omma idia. Leg-
pai s 3–7 s ongly enla ged and modi ied. Leg-pai 6 wi h
a cha ac e is ic p oximal do sal bulge on he a si, co e ed
by nipples. Leg-pai s 10 and 11 wi h coxal glands. Leg-pai
10 wi h pos e odis al p ocess co e ed by nipples. An e io
gonopods complex; s e num wi h s e nal lamella wi h h ee
lobes and a pai o la e al p ocesses; angiocoxi es in o m o
a pai o shields and bo le shaped synangiocoxi e an e io ly;
a pai o coxal esicles p esen pos e io ly. Pos e io gono-
pods wi hou elopodi es; angiocoxi es in he o m o i e
long lagelli o m p ocesses; colpocoxi es poo ly scle o ized,
memb anous, a he sub iangula .
Type species. P o ube a o no aki gen. e sp. no ., by
p esen designa ion and mono ypy.
P o ube a o no aki An ić & Maka o , gen. e sp. no .
Figs 1–5
Diagnosis. As o he genus.
Name. The species is named a e colleague János
No ák, a Hunga ian a achnologis who collec ed his e-
ma kable specimen. The name is ea ed as a noun in he
geni i e case.
Ma e ial examined. Holo ype male (NHMW-ZOO-
MY-10660) – Geo gia, Adja a, Machakhela Na ional Pa k,
be ween Uchkhi i and Acha isaghma i, lea li e si ing in
a o es ed a ea, 10-11-2023, leg. J. No ák.
Desc ip ion. Numbe o body segmen s and size. Body
wi h 31 segmen s (including collum and elson). Body leng h
13 mm, e ical diame e o he la ges segmen 1.1 mm.
Colo a ion. Whi ish (Figs 1, 2).
Head (Figs 1A, 2A–E). Mos ly smoo h, wi h spa se
se ae; s ongly modi ied, wi h a la ge an e io bulge; do -
sally, wi h a longi udinal medial and wo longi udinal la e al
idges, and wo longi udinal dep essions in be ween; an e-
io side wi h a cen al dep ession and wo smalle la e al
bulges. Lab um wi h h ee lab al ee h, and 4+4 lab al and
2+2 sup alab al se ae. Gna hochila ium no mal, wi hou pe-
culia i ies. An ennae 2.2 mm long. Leng h o an ennome es
(in mm): I (0.11), II (0.25), III (0.60), IV (0.31), V (0.55), VI
(0.20), VII (0.16) and VIII (0.02). Leng h/b ead h a ios o
an ennome es I–VII: I (1.0), II (2.0), III (5.5), IV (2.6), V
(4.0), VI (1.3) and VII (1.6). An ennome es II, IV, V, VI and
VII wi h one, h ee, one, ou and one long sensillum i-
choideum, espec i ely. Omma idia poo ly de eloped, ligh
b ownish, ba ely isible; le side wi h 4 omma idia in wo
ows, igh side wi h 5 omma idia in one line.
Collum. Na owe han head, wi h six mac ochae ae as
on all body segmen s. An e io edge semi-ci cula , pos e io
ma gin gen ly conca e.
Body segmen s (Figs 1, 2F). La e al keels ounded
in do sal iew. Mac ochae ae a he sho and ichoid in
midbody segmen s, somewha longe in he an e io pa
o body.
Telson. No mal, wi hou peculia i ies.
Leg-pai s 1 and 2. Wi h a sal combs; emo a, pos -
emo a and ibiae wi h long and obus se ae (Fig. 3A).
Biologia Se bica 47 55
P o ube a o no aki gen. no ., sp. no ., a new an h oleucosoma id millipedeD. An ić e S. Maka o
Fig. 1. P o ube a o no aki gen. no ., sp. no ., holo ype male (NHMW-ZOO-MY-10660). A, habi us, la e al iew; B, an e io pa o body,
la e al iew. (Scale ba s: 1 mm).
56 Biologia Se bica 47
P o ube a o no aki gen. no ., sp. no ., a new an h oleucosoma id millipede
Fig. 2. P o ube a o no aki gen. no ., sp. no ., holo ype male (NHMW-ZOO-MY-10660). A–E, head in an e io , do sal, do sola e al, an-
e ola e al and la e al iew, espec i ely (whi e a ow indica es omma idia); F, midbody segmen s, do sal iew. (Scale ba s: 0.5 mm).
Biologia Se bica 47 57
P o ube a o no aki gen. no ., sp. no ., a new an h oleucosoma id millipedeD. An ić e S. Maka o
Fig. 3. P o ube a o no aki gen. no ., sp. no ., holo ype male (NHMW-ZOO-MY-10660), legs. A, igh leg 2, an e io iew; B, igh leg
3, pos e io iew (whi e a ow indica es small p oximal en al bulge on emu ); C, leg-pai 5, an e io iew; D, leg-pai 6, an e io iew
(whi e a ow indica es p oximal do sal bulge on a sus); E, leg-pai 7, pos e io iew; F, leg-pai 10, pos e io iew (whi e a ow indica es
dis al pos e io p ocess on coxa). (Scale ba s: 0.5 mm).
58 Biologia Se bica 47
P o ube a o no aki gen. no ., sp. no ., a new an h oleucosoma id millipede
Fig. 4. P o ube a o no aki gen. no ., sp. no ., holo ype male (NHMW-ZOO-MY-10660), legs. A, igh leg 6, pos e io iew; B, coxae 7,
pos e io iew (whi e a ow indica es small pos e io iangula oo h); C, igh a sus 7, pos e io iew; D, coxae 10, pos e io iew; E,
coxae 11, pos e io iew. (Scale ba s: 0.5 mm).
Male sexual cha ac e s. Gonopo es opening mesally
on coxae 2. Leg-pai s 3–7 s ongly enla ged and modi ied
(Fig. 1). Leg-pai 3 s ou , p e emo a and emo a hick, p e-
emo a wi h a p oximal do sal p o usion, emo a wi h a
small p oximal en al bulge on pos e io side (Fig. 3B). Leg-
pai 4 wi h hick p e emo a and emo a, p e emo a wi h a
p oximal do sal p o usion. Leg-pai 5 wi h obus emo a
and a ain p oximal do sal bulge on a si (Fig. 3C). Leg-
pai 6 wi h s ongly de eloped emo a and wi h peculia
a si a wi h well-de eloped p oximal do sal bulge co e ed
Biologia Se bica 47 59
P o ube a o no aki gen. no ., sp. no ., a new an h oleucosoma id millipedeD. An ić e S. Maka o
by nipples and en al side also co e ed by nipples (Figs 3C,
4A). Leg-pai 7 he mos obus , coxae wi h a small, pos e-
io , iangula , den icula e p ocess, an e io ly wi h a small
en al dis al bulge; p e emo a wi h dis al en al bulge; a si
elonga e, en al side la ened wi h small nipples (Figs 3E,
4B, C). Leg-pai 10 wi h coxal glands and dis al pos e io
coxal p ocesses wi h small den icles (Figs 3F, 4D). Leg-pai
11 wi h coxal glands and small dis al pos e io p ocess on
coxae. Leg-pai s 12 and 13 wi h ounded, dis al bulge on
coxae, mo e p onounced on leg-pai 12. Legs no mal om
leg-pai 14 onwa ds.
An e io gonopods (Fig. 5A–C). S e num (s) s ongly
de eloped, wi h a well-de eloped, imb ia e, ilobed s e -
nal lamella (sl) on an e io side, la e al lobes highe han
he medial one; a pai o an e ola e al, sub iangula , s e -
nal p ocesses (sp) p esen . Angiocoxi es (a) shieldlike, wi h
hin inged la e al lamellae (ll); medially, angiocoxi es
nea ly used, o ming pos e io ly a pai o mesal lamellae
(ml); dis ally, shield na owing in he o m o a ounded ip;
pos e io ly, bo h angiocoxi es wi h in e media e lamella (il).
An e io ly, s ongly de eloped, bo le-shaped synangiocoxi e
(sa) p esen ing wi h na owing dis al pa (b oken). Coxal
Fig. 5. P o ube a o no aki gen. no ., sp. no ., holo ype male (NHMW-ZOO-MY-10660), gonopods. A–C, an e io gonopods in an e io ,
la e al and pos e io iew espec i ely; D, pos e io gonopods, an e io iew. Abb e ia ions: a: angiocoxi e; c: colpocoxi e; c : coxal esicle;
cx: coxi e; d: an e io dep ession; il: in e media e lamella; ll: la e al lamella; ml: mesal lamellae; s: s e num; sa: synangiocoxi e; sl: s e nal
lamella; sp: s e nal p ocess. (Scale ba s: 0.2 mm).
60 Biologia Se bica 47
P o ube a o no aki gen. no ., sp. no ., a new an h oleucosoma id millipede
esicles (c ) well-de eloped, wing-like, p oximal pa s na -
owing wi h inged la e al ma gins.
Pos e io gonopods (Fig. 5D). Wi h de eloped s e num
(s). Coxi es (cx) sho , wi h semici cula an e io idge wi h
dis ola e al ele a ion co e ed by minu e se ae. A semici cula
an e io idge bo de s he an e io dep ession (d). Angiocox-
i es (a) la e al in posi ion, consis ing o i e lagelli o m p o-
cesses, sligh ly cu ed pos e iad. Colpocoxi es (c) a ise om
pos e io side o he coxi al dep ession, weakly scle o ized,
lamella , mo e o less sub iangula , wi h an an e odis al old;
sho e han angiocoxi es. No aces o elopodi es.
DISCUSSION
An ić and Maka o (2016) es ablished eigh complexes
o gene a, in which hey classi ied all Caucasian An h oleu-
cosoma idae. The new genus can be p o isionally placed
wi hin he He culina complex. Acco ding o An ić and Ma-
ka o (2016), his complex includes only he genus He culina
An ić & Maka o , 2016, wi h wo species, which is diagnosed
by “... he p esence o a e y obus and biza e male leg-pai
6, and by he p esence o nume ous a ow-like ilamen s
on he coxi es o he pos e io gonopods”. In addi ion, bo h
He culina species ha e a obus leg-pai 5. In he new genus,
leg-pai s 5 and 6 a e no as obus as in He culina, bu hey
do ha e somewha enla ged emo a. The second diagnos ic
cha ac e o he He culina complex in he pos e io gono-
pods is he p esence o angiocoxi es in he o m o nume ous
lagelli o m p ocesses. The same is ue o he new genus,
which has angiocoxi es in he o m o i e lagelli o m s uc-
u es. In addi ion, he colpocoxi es a e e y simila in bo h
He culina and he new genus. While bo h He culina species
ha e small elopodi es on he pos e io gonopods, hese a e
absen in he only ep esen a i e o P o ube a o gen. no . I
is no ewo hy o men ion ha he elopodi es a e also absen
om he pos e io gonopods o he Caucasian monospeci ic
genus Ra cheuma Golo a ch, 1985 (Golo a ch 1985; An ić
and Maka o 2017). Mo e gene ally, wi h he excep ion o
hese wo Caucasian gene a, wi hin he amily An h oleu-
cosoma idae, he absence o elopodi es on he pos e io
gonopods also cha ac e izes he Balkan ep esen a i es o
he Bulga osoma complex and ou species o he genus
Anamas igona Sil es i, 1898, as well as he No h Ame ican
species Leschius mcallis e i Shea & Leona d, 2004 (Shea
and Leona d 2004; An ić 2017). The absence o elopodi es
on he gonopods o Diplopoda is a de i ed (apomo phic)
cha ac e s a e; he e o e, he new genus belongs o hose
gene a wi hin An h oleucosoma idae ha possess de i ed
pos e io gonopods. Fu he mo e, when compa ing he new
genus wi h He culina, he s uc u e o he an e io gono-
pods should also be conside ed. Bo h gene a sha e a simila
con igu a ion o hei angiocoxal shields, which bea h ee
ypes o lamellae: la e al, in e media e, and mesal. In He cu-
lina oligosagi ae An ić & Maka o , 2016, he la e al lamellae
a e ben pos e io ly, whe eas in He culina polysagi ae An ić
& Maka o , 2016, and in he new genus, hey a e di ec ed
la e ad. Bo h gene a also ha e e y simila s uc u es in he
dis al po ion o he angiocoxal shields and possess a pai o
coxal esicles pos e io ly. Howe e , he new genus clea ly di
-
e s om He culina in ha ing a s ongly de eloped, iloba e
s e nal lamella and a pai o an e ola e al s e nal p ocesses,
and by he p esence o a s ongly de eloped synangiocoxi e.
Among he An h oleucosoma idae, a synangiocoxi e in he
o m o an an e io medial p ocess be ween he angiocoxal
shields is also p esen in he monospeci ic Caucasian genus
B achychae osoma An ić & Maka o , 2016, belonging o he
Caucaseuma complex (An ić and Maka o 2017: 27, ig.
17A–C). Howe e , he wo gene a di e ma kedly in nu-
me ous ex e nal and gonopodal cha ac e s. Finally, i should
be no ed ha P o ube a o gen. no . and bo h species o he
genus He culina a e geog aphically close and inhabi o es
habi a s in he Lesse Caucasus in Geo gia.
As men ioned abo e, he new genus is cha ac e ized
by a s ongly modi ied male head. A modi ied male head is
common among cho deuma idans, and is mainly e lec ed
by he ac ha males ha e ei he a la ened o sligh ly con-
ca e an e io pa o he head, whe eas in emales he head is
no mal and sligh ly con ex. Howe e , he p esence o a mo e
o less p onounced an e io bulge on he head appea s o be
e y a e among Cho deuma ida. To ou knowledge, a bulge
on he male head cha ac e izes some ep esen a i es o he
amily An h oleucosoma idae. Among Caucasian an h oleu-
cosoma ids, apa om he new genus, he only axon wi h
an an e io bulge on he male head is he monospeci ic genus
Adsha dicus Golowa ch, 1981 (see An ić and Maka o 2016:
20, ig. 10B, C). An an e io bulge is also p esen in some
Balkan an h oleucosoma ids. I is mos p onounced in Bul-
ga osoma bu eschi Ve hoe , 1926 (S asse 1962: 450, ig. 17;
pe sonal obse a ion) and Bana osoma ocella um (Tabaca u,
1967; pe sonal obse a ion); whe eas a much smalle bulge
can be seen, o example, in males o h ee Se bosoma species
(Ću čić and Maka o 2000; pe sonal obse a ion). The ex-
amples discussed abo e e e o head modi ica ions cha ac-
e is ic only o males. Howe e , he e is also a case o a head
modi ica ion ha occu s in bo h sexes: he No h Ame ican
oglobi ic cleidogonid Pseudo emia peponoc anium Shea ,
2011. Adul s o bo h sexes exhibi a la ge, squa e bulge on he
head, which is smalle in emales (see Shea 2011: 15, igs.
34–35). The new genus ep esen s he mos s iking example
o head modi ica ion known among an h oleucosoma ids.
Un o una ely, in he absence o collec ed emales, i emains
unce ain whe he his modi ica ion is es ic ed o males.
I should be no ed ha male head modi ica ions also
Biologia Se bica 47 61
D. An ić e S. Maka o
occu in he o de s Callipodida and Polydesmida. Among
Callipodida, his is pa icula ly cha ac e is ic o he genus
Pa aco ina Wang & Zhang, 1993 (Akka i e al. 2023), wi h
pe haps he mos biza e example ound in he Vie namese
oglomo phic species Pa aco ina ky ang Nguyen, S oe ,
Nguyen & Vu, 2023 (see Nguyen e al. 2023: 185, ig. 2B).
Among Polydesmida, head modi ica ions a e obse ed in
some A o- opical Py godesmidae and T ichopolydesmidae,
occu ing ei he only in males o in bo h sexes (Golo a ch
2018; Golo a ch e al. 2018, 2019).
The unc ion o he modi ied head in males emains
unknown. In addi ion o head modi ica ions, i is wo h no -
ing he p esence o modi ica ions o he p egonopodal legs in
male Cho deuma ida and Callipodida. These legs, which a e
o en g ea ly hickened and bea ing a ious ex ensions, a e
assumed o play an impo an ole du ing ma ing by p o id-
ing i me s abiliza ion o he emale. I is possible ha he
modi ied heads o males se e a simila unc ion, con ib-
u ing o he s abiliza ion o he emale du ing copula ion.
Howe e , such modi ica ions a e appa en ly a e, and a e no
genus-speci ic: occu ing independen ly in se e al lineages.
Fu he mo e, he p esence o simila head modi ica ions in
bo h sexes o Pseudo emia peponoc anium aises addi ional
ques ions. I head modi ica ions unc ion in ma ing oge he
wi h he p egonopodal legs, why do hey occu in bo h sexes
in his species, and why a e hey absen in o he species ha
e iden ly achie e success ul ma ing wi h “only” he modi ied
p egonopodal legs o males? I is ema kable ha , al hough
hese head and p egonopodal leg modi ica ions a e bo h con-
spicuous and well-documen ed, hei unc ional signi icance
emains unce ain.
ACKNOWLEDGMENTS
We a e g a e ul o János No ák o p o iding he alu-
able specimen used o his s udy. We a e also e y g a e ul
o Bill Shea o d awing ou a en ion o he unusual head
o Pseudo emia peponoc anium. Many hanks as well o he
e iewe s Nes ine Akka i (Aus ia) and Se gei Golo a ch
(Russia) o hei help ul commen s and co ec ions ha im-
p o ed he manusc ip .
This s udy was pa ly suppo ed by he Minis y o Sci-
ence, Technological De elopmen and Inno a ion o he Re-
public o Se bia (451-03-137/2025-03/ 200178).
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