1
Two new species o he genus Ache on iella (Collembola,
Hypogas u idae) om ca es o he Albo z and Kope Dagh
moun ains in I an
Mahmood Meh a ooz May an1, Ľubomí Ko áč1
1 Depa men o Zoology,Ins i u eo BiologyandEcology,Facul yo Science,Pa olJoze Ša á ikUni e si y,Š obá o a2,04154Košice,Slo akia
Co espondingau ho s:MahmoodMeh a oozMay an(mahmood_meh a [email p o ec ed]);Ľubomí Ko áč(lubomi [email p o ec ed])
Copy igh : ©
Mahmood Meh a ooz May an & Ľubomí Ko áč.
This is an open access a icle dis ibu ed unde
e ms o he C ea i e Commons A ibu ion
License (A ibu ion 4.0 In e na ional – CC BY 4.0).
Resea ch A icle
Abs ac
Two new sub e anean species o he genus Ache on iella Absolon, 1913 om I an
a e desc ibed, illus a ed, and mo phologically di e en ia ed om simila species.
Ache on iella nezamdoos i sp. no . has la ge dis ibu ion ange co e ing ca es o No h
Kho asan, Raza i Kho asan, and Teh an p o inces, while A. palaciosi sp. no . is p obably
es ic ed o Moghan Ca e in Raza i Kho asan p o ince. Bo h new species a e cha -
ac e ized by absence o eyes, PAO, empodium, u ca, and e inaculum. Ache on iella
nezamdoos i sp. no . is dis inguished om o he species by ha ing wo small anal
spines, Th II–III wi h se ae m1, m4, and m5, and ou hickened an e io se ae on he
lab um. Ache on iella palaciosi sp. no . is cha ac e ized by bea ing wo medial se ae in
he an e io ow o he lab um hickened in o spines and apically den a e, and by he ab-
sence o se ae m3 on Th I and p3 on Th II–III. An upda ed key and a able summa izing
diagnos ic cha ac e s o wo ld species o he genus Ache on iella a e p o ided.
Key wo ds: Mo phology, se al pa e n, Sou hwes Asia, sub e anean auna, oglobion
In oduc ion
Ache on iella Absolon, 1913 is a cosmopoli an genus belonging o he amily
Hypogas u idae and desc ibed om a ca e in Alge ia (Absolon 1913; Thibaud
1990). Species o his genus a e cha ac e ized by hei usi o m, unpigmen -
ed body, and he absence o eyes, pos -an ennal o gan, e inaculum, and u ca
(Thibaud 1990; Thibaud e al. 2004). A subs an ial numbe o species ha e been
desc ibed om ca es. To da e, A. bougisi Cassagnau & Delama e Debou e ille,
1955 is he only species o he genus Ache on iella eco ded om I an, whe e
i was ound in he soil o whea ields in Khuzes an p o ince (Meh a ooz
May an e al. 2023).
I an is a semi-a id coun y in Sou hwes Asia co e ing app oxima ely 1.64
million km2 and zoogeog aphically assigned o he Palea c ic egion (Lomolino
e al. 2017; Azizi Jalilian e al. 2020; Meh a ooz May an e al. 2021). The
Albo z Moun ain ange in he no he n I an has biodi e si y- ich Hy canian
o es s, and he Kope Dagh moun ain ange in no heas e n I an is especially
ich in ka s ca es, he mos common ype o ca es wo ldwide (Cul e and
Academic edi o : Wanda M. Weine
Recei ed:
4 July 2025
Accep ed:
15 Augus 2025
Published:
9 Oc obe 2025
ZooBank: h ps://zoobank.o g/
BC96F05D-AB55-4370-A895-
CFB0E77B7F18
Ci a ion: Meh a ooz May an M,
Ko áč Ľ (2025) Two new species o
he genus Ache on iella (Collembola,
Hypogas u idae) om ca es o he
Albo z and Kope Dagh moun ains in
I an. ZooKeys 1255: 1–25. h ps://doi.
o g/10.3897/zookeys.1255.164170
ZooKeys 1255: 1–25 (2025)
DOI: 10.3897/zookeys.1255.164170
2
ZooKeys 1255: 1–25 (2025), DOI: 10.3897/zookeys.1255.164170
Mahmood Meh a ooz May an & Ľubomí Ko áč: Two new oglobion species o Ache on iella om Sou hwes Asia
Pipan 2009; Mema iani 2020; Da oudnia e al. 2024). This con ibu ion is
based on ecen speleobiological ieldwo k conduc ed in ca es o no he n
and no heas e n I an wi h e y limi ed a ailable da a o sub e anean auna.
In his pape , we epo on he disco e y o wo new species o he genus
Ache on iella. They we e ound in I anian ca es wi hin he Albo z (Hy canian
o es s) and Kope Dagh moun ain anges. Addi ionally, we p o ide an upda -
ed iden i ica ion key and a able summa izing he diagnos ic cha ac e is ics
o wo ld species o he genus Ache on iella.
Ma e ial and me hods
Ca e desc ip ions
Table 1
Aghol Ca e is a e y small ca e on a moun ain slope nea Kenan Ca es, in Ra-
za i Kho asan p o ince, close o Chena an ci y (36°33'36"N, 59°04'58"E). I is a
an al i ude o 1,445 m a.s.l. The ca e ea u es a co ido ha is 25 m long, wi h
a high ceiling, and one co ne con ains a la ge amoun o ba guano.
Al Ca e, loca ed in Raza i Kho asan p o ince, is abou 50 km eas o Mash-
had, nea Ka deh illage, in he sou he n sec ion o he Kope Dagh Moun ains
(36°40'0.2"N, 59°39'19"E). I is wi hin he Mozdou an geological o ma ion o
no heas e n I an. The ca e is a an ele a ion o 1,477 m a.s.l. and has wo
adjacen en ances on a moun ain slope in he middle o a alley. The i s , well-
shaped en ance leads o wo ela i ely la ge halls.
Bazangan Ca e is in no heas e n I an, in he Sa akhs egion o Raza i Kho-
asan p o ince, 140 km no heas o Mashhad (36°18'29"N, 60°22'19"E). I is
in he Kope Dagh moun ains a an al i ude o 1,401 m a.s.l.. I s en ance is
ela i ely la ge. F om he opening, he ca e con inues downwa d in a gen le
slope owa ds a lake a he bo om. The ceiling is ocky, while he loo is hea ily
co e ed wi h ba guano and bi d (mos ly pigeon) d oppings. In some pa s o
he ca e, he ceiling heigh eaches o 25 m. The ca e ex ends o 204 m.
Bidak Ca e is in No h Kho asan p o ince, 4 km sou heas o Bidak illage
and 12 km om Bojnou d ci y in an open a ea (37°26'30"N, 57°13'14"E). The
ca e en ance is loca ed a an ele a ion o 1,327 m a.s.l. I appea s a g ound
le el as a e ical sha esembling a well, wi h a s one-walled opening ap-
p oxima ely 6 m high. The ca e ex ends o abou 80 m in leng h, wi h he i s
20 m descending s eeply a an almos e ical angle, ollowed by a passage
ha leads o a co ido and a chambe a he e minal pa o he ca e.
Bu nik Ca e is wi hin a limes one o ma ion nea he ci y o Fi ouzkouh
and he illage o He andeh (35°41'14"N, 52°41'24"E) a an al i ude o ap-
p oxima ely 2,000 m a.s.l., abou 100 m abo e he Bu nik plain (Fig. 1). I s
en ance is on a s eep moun ain slope and, a 4022 m, is he longes ca e
(Fig. 2) in Teh an p o ince and he ou h longes ca e in I an. I ea u es a
la ge, iangula en ance, app oxima ely 15 m long and 10 m high (Fig. 10A).
The ca e con ains ou ex ensi e halls, wi h he S alagmi e Hall (Fig. 10B)
con aining a 30 m deep well nea i s owe ing cli s. The empe a u e and
ela i e humidi y we e measu ed in he e minal pa o he ca e in June
2024 using a TFA 30.5015 DTH he mo-hyg ome e ; he empe a u e was
ound o be 8 °C and humidi y was 87%.
3
ZooKeys 1255: 1–25 (2025), DOI: 10.3897/zookeys.1255.164170
Mahmood Meh a ooz May an & Ľubomí Ko áč: Two new oglobion species o Ache on iella om Sou hwes Asia
Danial Ca e is in he Albo z Moun ain ange and he Hy canian Fo es wi hin
he Mazanda an ka s a ea on he sou he n edge o he Caspian Sea in Salman
Shah , Abbasabad ci y (36°39'35"N, 51°10'53"E). The ca e is a an al i ude o
164 m a.s.l. The ca e has a leng h o 2,158 m, and a sub e anean i e lows
h oughou he ca e’s en i e leng h (Ghelich Khani 2017; Yuse inia 2017; Ba jadze
e al. 2024). I had cons an ai empe a u e o 16 °C and ela i e humidi y o e
84% in July 2024 (measu ed wi h a he mo-hyg ome e TFA 30.5015 DTH).
Ka deh Ca e, also known as Heza Dalan (Thousand co ido ), is 3 km om
Ka deh illage (36°40'15"N, 59°40'37"E) and 48 km om Mashhad ci y in Raza i
Kho asan p o ince. The en ance, loca ed a an ele a ion o 1,458 m a.s.l., ea-
u es a s one gap. The ca e is 425 m in leng h.
Kenan 1 Ca e, a an al i ude o 1451 m a.s.l., is in he Zoukenan egion, app ox-
ima ely 5 km sou h o Chena an ci y in Raza i Kho asan p o ince (36°33'58"N,
59°04'19"E). Al hough mos o he ca e emains d y, humidi y can be obse ed
in some a eas du ing he ainy season. The ca e is 541 m in leng h and is di id-
ed in o wo sepa a e loo s. I has six en ances and con ains h ee wells.
Moghan Ca e is 35 km sou hwes o Mashhad in Raza i Kho asan P o ince
a an al i ude o 2,193 m a.s.l. in eas e n pa o he Kope Dagh Moun ains
(36°06'59"N, 59°22'06"E) (Figs 9, 10C). The ca e is app oxima ely 500 m long
and has a maximum dep h o 44 m. I ea u es i e chambe s in wo loo s. The e
is a wa e pool in he e minal pa o he i h chambe on he second loo .
This pool measu es 5.66 × 6.93 m and has a dep h o 0.3–0.6 m (Meh a ooz
May an e al. 2024). The ai empe a u e in his chambe is a cons an ly 9 °C,
and he ela i e humidi y exceeds 84% in July 2024 (measu ed by HTC-2 digi al
he mo-hyg ome e ).
Me hods
Collembola, including Ache on iella specimens, we e collec ed ca e walls, sedi-
men , o en wood, and ba guano using an aspi a o . Pi all aps equipped wi h
plas ic cups con aining p opylene glycol we e used as a complemen a y collec ion
me hod. Fu he mo e, om wooden sha ings ha we e used as a bai exposed in
a ca e o abou six mon hs, he specimens we e ex ac ed in Be lese unnels in
he labo a o y o he Zoology Museum o Fe dowsi Uni e si y o Mashhad in I an.
In he labo a o y, he specimens we e placed in con aine s illed wi h 96% e hyl
alcohol o p ese a ion. Fo obse a ion unde an op ical mic oscope wi h phase
Table 1. Lis o I anian ca es wi h eco ds o he new species o Ache on iella. Al i ude is abo e mean sea le el.
Ca e name P o ince GPS Coo dina es (La i ude, Longi ude) Leng h Al i ude
(m a.s.l.) Tempe a u e (°C) Rela i e
humidi y (%)
Aghol Raza i Kho asan 36°33'36"N, 59°04'58"E 25 1445 — —
Al Raza i Kho asan 36°40'0.2"N, 59°39'19"E 75 1447 — —
Bazangan Raza i Kho asan 36°18'29"N, 60°22'19"E 204 1401 — —
Bidak No h Kho asan 37°26'30"N, 57°13'14"E 80 1237 — —
Bu nik Teh an 35°41'14"N, 52°41'24"E 4022 2000 8 87
Danial Mazanda an 36°39'35"N, 51°10'53"E 2158 164 16 84
Ka deh Raza i Kho asan 36°40'15"N, 59°40'37"E 425 1458 — —
Kenan 1 Raza i Kho asan 36°33'58"N, 59°04'19"E 541 1451 — —
Moghan Raza i Kho asan 36°06'59"N, 59°22'06"E 500 2193 9 84
4
ZooKeys 1255: 1–25 (2025), DOI: 10.3897/zookeys.1255.164170
Mahmood Meh a ooz May an & Ľubomí Ko áč: Two new oglobion species o Ache on iella om Sou hwes Asia
Figu e 1. A. Loca ion o Teh an p o ince in I an ( ed) and he Bu nik Ca e ( ed as e isk), ype locali y o A. nezamdoos i
sp. no .; B. En ance o he Bu nik Ca e (pho o: M. Meh a ooz May an).
con as , he ini ial specimens we e moun ed in Heinz’s medium a e clea ing in
Nesbi ’s luid. To compa e he e ec i eness o di e en clea ing me hods, addi-
ional specimens we e gen ly boiled in a glass e apo a ing dish con aining 96%
e hyl alcohol on an elec ic ho pla e o app oxima ely 1 min o emo e a om
he body. Fo clea ing, he specimens we e anspo ed o conca e glass dish wi h
10% aqueous solu ion o KOH o 1 min, a e which hey we e ans e ed o a
dish con aining chlo ophenol un il hey became anspa en . Finally, he specimens
we e moun ed in Swann medium. Once d y, he co e glasses we e sealed wi h nail
polish o Canada balsam o p o ec ion o he moun ing medium agains d ying.
The specimens we e obse ed in Leica DM 2500 ligh mic oscope equipped wi h
phase and DIC con as s, and a d awing ube.
Abb e ia ions o mo phological e m used in he ex and igu es a e as ollows:
Abd abdominal segmen
An an ennal segmen
An IIIO enso y o gan An III
ASp anal spines
Cl claws
sm se a e mesose a
ms mic osensillum
os subapical o gani e (sub-apical senso y peg)
PAO pos -an ennal o gan
S sensillum/sensilla
ss senso y se ae
Tib ibio a sus
VT en al ube
5
ZooKeys 1255: 1–25 (2025), DOI: 10.3897/zookeys.1255.164170
Mahmood Meh a ooz May an & Ľubomí Ko áč: Two new oglobion species o Ache on iella om Sou hwes Asia
Ins i u ional abb e ia ions:
CoPJSU Collembola collec ion o he Depa men o Zoology, Ins i u e o Bi-
ology and Ecology, Facul y o Science, Pa ol Joze Ša á ik Uni e si y,
Košice, Slo akia;
ZMFUM Zoology Museum o Fe dowsi Uni e si y o Mashhad, Mashhad, I an
Resul s
Taxonomy
Class Collembola Lubbock, 1873
O de Podu omo pha Bö ne , 1913
Family Hypogas u idae Bö ne , 1906
Genus Ache on iella Absolon, 1913
Diagnosis. Small Hypogas u idae wi h a usi o m habi us. Whi e species, wi h-
ou eyes, pos -an ennal o gan, e inaculum and u ca. Tibio a si wi h 16–20
se ae, including 1 o 2 knobbed o poin ed enen hai s. Claws wi h o wi hou
ee h. Se al pa e n o ype I, wi h mesose ae mo e o less se a ed and senso y
se ae some imes longe and smoo h (Thibaud e al. 2004).
Type species. Ache on iella onychiu i o mis Absolon, 1913.
New axa
Ache on iella nezamdoos i Meh a ooz May an & Ko áč, sp. no .
h ps://zoobank.o g/1206FFB3-B5C4-481B-8809-F8EA9CE3122F
Figs 3–8, 17A, Table 2
Diagnosis. Colou whi e, wi hou pigmen . Eyes, PAO, u ca and e inaculum ab-
sen . Lab um wi h 4/554 se ae. Fou an e io se ae on lab um hickened. An
IV wi h simple apical esicle and 4 subapical subglobula sensilla. VT wi h 4+4
apical se ae. Tibio a si wi h 2 knobbed enen hai s. Claws wi h s ong in e nal
subapical oo h, empodium absen . Two small anal spines p esen . Se ae m1,
m4, and m5 on Abd IV p esen .
Type ma e ial. Holo ype. • ♂ on slide; I an, Teh an p o ince, Fi ouzkouh ci y,
He andeh illage, Bu nik Ca e; 35°41'14"N, 52°41'24"E; 2000 m a.s.l.; 02 Aug.
2024; M. Meh a ooz May an leg.; da k zone, collec ed om su ace o wa e ;
empe a u e = 8 °C, ela i e humidi y = 87.0%; slide no. I BuAc110. Pa a ypes.
• 5 ♀♀, 2 ju eniles, on slides; I an, same collec ion da a as o holo ype; slide
no. I BuAc111–117; Holo ype and 5 pa a ypes deposi ed in CoPJSU, 2 pa a-
ypes (I BuAc116–117) deposi ed in ZMFUM.
Addi ional ma e ial. • 4 ♂♂, 5 ♀♀, on slides; I an, Mazanda an p o ince,
Abbas Abad Coun y, close o he Danial illage, Danial Ca e; 36°39'35"N,
51°10'53"E; 203 m a.s.l.; 13 Jul. 2024; M. Meh a ooz May an leg.; da k zone,
deep sec ion o he ca e, Rizan chambe , collec ed om su ace o wa e ; em-
pe a u e = 16 °C, ela i e humidi y = 84.0%; slide no. I DaAc118–126; deposi ed
in CoPJSU.
6
ZooKeys 1255: 1–25 (2025), DOI: 10.3897/zookeys.1255.164170
Mahmood Meh a ooz May an & Ľubomí Ko áč: Two new oglobion species o Ache on iella om Sou hwes Asia
• 10 ♂♂, 8 ♀♀, on slides; I an, No h Kho asan p o ince, Bojnou d ci y, close o
he Bidak illage, Bidak Ca e; 37°26'30"N, 57°13'14"E; 1327 m a.s.l.; 23 Jun. 2022
and 26 Jul. 2023; M. Meh a ooz May an leg.; da k zone, e minal pa o he
ca e, collec ed om ba guano; slide no. I BiAc127–145; deposi ed in CoPJSU.
• 7 ♂♂, 3 ♀♀, on slides; I an, Raza i Kho asan p o ince, Sa akhs coun y, close
o Bazangan illage, Bazangan Ca e; 36°18'29"N, 60°22'19"E; 1401 m a.s.l.;
22 No . 2021, 18 Jun. 2022 and 27 Sep. 2023; M. Meh a ooz May an leg.; da k
zone, e minal pa o he ca e, collec ed om ba guano, o en wood and pi -
all aps wi h p opylene glycol; slide no. I BaAc146–155; deposi ed in CoPJSU.
• 2 ♂♂, 2 ♀♀, on slides; I an, Raza i Kho asan p o ince, Chena an ci y, nea he
indus ial own, Aghol Ca e; 36°33'36"N, 59°04'58"E; 1445 m a.s.l.; 07 Jul. 2022
and 08 Aug. 2023; M. Meh a ooz May an leg.; da k zone, e minal pa o he
ca e, collec ed om ba guano; slide no. I AgAc156–159; deposi ed in CoPJSU.
• 16 ♂♂, 1 ju enile, on slides; I an, Raza i Kho asan p o ince, Mashhad ci y,
Ka deh illage, Ka deh Ca e; 36°40'15"N, 59°40'37"E; 1458 m a.s.l.; 01 Jul. 2022,
10 Jul. 2023 and 18 Aug. 2024; M. Meh a ooz May an leg.; da k zone, collec ed
om ba guano in Ba chambe and om o en wood in 2nd ca e loo ; empe a u e
= 16 °C, ela i e humidi y = 80.0%; slide no. I KaAc160–176; deposi ed in CoPJSU.
Figu e 2. G ound plan o he Bu nik Ca e (ske ched by E. Geye ).
7
ZooKeys 1255: 1–25 (2025), DOI: 10.3897/zookeys.1255.164170
Mahmood Meh a ooz May an & Ľubomí Ko áč: Two new oglobion species o Ache on iella om Sou hwes Asia
Table 2. Di e en ial cha ac e s o wo ld species o Ache on iella. Cha ac e s: 1, Anal spines: 0 = absen , 1 = p esen ; 2, Tibio a sal enen hai (s) numbe ; 3, Tibio a sal
enen hai o m: 1 = poin ed, 2 = knobbed; 4, An IV sensilla numbe ; 5, in e nal oo h on claw: 0 = absen , 1 = weekly de eloped, 2 = s ong; 6, VT – numbe o apical
se ae; 7, Th I m se ae; 8, Th II–III a ow se ae; 9, Th II–III m ow se ae; 10, Th II–III p ow se ae; 11, Abd I–III a ow se ae; 12, Abd I–III p ow se ae; 13, Abd IV a ow se ae;
14, Abd IV m ow se ae; 15, Abd IV p ow se ae; 16, Abd V a ow se ae; 17, Abd V m ow se ae; 18, Abd V p ow se ae. “-“= absence o da a.
Species/Cha ac e s 123 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18
A. aokii Tamu a & Yue, 1999 10 – 4 0 4+4 1 3 4 1 3 4 5 1 3 4 1 3 4(s) 1 3 4 5 1–5(s) 1 2 5 0 1–4(s) 1 3 0 1–3(s)
A. bougisi Cassagnau & Delama e Debou e ille,
1955
11–2 2 4 14+4 1 3 4 1 2 4 5 1 3 5 1–4(s) 1 2 3 4 5 1–5(s) 1 2 4 5 1 3 5 1–5(s) 1 3 0 1–3 (s)
A. candida (Delama e Debou e ille, 1952) 0 0 – 5 0 4+4 1 3 4 1 2 4 5 1 3 5 1–4(s) 1 2 3 4 5 1–5(s) 1 2 4 5 1 3 5 1–4(s) 1 3 0 1–3(s)
A. ca usoi Dallai, 1978 0 2 2 4 2 4+4 1 (2) 3 4 1 2 4 5 1 4 5 1 2 4(s) 1 2 4 5 1 2 3 5(s) 1 2 4 5 4 1–4(s) 1 2 3 0 1–3(s)
A. cassagnaui Thibaud, 1967 0 2 2 4 2 4+4 1 3 4 1 2 4 5 1 4 5 1 2 4(s) 1 2 3 4 5 1–5(s) 1 2 4 5 1 4 5 1–4(s) 1 2 3 1 3 1–3(s)
A. ca e nicola (Ta sia in Cu ia, 1941) 12 2 4 0 4+4 – – – – – – – – – – 0 –
A. colo lipana Palacios-Va gas & Thibaud, 1985 0 1 1 5 0 4+4 1 3 4 1 2 4 5 1 4 5 1–4(s) 1 2 3 4 5 1–5(s) 1 2 4 5 1 3 1–4(s) 1 3 0 1–3(s)
A. den a a Djanasch ili, 1971 0 2 2 4 2 4+4 1 3 4 1 2 4 5 1 4 5 1 2 4(s) 1 2 3 4 5 1–5(s) 1 2 4 5 2 1–4(s) 1 2 3 0 1–3(s)
A. epigea Bone , 1945 112 4 2 4+4 1 3 4 1 2 3 4 5 1 3 4 1–4(s) 1 2 3 4 5 1–5(s) 1 2 4 5 1 3 5 1–4(s) 1 3 0 1–3(s)
A. globula a Thibaud & Massoud, 1980 0 1 1 4 0 4+4 1 3 4 1 2 4 5 1 3 5 1–4(s) 1 2 4 5 1–5(s) 1 2 4 5 1 4 1–4(s) 1 3 0 1–2(s)
A. kowalskio um Weine & Naj , 1998 0 0 – 5 0 4+4 1 3 4 1 2 4 5 1 4 5 1–4(s) 1 2 3 4 5 1–5(s) 1 2 4 5 1 3 5 1–5(s) 1 3 0 1–3(s)
A. mac (Palacios-Va gas & Thibaud, 1985) 111 4 0 4+4 1 3 4 1 2 4 5 1 4 5 1 2 4(s) 1 2 4 5 1–5(s) 1 2 4 5 3 5 1–4(s) 1 3 0 1–3(s)
A. massoudi Thibaud, 1963 0 2 2 8 0 4+4 1 3 4 1 2 4 5 4 1 2 4(s) 1 2 3 5 1 2 3 5(s) 1 2 4 5 3 1 2 4(s) 1 2 3 0 1–2(s)
A. nezamdoos i sp. no . 12 2 4 2 4+4 1 3 4 1 2 4 5 1 4 5 1–4(s) 1 2 3 4 5 1–5(s) 1 2 4 5 1 4 5 1–4(s) 1 2 3 0 1–3(s)
A. onychiu i o mis Absolon, 1913 0 1 1 4 0 4+4 1 3 4 1 2 4 5 1 4 5 1 2 4(s) 1 2 3 4 5 1–5(s) 1 2 4 5 1 4 5 1–4(s) 1 2 3 1 3 1–3(s)
A. palaciosi sp. no . 0 2 2 4 2 4+4 1 4 1 2 4 5 1 4 1 2 4(s) 1 2 3 4 5 1–5(s) 1 2 4 5 4 (5) 1–4(s) 1 3 0 1–3(s)
A. p ominen ia Thibaud & Weine , 1997 10 – 6 0 5+5 1 3 4 1 2 4 1 3 4 1 3 4(s) 1 2 3 5 1–5(s) 1 2 4 5 3 4 1–4(s) 1 2 3 0 1–2(s)
A. sabina Bone , 1945 12 2 4 2 4+4 1 2 4 1 2 4 5 1 3 5 1–4(s) 1 2 3 4 5 1–6(s) 1 2 4 5 1 3 4 1–4(s) 1 2 3 0 1–3(s)
A. hai Thibaud, 1990 111 4 0 4+4 1 3 4 1 2 4 5 1 3 5 1–4(s) 1 3 4 5 1 2 4 5(s) 1 2 4 5 0 1 3 4(s) 1 3 0 2(s)
A. hibaudi Ba a, 1994 10 – 5 0 5+5 1 3 4 1 (2) 3 4 5 1 3 4 1 2 4(s) 1 2 3 4 5 1–5(s) 1 2 4 5 3 5 1–4s) 1 3 2 1–3(s)
A. a iabilis Delama e Debou e ille, 1948 0 2 2 4–7 0 4+4 1 3 4 1 2 4 5 II 1 (4)
III (1) 4
1 2 4(s) 1 2 3 4 5 I–II 1–4(s)
III 1–5(s)
1 2 4 3 1 2 4(s) 1 2 3 0 1–2(s)
A. xenylli o mis Gisin, 1952 0 2 2 4 14+4 1 3 4 1 2 4 5 1 4 5 1–4(s) 1 2 3 4 5 1–5(s) 1 2 3 (3) 4 1–5(s) 1 2 3 3 1–3(s)
8
ZooKeys 1255: 1–25 (2025), DOI: 10.3897/zookeys.1255.164170
Mahmood Meh a ooz May an & Ľubomí Ko áč: Two new oglobion species o Ache on iella om Sou hwes Asia
Figu e 3. Ache on iella nezamdoos i sp. no . A. An I–IV ( igh an enna), se al pa e n; B. An III senso y o gan; C. Head,
se al pa e n o do sal side. Scale ba s: 0.01 mm (A); 0.05 mm (B); 0.025 mm (C).
Figu e 4. Ache on iella nezamdoos i sp. no . A. Head, se al pa e n o en al side; B. Lab um, se al pa e n; C. Labium
and labial iangle, se al pa e n; D. Mandibles; E. Maxilla y ou e lobe. Scale ba s: 0.025 mm (A); 0.01 mm (B–E).
9
ZooKeys 1255: 1–25 (2025), DOI: 10.3897/zookeys.1255.164170
Mahmood Meh a ooz May an & Ľubomí Ko áč: Two new oglobion species o Ache on iella om Sou hwes Asia
Figu e 5. Ache on iella nezamdoos i sp. no . A. An IV sensilla; B. An IV apical esicle; C. Lab al se ae; D. Se a e mesose ae
om Abd I; E. Claw wi h subapical in e nal oo h; F. Knobbed enen hai on ibio a sus. Scale ba s: 0.01 mm (A–F).
16
ZooKeys 1255: 1–25 (2025), DOI: 10.3897/zookeys.1255.164170
Mahmood Meh a ooz May an & Ľubomí Ko áč: Two new oglobion species o Ache on iella om Sou hwes Asia
collec ed om o en wood in 2nd ca e loo and om a s alagmi e column in
1s loo ; empe a u e = 9 °C, ela i e humidi y = 84.0%; slide no. I MoAc101.
Pa a ypes. • 4 ♂♂, 4 ♀♀, 1 ju enile, on slides; I an, same collec ion da a as o
holo ype; slide no. I MoAc 102–109; Holo ype and 8 deposi ed in CoPJSU, 1
pa a ype (I MoAc 109) deposi ed in ZMFUM.
Desc ip ion. Body. Leng h 1.0 mm on a e age (n = 9, maximum leng h
1.18 mm). Habi us and buccal cone ypical o he genus Ache on iella.
Eyes, PAO, e inaculum, and u ca absen . Body usi o m, co e ed by se a e
Figu e 13. Ache on iella palaciosi sp. no . A. An IV apical esicle; B. An IV sensillum; C. Two medial se ae in an e io ow
o lab um; D. Se a e mesose ae o Th II; E. Knobbed enen hai on ibio a si; F. Claw wi h subapical oo h. Scale ba s:
0.01 mm (A–F).
17
ZooKeys 1255: 1–25 (2025), DOI: 10.3897/zookeys.1255.164170
Mahmood Meh a ooz May an & Ľubomí Ko áč: Two new oglobion species o Ache on iella om Sou hwes Asia
Figu e 14. Ache on iella palaciosi sp. no . A–C. Legs I–III, se al pa e n o coxa, ochan e , emu and ibio a sus;
D–F. Legs I–III, oo complex. Scale ba s: 0.01 mm (A–F).
Figu e 15. Ache on iella palaciosi sp.no . Th I–III, se al pa e n o do sal side. Scale ba : 0.025 mm.
18
ZooKeys 1255: 1–25 (2025), DOI: 10.3897/zookeys.1255.164170
Mahmood Meh a ooz May an & Ľubomí Ko áč: Two new oglobion species o Ache on iella om Sou hwes Asia
mesose ae (Fig. 13D), anal spines absen . Colou whi e, wi hou aces o pig-
men . Cu icula g anula ion a he uni o m.
An ennae. An ennae sligh ly sho e han head, 0.18 mm and 0.20 mm, e-
spec i ely (Fig. 11A). Leng h o An I: II: III and IV (do sally used) as 0.034:
0.036: 0.11 mm, espec i ely. All se ae on An I–II sligh ly se a e. An I wi h 7
se ae; An II wi h 11 se ae. An III and IV do sally used, in e segmen al su u e
on en al side well ma ked, 26 do sal and 25 en al se ae, all no mal, smoo h
(la e al se ae longe han o he se ae). An IIIO wi h 2 cen al sho and cu ed
ods (mic osensilla) lanked by 2 long gua d sensilla o cha ac e is ic cons i u-
ion (Deha eng and Díaz 1984) (Fig. 11B), a small en ola e al mic osensillum
in a en al pi . An IV wi h a p o usible en oapical simple esicle (Fig. 13A),
4 globula sensilla (Fig. 13B) – 2 do sal (1 la e oex e nal, 1 la e oin e nal) and
2 en al (bo h la e oex e nal), do soex e nal subapical ms and apical os nea
ex e nal sensilla (see D’Haese 2003).
Head. Eyes and PAO absen . Se al o mula o lab um 4/5,5,4, wo medi-
al se ae in an e io ow on lab um hickened as spines and apically den a e
(Figs 12B, 13C). Labium and labial iangle as in Fig. 12C, wi h 5 main papillae
(A–E), 6 p oximal smoo h se ae, 4 basomedial and 5 basola e al smoo h se ae.
Maxilla y ou e lobe wi h 1 sublobal hai (Fig. 12E). Le mandible wi h 3 ee h
and igh mandible wi h 4 ee h (Fig. 12D).
Legs. Legs I–III wi h no mal se ae, ibio a si wi h 2 knobbed enen hai s
(Fig. 13E). Se al pa e n as in Fig. 14A–C. Leg I: coxa wi h 3 se ae, ochan e
wi h 6 se ae, emu wi h 12 se ae, Tib wi h 18, including 2 knobbed apical enen
Figu e 16. Ache on iella palaciosi sp.no . A. Abd I–VI se al pa e n o do sal side; B. Abd I–VI se al pa e n o en al side.
Scale ba s: 0.025 mm (A, B).
19
ZooKeys 1255: 1–25 (2025), DOI: 10.3897/zookeys.1255.164170
Mahmood Meh a ooz May an & Ľubomí Ko áč: Two new oglobion species o Ache on iella om Sou hwes Asia
hai s. Leg II: coxa wi h 8 se ae, ochan e wi h 6 se ae, emu wi h 12 se ae,
Tib wi h 18, including 2 knobbed apical enen hai s. Leg III: coxa wi h 7 se ae,
ochan e wi h 6 se ae, emu wi h 11 se ae, Tib wi h 17, including 2 knobbed
apical enen hai s; claw slende (Fig. 14D–F), in e nal edge wi h s ong sub-
apical oo h (Fig. 13F); empodium absen . P e a sus wi h 2 se ae (1 in e nal, 1
ex e nal), bo h eaching he hal leng h o in e nal lamella.
Abdomen. Ven al ube wi h 4+4 se ae. Re inaculum and u ca absen . Anal
spines absen , subs i u ed by 2 se a e se ae (Fig. 17B).
Se al pa e n. Do sal se al pa e n gene ally o ype I wi h mesose ae (a e
Jo dana e al. 1997). Se ae on head and body sligh ly se a e, senso y se ae
longe and smoo h.
Head. Do sal se al pa e n as in Fig. 11C. Se a a0 p esen ; d1–d5; sd3–sd5
(sd1 and sd2 absen ); 1– 2; c2–c5 (c1 absen ); p1–p5; oc1–oc2 (oc3 absen ).
Ven al se al pa e n as in Fig. 12A, all se ae smoo h.
Tho ax. Do sal se al pa e n as in Fig. 15. Posi ion o ss in p ow on Th II–III
is 4 and 4, espec i ely. Th I wi h se ae m1 and m4 [m2 and m3 absen ]; Th II
and III wi h 3 ows o se ae – a: 4+4 as a1, a2, a4 and a5 [a3 absen ]; m: 3+3
be ween m6(s) as m1, m4 and m5 [m2 and m3 absen ]; p: 2+2 be ween p4(s)
as p1 and p2 [p3 absen ]. Tho acic s e na wi hou se ae.
Abdomen. Do sal se al pa e n as in Fig. 16A. Posi ion o ss in p ow on Abd
I–V is 5, 5, 5, 4 and 3, espec i ely. Abd I–III wi h 2 ows o se ae – a: 5+5 as
a1–a5; p: 4+4 be ween p5(s) as p1–p4. Abd IV wi h 3 ows o se ae – a: 3+3
as a1, a2 and a4 [a3 absen ]; m: 1+1 (o 2+2) as m4 (m5) [m1, m2 and m3 ab-
sen ]; p: 3+3 be ween p4(s) as p1–p3. Abd V wi h 2 ows o se ae – a: 3+3 as
a1, a3 and a4 [a2 absen ]; p: 2+2 be ween p3(s) as p1–p2. Ven al abdominal
se al pa e n as in Fig. 16B.
Figu e 17. A. Ache on iella nezamdoos i sp. no . – wo small anal spines on Abd VI; B. Ache on iella palaciosi sp. no . –
wo se a e mesose ae in posi ion o anal spines on Abd VI. Scale ba s: 0.01 mm (A, B).
20
ZooKeys 1255: 1–25 (2025), DOI: 10.3897/zookeys.1255.164170
Mahmood Meh a ooz May an & Ľubomí Ko áč: Two new oglobion species o Ache on iella om Sou hwes Asia
E ymology. The new species is dedica ed o ou colleague and iend p o es-
so José G. Palacios-Va gas (Uni e sidad Nacional Au ónoma de México) o
his immense con ibu ion o axonomy o soil and sub e anean Collembola.
Taxonomic ema ks. Ache on iella palaciosi sp. no . is simila o 11 species
in lack o anal spines: A. candida (Delama e Debou e ille, 1952), A. ca usoi
Dallai, 1978, A. cassagnaui Thibaud, 1967, A. colo lipana Palacios-Va gas &
Thibaud, 1985, A. den a a Djanasch ili, 1971, A. globula a Thibaud & Massoud,
1979, A. kowalskio um Weine & Naj , 1998, A. massoudi Thibaud, 1963,
A. onychiu i o mis Absolon, 1913, A. a iabilis Delama e Debou e ille, 1948,
and A. xenylli o mis Gisin, 1952. Among hem, A. palaciosi sp. no ., A. ca usoi,
A. cassagnaui, A. den a a, A. massoudi, A. a iabilis, and A. xenylli o mis di e
om o he s by 2 knobbed enen hai s on ibio a si, while he es o he spe-
cies ha e 1–2 sha p enen hai s on ibio a si. Mo eo e , claws in A. palaciosi
sp. no ., A. ca usoi, A. cassagnaui and A. den a a a e equipped wi h a s ong
subapical oo h; howe e , claws a e oo hless in A. massoudi, A. a iabilis, and
A. xenylli o mis. In A. palaciosi sp. no ., A. cassagnaui, and A. den a a, 2 medial
se ae in an e io ow on lab um a e hickened as spines and apically den a e,
while in A. ca usoi all 4 an e io lab al se ae a e sha p and smoo h. Ache on iella
palaciosi sp. no . di e s om A. cassagnaui by Th I wi h 2+2 m se ae, absence
o se a a3 on Abd IV, p esence o se a m4 (m5) on Abd IV, and absence o se a
a2 and m se ae on Abd V. In A. cassagnaui 3+3 m se ae on Th I and se a a3 on
Abd IV a e p esen , as well as se ae m1 (m3) and m4 on Abd IV and se a a2 and
m se ae on Abd V. Finally, A. palaciosi sp. no . di e s om A. den a a by ollow-
ing cha ac e s: se ae a0 and sd3 a e p esen , while oc3 is absen on do sal side
o head, maxilla y ou e lobe wi h one sublobal hai , se a m3 on Th I is absen ,
se a m4 on Abd IV is p esen , and se a a2 on Abd V is absen . In con as , in
A. den a a se ae a0 and sd3 a e absen , while oc3 is p esen on he do sal side
o head; maxilla y ou e lobe wi h 2 sublobal hai s, se a m3 on Th I is p esen ,
Abd IV wi h se a m2 in he m ow, and se a a2 on Abd V is p esen .
Iden i ica ion key o wo ld species o he genus Ache on iella
The ollowing iden i ica ion key is based on he diagnos ic cha ac e s o all de-
sc ibed Ache on iella species (adap ed om Dallai 1978; Bone 1945; Thibaud
1990; Thibaud e al. 2004).
1 ASp absen .....................................................................................................2
– ASp p esen .................................................................................................13
2 Tib wi hou o wi h 1 poin ed enen hai .....................................................3
– Tib wi h 2 mo e o less knobbed enen hai s .............................................7
3 An IV wi h 4 sensilla .....................................................................................4
– An IV wi h 5 sensilla .....................................................................................5
4 Two an e io medial lab al se ae hickened o den a e spines. Abd V wi h
1–3 se ae in m ow. Cl gene ally wi hou ee h, some imes wi h a small
en oapical in e nal oo h ..............................................................................
........................................A. onychiu i o mis Absolon, 1913 (Alge ia, ca e)
– An e io lab al se ae no mal. Abd V wi hou se ae in m ow. Cl wi hou
ee h .................................................................................................................
....... A. globula a Thibaud & Massoud, 1979 (Ma ie-Galan e Island, ca e)
21
ZooKeys 1255: 1–25 (2025), DOI: 10.3897/zookeys.1255.164170
Mahmood Meh a ooz May an & Ľubomí Ko áč: Two new oglobion species o Ache on iella om Sou hwes Asia
5 Tib wi h 1 poin ed enen hai . Tib I–III wi h 18, 18, and 17 se ae, espec-
i ely. Subcoxae “2” wi h 1, 2, and 2 se ae .....................................................
............... A. colo lipana Palacios-Va gas & Thibaud, 1985 (Mexico, ca e)
– Tib wi hou enen hai . Tib I–III wi h 19, 19, and 18 se ae, espec i ely.
Subcoxae “2” wi h 1, 2, and 3 se ae .............................................................6
6 Abdominal s e num IV wi hou se a m1 ........................................................
...........................A. candida (Delama e Debou e ille, 1952) (F ance, soil)
– Abdominal s e num IV wi h se a m1 ..............................................................
.....................................A. kowalskio um Weine & Naj , 1998 (Kenya, soil)
7 Cl wi h s ong subapical oo h ......................................................................8
– Cl wi hou ee h o wi h 1 weakly de eloped oo h ...................................11
8 Two lab al an e io medial se ae hickened in o den a e spines ...............9
– Fou lab al an e io se ae smoo h and equally long and hick .....................
............................................................... A. ca usoi Dallai, 1978 (I aly, ca e)
9 Abd IV wi hou se a m1. Abd V wi hou se ae in m ow ...........................10
– Abd IV wi h se a m1. Abd V wi h 1–3 se ae in m ow ...................................
.............................A. cassagnaui Thibaud, 1967 (F ance & Romania, ca e)
10 Do sal head wi hou se ae a0 and sd3. Th I wi h se a m3. Maxilla y ou e
lobe wi h 2 sublobal hai s. Abd V wi h se a a2 .............................................
.........................................A. den a a Djanasch ili, 1971 (Aze baijan, ca e)
– Do sal head wi h se ae a0 and sd3. Th I wi hou se a m3. Maxilla y ou e
lobe wi h 1 sublobal hai . Abd V wi hou se a a2 ..........................................
.................................................................... A. palaciosi sp. no . (I an, ca e)
11 Fou lab al an e io se ae smoo h and equally long and hick .....................
............................................. A. massoudi Thibaud, 1963 (I o y coas , soil)
– Two lab al an e io medial se ae hickened in o den a e spines .............12
12 An IV wi h 4–7 sensilla. Th II–III wi h se a m1 in m ow. Cl wi hou ee h .
............................A. a iabilis Delama e Debou e ille, 1948 (F ance, soil)
– An IV wi h 4 sensilla. Th II–III wi hou se a m1 in m ow. Cl wi h 1 small
in e nal oo h de eloped a 2/3 o he lamella ..............................................
.................A. xenylli o mis Gisin, 1952 (Mo oco, Po ugal & Spain, ca e)
13 Cl wi h a s ong oo h ..................................................................................14
– Cl wi hou ee h ...........................................................................................17
14 Tib wi h 2 knobbed enen hai s .................................................................15
– Tib wi h 1 mo e o less knobbed enen hai .............................................16
15 An IV wi h 4 sho subglobula sensilla. Th1 wi h se ae m1, m3, and m4 in m
ow. Abd I–III wi h ss in posi ion p5 ........ A. nezamdoos i sp. no . (I an, ca e)
– An IV wi h 4 elonga ed cylind ical sensilla. Th1 wi h se ae m1, m2, and
m4 in m ow. Abd I–III wi h ss in posi ion p6 ................................................
...............................................A. sabina Bone , 1945 (Mexico & USA, ca e)
16 An IV wi h 4 cylind ical and hick sensilla, pa ially hidden by egumen a y
olds. Abd IV wi h se ae m1, m3, and m4 in m ow ...................... A. bougisi
Cassagnau & Delama e Debou e ille, 1955 (Lebanon, soil & ca e)
– An IV wi h 4 cylind ical cu ed sensilla no concealed by egumen a y
olds. Abd IV wi h se ae m1, m4, and m5 in m ow .......................................
...........................................................A. epigea Bone , 1945 (Mexico, ca e)
17 Tib wi h 2 mo e o less knobbed enen hai s ...............................................
.......................................A. ca e nicola (Ta sia in Cu ia, 1941) (I aly, ca e)
– Tib wi hou o wi h 1 poin ed enen hai ...................................................18
22
ZooKeys 1255: 1–25 (2025), DOI: 10.3897/zookeys.1255.164170
Mahmood Meh a ooz May an & Ľubomí Ko áč: Two new oglobion species o Ache on iella om Sou hwes Asia
18 An IV wi h 6 sensilla. VT wi h 5+5 se ae ...................................................19
– An IV wi h 4 sensilla. VT wi h 4+4 se ae ...................................................20
19 Apical esicle on An IV ilobed. Abd V wi h ss in posi ion p2 ....................
........A. p ominen ia Thibaud & Weine , 1997 (New Caledonia Island, soil)
– Apical esicle on An IV simple. Abd V wi h ss in posi ion p3 ......................
.................................................. A. hibaudi Ba a, 1994 (Sou h A ica, soil)
20 Fou lab al an e io se ae hickened in o den a ed spines. Abd V wi h ss in
posi ion p2 ...................................... A. hai Thibaud, 1990 (Thailand, ca e)
– Fou lab al an e io se ae no den a e. Abd V wi h ss in posi ion p3 .......21
21 Tib I–III wi h 19, 19, and 18 se ae, espec i ely. An I wi h 7 se ae. Fou
lab al an e io se ae sha p, smoo h and equal ..............................................
..........................A. mac (Palacios-Va gas & Thibaud, 1985) (Mexico, soil)
– Tib I–III wi h 15, 15, and 14 se ae, espec i ely. An I wi h 6 se ae. Fou
lab al an e io se ae s ou ..........A. aokii Tamu a & Yue, 1999 (China, soil)
Discussion
Ache on iella is a cosmopoli an genus o he amily Hypogas u idae
(Podu omo pha) including bo h edaphic and sub e anean species. O he
22 known species o Ache on iella wo ldwide, 13 a e ca e-dwelling and
eigh a e soil-dwelling. No ably, A. bougisi is he only species ha has been
collec ed om en i onmen s bo h wi hin and ou side o ca es (Cassagnau
and Delama e Debou e ille 1955; Bellinge e al. 1996–2025). Among
he sub e anean species, eigh species, including A. nezamdoos i sp.
no ., A. cassagnaui, A. colo lipana, A. globula a, A. onychiu i o mis, A. hai,
A. a iabilis, and A. xenylli o mis ha e been ound on ba guano. This is in
con as o A. nezamdoos i sp. no . and A. palaciosi sp. no ., which ha e been
ound on o en wood in Moghan Ca e wi h a gene al lack o guano. Un o u-
na ely, due o he excessi e numbe o isi o s o his ca e in di e en sea-
sons and he lack o p ope p o ec ion laws, he popula ion o ba s in his
ca e has become e y a e. Also, A. nezamdoos i sp. no . is limi ed o deep
sec ions o ca es and, hus, can be conside ed o be a oglobio ic species.
Acknowledgemen s
We would like o hank p o essional speleologis Hamid Nezamdoos and Vahid
Ash a i o p epa ing he map and pho o o he ca e. Also, we a e g a e ul o Ka-
oshga an Ca ing Club in Mashhad, Hamid Sha i i, Sahand Sa a i, Sabe Kalho ,
Reza Hassani, and Ali Fa hi Loshani o assis ance du ing he expedi ions o he
ca es in no h and no heas e n I an. We also hank o Mohammad Ali Ja i (head
o he climbing boa d o Abbas Abad Coun y, I an), D Saeid Esmaeilpou , Sahand
Sa a i, and Mohammad Esmaeilpou o expedi ions o Danial Ca e. We a e g a e-
ul o hank wo anonymous e e ees o aluable commen s on ou manusc ip .
Addi ional in o ma ion
Con lic o in e es
The au ho s ha e decla ed ha no compe ing in e es s exis .
23
ZooKeys 1255: 1–25 (2025), DOI: 10.3897/zookeys.1255.164170
Mahmood Meh a ooz May an & Ľubomí Ko áč: Two new oglobion species o Ache on iella om Sou hwes Asia
E hical s a emen
No e hical s a emen was epo ed.
Use o AI
No use o AI was epo ed.
Funding
The s udy was suppo ed om he Slo ak Resea ch and De elopmen Agency, p ojec
APVV-21-0379, and om he Slo ak Scien i ic G an Agency, p ojec VEGA 1/0438/22.
The i s au ho was suppo ed om he pos doc o al p og am o he Facul y o Science,
Pa ol Joze Ša á ik Uni e si y in Košice, Slo akia.
Au ho con ibu ions
Mahmood Meh a ooz May an: Collec ing and so ing o specimens, mo phological
analysis o specimens, obse ing o specimens unde mic oscope, line d awings p epa-
a ion, w i ing o manusc ip . Ľubomí Ko áč: obse ing o specimens unde mic o-
scope, e iew and edi ing o manusc ip .
Au ho ORCIDs
Mahmood Meh a ooz May an h ps://o cid.o g/0000-0002-7494-0981
Ľubomí Ko áč h ps://o cid.o g/0000-0001-8194-2128
Da a a ailabili y
All o he da a ha suppo he indings o his s udy a e a ailable in he main ex .
Re e ences
Absolon K (1913) Uebe eine neue sub e ane Collembole (Insec a Ap e ygo a)—
Ache on iella onychiu i o mis n. g. n. sp. aus den Höhlen Algie s. A chi es de Zoologie
Expé imen ale e Géné ale 51(1): 1–7.
Azizi Jalilian M, Shayes eh K, Danehka A, Salmanmahiny A (2020) A new
ecosys em-based land classi ica ion o I an o conse a ion goals. En i onmen al
Moni o ing and Assessmen 192: 1–17. h ps://doi.o g/10.1007/s10661-020-8145-1
Ba jadze S, Meh a ooz May an M, Pa imucho á A, Magh adze E, Ko áč Ľ (2024) Two
new species o Plu omu us Yosii (Collembola: Tomoce idae) om Geo gia and I an.
Zoo axa 5463(4): 559–572. h ps://doi.o g/10.11646/zoo axa.5463.4.6
Ba a JA (1994) Nou eaux collemboles podu omo phes de la p o ince du Na al (Rép.
Sud A icaine) (Insec a: Collembola). Re ue de Zoologie A icaine 108(2): 181–189.
Bellinge PF, Ch is iansen KA, Janssens F (1996–2024) Checklis o he Collembola o
he wo ld. h p://www.collembola.o g [accessed June 2025]
Bone F (1945) Nue os géne os y especies de hipogas ú idos de México (Colembolos).
Re is a de la Sociedad Mexicana de His o ia Na u al 6(1–2): 13–45.
Bö ne C (1906) Das Sys em de Collembolen nebs Besch eibung neue Collembolen des
Hambu ge Na u his o ischen Museums. Mi eilungen aus dem Na u his o ischen
Museum in Hambu g 23: 147–188.
Bö ne C (1913) Die Familien de Collembolen. Zoologische Anzeige 41: 315–322.
Cassagnau P, Delama e Debou e ille C (1955) Mission Hen i Coi ai (1951). 3.
Collemboles. A chi es de Zoologie Expé imen ale e Géné ale 91: 365–395.
24
ZooKeys 1255: 1–25 (2025), DOI: 10.3897/zookeys.1255.164170
Mahmood Meh a ooz May an & Ľubomí Ko áč: Two new oglobion species o Ache on iella om Sou hwes Asia
Cul e DC, Pipan T (2009) The Biology o Ca es and O he Sub e anean Habi a s. Ox o d
Uni e si y P ess, Ox o d, 256 pp.
D’Haese CA (2003) Homology and mo phology in Podu omo pha (Hexapoda,
Collembola). Eu opean Jou nal o En omology 101: 385–407. h ps://doi.
o g/10.14411/eje.2003.060
Dallai R (1978) Rice che sui Collemboli. XXIII. Una nuo a species ca e nicola della
Sicilia (Ache on iella ca usoi n. sp.). Animalia 5(113): 345–355.
Da oudnia B, Dadkhodaie A, Nade i R, an Slage en M, Pou kho shid Z, Ekhlasi M (2024)
Di e si y and eco-geog aphical dis ibu ion pa e n o some Aegilops species om
he Zag os and Albo z Moun ain anges o I an. Plan Ecology 225: 761–774. h ps://
doi.o g/10.1007/s11258-024-01426-5
Deha eng L, Díaz A (1984) Collemboles du Kenya. I: lis e des s a ions. II: Hypogas u idae.
Bulle in du Muséum Na ional d’His oi e Na u elle. Sec ion A. Zoologie, Biologie, e
Écologie Animales 6(2): 335–347. h ps://doi.o g/10.5962/p.285922
Delama e Debou e ille C (1948) Su la p ésence du gen e Ache on iella Absolon dans
les g o es de l’A dèche e du Ga d. No es Biospéologiques 3: 49–56.
Delama e Debou e ille C (1952) É ude sys éma ique. I. Collemboles. In: Delama e C, Paulian
R (Eds) Faune des nids e e ie s en Basse Cô e d’I oi e. Leche alie , Pa is, 62–77.
Djanasch ili R (1971) Hypogas u idae (Collembola) de oid o eyes and pos an ennal
o gan om he T anscaucasian ca es. I. Zoologicheskij Zhu nal 50(5): 666–676.
[in Russian wi h English abs ac ]
Ghelich Khani P (2017) Danial Ca e. Zis Aein 6: 51–53. [in Pe sian]
Gisin H (1952) Collemboles écol és dans des g o es du moyen A las. Bulle in de la
Socié é des Sciences Na u elles du Ma oc 31: 53–56.
Jo dana R, A bea JI, Simón JC, Luciáñez MJ (1997) Fauna Ibe ica Vol. 8: Collembola,
Podu omo pha. Museo Nacional de Ciencias Na u ales, Consejo Supe io de
In es igaciones Cien í icas, Mad id, 1–807.
Lomolino MV, Riddle BR, Whi ake RJ (2017) Biogeog aphy: Biological Di e si y Ac oss
Space and Time. 5 h Edi ion. Sinaue Associa es, Sunde land, 759 pp. h ps://doi.
o g/10.1093/hesc/9781605354729.001.0001
Lubbock J (1873) Monog aph o he Collembola and Thysanu a. Ray Socie y, London.
h ps://doi.o g/10.5962/bhl. i le.11583
Meh a ooz May an M, Sadeghi-Namaghi H, Shayanmeh M, G eenslade P (2021) An
anno a ed ca alog o I anian Symphypleona and Neelipleona (Hexapoda: Collembola):
new eco ds and key o species. Jou nal o Asia-Paci ic Biodi e si y 14: 501–513.
h ps://doi.o g/10.1016/j.japb.2021.07.006
Meh a ooz May an M, G eenslade P, Sadeghi-Namaghi H (2023) An anno a ed checklis
o he Collembola (Hexapoda) om I an. Zoo axa 5275(1): 001–101. h ps://doi.
o g/10.11646/zoo axa.5275.1.1
Meh a ooz May an M, Pa imucho á A, Ko áč Ľ (2024) A new sub e anean species
o Oncopodu a Ca l & Lebedinsky, 1905 (Collembola, En omob yomo pha,
Oncopodu idae) om a ca e in no heas e n I an. Sub e anean Biology 49: 31–51.
h ps://doi.o g/10.3897/sub biol.49.118293
Mema iani F (2020) The Kho assan-Kope Dagh Moun ains. In: No oozi J (Ed.) Plan
Biogeog aphy and Vege a ion o High Moun ains o Cen al and Sou h-Wes Asia.
Plan and Vege a ion, Vol.17. Sp inge , Cham, 93–116. h ps://doi.o g/10.1007/978-
3-030-45212-4_3
Palacios-Va gas JG, Thibaud JM (1985) Nue os Hypogas u idae ano almos
(Collembola) de cue as y suelos de México. Folia En omologica Mexicana 66: 3–16.
25
ZooKeys 1255: 1–25 (2025), DOI: 10.3897/zookeys.1255.164170
Mahmood Meh a ooz May an & Ľubomí Ko áč: Two new oglobion species o Ache on iella om Sou hwes Asia
Tamu a H, Yue QY (1999) A new species o he genus Ache on iella om M . Yulongxue,
Yunnan, sou hwes China (Insec a: Collembola; Hypogas u idae). Na u al His o y
Bulle in o Iba aki Uni e si y 3: 1–3.
Ta sia in Cu ia I (1941) Due specie nuo e ed una poco no a di Collembola ca e nicoli
d’I alia. Annua io del Museo Zoologico della Uni e si à di Napoli 7(10): 1–7.
Thibaud JM (1963) Documen s su le gen e Ache on iella Absolon (Collembola) e
desc ip ion d’une espèce de Cô e d’I oi e. Annls Spéléoi 18(3): 287–297.
Thibaud JM (1967) Desc ip ion d’une espèce nou elle de Collembole Ache on iella
cassagnaui n. sp. Annales de Spélèologie 22: 393–400.
Thibaud JM (1990) Ré ision du gen e Ache on iella Absolon, 1913 (Insec a,
Collembola). Bulle in du Muséum Na ional d’His oi e Na u elle 4: 401–414. h ps://
doi.o g/10.5962/p.289587
Thibaud JM, Massoud Z (1979) Les collemboles des Pe i es An illes. 1—Hypogas u idae
e Onychiu idae (à sui e). Re ue d’Écologie e de Biologie du Sol 16(4): 547–557.
Thibaud JM, Weine WM (1997) Collemboles in e s i iels des sables de Nou elle-Calédonie.
Mémoi es du Muséum Na ional d’His oi e Na u elle 171: 63–89.
Thibaud JM, Schulz HJ, da Gama Assalino MM (2004) Hypogas u idae. In: Dunge W
(Ed.) Synopses on Palaea c ic Collembola, Vol. 4. Abhandlungen und Be ich e des
Na u kundemuseums Gö li z75(2): 1–287.
Weine WM, Naj J (1998) Collembola (En ogna ha) om Eas A ica. Eu opean Jou nal
o En omology 95: 217–237.
Yuse inia AA (2017) Ancien langa: Kela abad, Salmanshah and Abas Abad. Bolu
Publishe , Rash , 61 pp. [in Pe sian].
