Con e gen e olu ion unde g ound - wo new
oglomo phic collembolan species o he genus
Schae e ia Absolon, 1900 (Collembola, Podu omo pha,
Hypogas u idae) om I an and Mon eneg o
Ľubomí Ko áč1, Ma ko Ka aman2, Mahmood Meh a ooz May an1
1Depa men o Zoology, Ins i u e o Biology and Ecology, Facul y o Science, Pa ol Joze Ša á ik Uni e si y
in Košice, Š obá o a 2, 041 54 Košice, Slo akia 2Na u al His o y Museum o Mon eneg o, T g oj ode Beći
bega Osmana ića 16, Podgo ica, Mon eneg o
Co esponding au ho : Mahmood Meh a ooz May an (mahmood_meh a [email protected])
Academic edi o : Louis Deha eng|Recei ed 28 Ap il 2025|Accep ed 21 Oc obe 2025|Published 5 No embe 2025
h ps://zoobank.o g/F2343D01-6810-4530-8D62-CDBB6BEC97DF
Ci a ion: Ko áč Ľ, Ka aman M, Meh a ooz May an M (2025) Con e gen e olu ion unde g ound - wo
new oglomo phic collembolan species o he genus Schae e ia Absolon, 1900 (Collembola, Podu omo pha,
Hypogas u idae) om I an and Mon eneg o. Sub e anean Biology 54: 1–22. h ps://doi.o g/10.3897/
sub biol.54.157228
Abs ac
Two new sub e anean sp ing ail species o he genus Schae e ia Absolon, 1900 a e desc ibed: S. g eenslade-
ae sp. no . om Danial Ca e in he Albo z Moun ains, I an, and S. mon eneg ina sp. no . om Baba uša
Ca e in Dina ic Ka s , Mon eneg o. Bo h species a e cha ac e ized by he absence o eyes and pos an en-
nal o gan (PAO) ha ing mo e han 5 lobes in he con a y o o he eyeless congene s ha ha e less han 5
lobes in PAO. Mo phological ai s o bo h species indica e high le el o associa ion wi h he sub e anean
en i onmen . The phylogene ic ela ionships and mo phological adap a ions o wo new species o sub-
e anean li e a e discussed. Schae e ia g eensladeae sp. no . and S. mon eneg ina sp. no . se e as a good
example o con e gen e olu ion ha is cha ac e is ic o in e eb a es highly adap ed o deep sub e anean
ealm. An upda ed key o he wo ld species o he genus and a able summa izing diagnos ic cha ac e s o
he wo ld species o Schae e ia a e p o ided.
Keywo ds
Ca e sp ing ails, con e gen e olu ion, phylogeny, Schae e ia, axonomy
Sub e anean Biology 54: 1–22 (2025)
doi: 10.3897/sub biol.22.157228
h ps://sub biol.penso .ne
Copy igh Ľubomí Ko áč e al. This is an open access a icle dis ibu ed unde he e ms o he C ea i e Commons A ibu ion License (CC
BY 4.0), which pe mi s un es ic ed use, dis ibu ion, and ep oduc ion in any medium, p o ided he o iginal au ho and sou ce a e c edi ed.
RESEARCH ARTICLE
Sub e anean
Biology Published by
The In e na ional Socie y
o Sub e anean Biology
A pee - e iewed open-access jou nal
Ľubomí Ko áč e al. / Sub e anean Biology 54: 1–22 (2025)
2
In oduc ion
The genus Schae e ia Absolon, 1900 (Hypogas u idae) was i s es ablished o
S. emuc ona a, a species desc ibed om a ca e in he Czech Republic. The genus is
cha ac e ized by he ollowing ea u es: body pigmen educed, eyes om 0+0 o
7+7 (mos ly less han 5+5), u ca educed o absen - i p esen , hen dens usually
wi h 3 o 4 se ae (some imes 5 o 6), anal spines long (mo e o less equal o claw
in leng h), An IV wi h a simple apical bulb and 5 o 12 sensilla, body co e ed
wi h well-di e en ia ed se ae, Ti I–III wi hou knobbed enen hai s, claws wi h
o wi hou inne oo h, empodium well-de eloped wi h basal lamella. Mo eo e ,
he genus belongs o he Ce a ophysella den icula a line, i.e. i s ep esen a i es a e
cha ac e ized by a speci ic se al pa e n (Thibaud e al. 2004).
Schae e ia emuc ona a Absolon, 1900 is he nominal species o he genus, de-
sc ibed om Ka eřinská jeskyně Ca e in he Mo a ian Ka s , Czech Republic.
La e , S. willemi (Bone , 1930) was desc ibed om he Roche o and Han ca es
in Belgium and wo species om ca es in Ge many: S. decemocula a (S ach, 1939)
om Kelle höhle, Wes alia and S. quad iocula a (S ach, 1939) om Diebshöhle
and Heimkehle ca es in Ha z Moun ains. The i s species ha was ound o in-
habi su ace en i onmen was S. pouadensis Delama e Debou e ille, 1945 om
F ance. Schae e ia gue e ense (Bone , 1945) was he i s species epo ed ou side
Eu ope, disco e ed in Mexico in he popula Cacahuamilpa show ca e.
The genus has Hola c ic dis ibu ion (Thibaud 1972) wi h 35 species desc ibed
up o da e (Jo dana e al. 2020), in ol ing bo h edaphic and sub e anean species.
The ep esen a i es o he genus o en inhabi ca es in low al i udes, while humus,
soil and mosses in moun ains, being equen in ela i ely cold mic oclima e wi h
ai empe a u e a ound 10 °C and ela i ely high humidi y (Thibaud 1972). A
ecen s udy, ca ied ou by Jo dana e al. (2020) in Sie a de Gauda ama Moun-
ains, Spain, yielded a species associa ed wi h collu ial “milieu sou e ain supe i-
ciel” (MSS - meso oid shallow subs a um) documen ing a ie y o sub e anean
habi a s ha he congene s o his genus occupy.
In his pape , we p o ide comp ehensi e desc ip ions o wo new eyeless
Schae e ia species ha we e disco e ed du ing he ecen ca e explo a ions in I an
and Mon eneg o, wi h hei mo phological ai s showing high le el o associa ion
wi h he sub e anean en i onmen . A able summa izing diagnos ic cha ac e s o
eyeless Schae e ia congene s is included oge he wi h an upda ed key o he wo ld
species o he genus. The phylogene ic ela ionships and mo phological adap a-
ions o wo new species o sub e anean li e a e discussed.
Two new species o eyeless ca e nicolous Schae e ia 3
Ma e ial and me hods
Ca e desc ip ions
Danial Ca e (IRAN)
Danial Ca e is he second la ges i e ca e in I an, which is loca ed in he Albo z
Moun ain ange in he Mazanda an ka s a ea, on he sou he n edge o he Caspian Sea
nea he Salman Shah o Abbasabad ci y (36°39'35.2"N, 51°10'53.2"E) a al i ude
o 204 m abo e sea le el (Fig. 1), i s name de i ed om he close illage. Al e na i e
name o his ca e is Ge de Kooh. The ca e is si ua ed in he Hy canian Fo es ype on
he slope o he moun ain wi h he main en ance 5 m long and 6 m high (Fig. 2). I s
leng h is 2,158 m, wi h a sub e anean i e lowing along he en i e ca e (Yuse inia
2017; Ghelich Khani e al. 2022; Ba jadze e al. 2024; Meh a ooz May an e al.
2025). I has cons an ai empe a u e (a ound 12 °C) and ela i e humidi y (RH) is
o e 73%, measu ed wi h a he mo-hyg ome e TFA 30.5015 DTH in Ap il 2024.
Baba uša Ca e (MONTENEGRO)
This ca e is si ua ed nea he illage o T no o, Vi paza , (42°17'56"N, 19°01'58"E) a al i-
ude o 381 m abo e sea le el (Fig. 7). The en ance is a ho izon al issu e abou 10 m wide
and abou 2 m high (Fig. 8) (Vujačić e al. 2024). The ca e is ela i ely sho (a li le o e 290
m), wi h he g ea main chambe ich in lows ones and small sin e pools (Dju o ic and
Dju o ic 2021). I s galle ies a e illed wi h clay deposi s up o 10 m hick (Vujačić e al. 2024).
Me hods
Fo obse a ion in an op ical mic oscope wi h phase con as , specimens we e i s ly
boiled so ly in e apo a ion glass dish wi h 95% e hyl alcohol on elec ic cooking
pla e o app oxima ely 1 minu e o emo e a om he body. Fo clea ing, he
specimens we e anspo ed o conca e glass dish wi h 10% wa e solu ion o KOH
o ca. 1 minu e and hen anspo ed o a dish con aining Chlo ophenol un il he
specimens we e ully anspa en . A he end, specimens we e moun ed on Swann
medium on pe manen slides. A e slide was su icien ly d ied ou , he co e glass
was inged wi h Canada balsam o seal he moun ing medium and p e en i s desic-
ca ion. 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 a m.
Ľubomí Ko áč e al. / Sub e anean Biology 54: 1–22 (2025)
4
Figu e 1. A map o I an, ed a ea - Mazanda an p o ince; loca ion o he Danial Ca e, locus ypicus o
Schae e ia g eensladeae sp. no ., is ma ked wi h a ed as e isk on an enla ged map o he p o ince (uppe )
B Hy canian Fo es nea he ca e in summe (Pho o: M. Meh a ooz).
Abb e ia ions used. a—an e io ow o se a(e); An —an ennal segmen (s);
Asp—anal spines; Cl—claw; m—median ow o se a(e); Mc—mac ose a(e); ms—mi-
c osensillum; os—subapical o gani e (sub-apical senso y peg); p—pos e io ow o
se a(e); PAO—pos an ennal o gan; s—senso y se a; Ti— ibio a sus; VT— en al
ube; CoPJSU—Collembola collec ion o he Depa men o Zoology, Ins i u e o
Biology and Ecology, Facul y o Science, Pa ol Joze Ša á ik Uni e si y, Košice, Slo a-
kia; ZMFUM—Zoology Museum o Fe dowsi Uni e si y o Mashhad, Mashhad, I an.
Resul s
Species desc ip ion
Schae e ia g eensladeae Meh a ooz May an & Ko áč, sp. no .
h ps://zoobank.o g/424A45B7-3B5F-41C2-992B-788B3B16DC29
Figs 3–6, Table 1
Diagnosis. Eyes absen , PAO la ge wi h 7 subequal lobes o med a ound cen al
ci cle. An IV wi h 7–8 sensilla and simple apical esicle. Claw enla ged, wi h an
in e nal oo h and pai o la e al ee h. Re inaculum wi h 3–4 ee h, u ca well-
de eloped, wi h 5 den al se ae.
Two new species o eyeless ca e nicolous Schae e ia 5
Type locali y. I an, Mazanda an p o ince, Salman Shah Coun y, Abbasabad ci y,
Danial illage, Danial Ca e, Coo dina es: 36°39'35.2"N, 51°10'53.2"E, ca e en ance
204 m a.s.l.
Type ma e ial. Holo ype: • Female on slide [DaSc101IR], da k zone, collec -
ed om su ace o wa e puddle, 15.x.2023, empe a u e = 11.9 °C, RH 73.0%,
leg. M. Meh a ooz. Pa a ypes: • 2 specimens on slides [DaSc102IR and DaS-
c103IR], da k zone, collec ed om su ace o wa e puddle, 15.x.2023, empe a-
u e = 12.0 °C, RH 73.4%, leg. M. Meh a ooz. Holo ype deposi ed in CoPJSU;
wo pa a ypes deposi ed in ZMFUM.
Desc ip ion. Body appea ance. Leng h 1.9 mm on a e age (n = 3, max. leng h
2 mm). Habi us ypical o Ce a ophysella Bö ne in B ohme , 1932 wi hou eyes.
Colou ali e whi e wi hou aces o pigmen . Cu icula g anula ion a he uni o m,
sligh ly coa se on Abd V–VI, mac o- and mesose ae di e en ia ed.
An enna. I is nea ly as long as head, wi h apical bulb (Fig. 3A–D). Leng h o
An I, II, III, IV in holo ype as 0.07, 0.06, 0.07, 0.1 mm, espec i ely. An I wi h
7 se ae; An II wi h 6 do sal and 6 en al se ae in a single ow and 1 sho smoo h
se a nea i s base (do sal se ae clea ly coa se and hicke ); An III and IV clea ly
Figu e 2. A g ound plan o he Danial Ca e (adop ed om Ghelich Khani e al. 2024) B ca e deco a-
ions in Rizan chambe (Pho o: M. Meh a ooz) C main en ance o he ca e (Pho o: M. Meh a ooz).
Ľubomí Ko áč e al. / Sub e anean Biology 54: 1–22 (2025)
6
sepa a ed, An III senso y o gan wi h 2 cen al sho and cu ed ods lanked by
2 long gua d sensilla (Fig. 3B), a small en ola e al mic osensillum in a en al
pi , and 7 do sal and 8 en al o dina y se ae; An IV wi h a simple apical bulb
(Fig. 3C), 7–8 cylind ical blun sensilla (a, b, c, d, e, , g and h) (a e Yosii 1962),
subapical ms nea base o sensillum a, apical o gani e os placed nea sensillum b
and 20 se ae a ound sensilla.
Head. Eyes absen . PAO nea base o An I well de eloped, la ge, abou 33 µm
in diame e , wi h 7 subequal lobes o med a ound cen al ci cle (Fig. 3F). Lab um
ypical o he genus wi h 2+2 p elab al and 5, 5, 4 lab al se ae. Labium wi h all
no mal papillae (A–E) and 6 p oximal se ae. Le mandible wi h 4 ee h and igh
mandible wi h 5 ee h.
Abdomen. Ven al ube wi h 4+4 apical se ae. Re inaculum wi h 3–4 ee h
(asymme y obse ed). Fu ca well-de eloped (Fig. 5B), leng h o manub ium, dens,
Figu e 3. Schae e ia g eensladeae sp. no . A an ennal segmen s I–IV B An III senso y o gan C ip o
an enna D An IV blun sensillum E do sal side o head, se al pa e n F pos an ennal o gan wi h he nea -
es se ae and g anula ion o cu icle G shape o mac ose a. Scale ba s: 0.025 mm (A); 0.01 mm (B, C);
0.025 mm (E); 0.02 mm (F–G).
Two new species o eyeless ca e nicolous Schae e ia 7
Figu e 4. Schae e ia g eensladeae sp. no . A–C se al pa e n o legs A leg I B leg II C leg III D–F oo
complex o legs D leg I E leg II F leg III. Scale ba s: 0.025 mm (A–C); 0.01 mm (D–F).
muc o in holo ype as 0.11, 0.06, 0.01 mm, espec i ely; manub ium wi h 12 se ae;
dens wi h 5 se ae (basal se a longe ); muc o abou 1/4 o dens leng h, wi h la e al
lamella, muc o and dens aken oge he a e as long as inne edge o claw III. Anal
spines long (86 µm), on papillae (Fig. 6C). Ra io anal spines (including papilla):
claw III = 1.6; a io anal spines: dens = 1.9.
Legs. Legs I–III wi h no mal se ae, enen hai acumina e. Se al pa e n as in
Fig. 4A–C. Leg I–III wi h 3, 8, 8 se ae on coxae, 7, 7, 7 se ae on ochan e s, 14, 13,
12 se ae on emo a and 19, 19, 18 se ae on ibio a si, espec i ely. Claw slende , 78 µm
long in holo ype, in e nal edge wi h s ong medial oo h and la e al edge wi h pai o
Ľubomí Ko áč e al. / Sub e anean Biology 54: 1–22 (2025)
8
Figu e 5. Schae e ia g eensladeae sp. no . A do sal side o ho ax, se al pa e n B u ca, se al pa e n.
Scale ba s: 0.05 mm (A); 0.01 mm (B).
Figu e 6. Schae e ia g eensladeae sp. no . A Do sal side o Abd I–III, se al pa e n B do sal side o Abd
IV–VI, se al pa e n C de ail o anal spine, la e al iew. Scale ba s: 0.05 mm (A, B); 0.01 mm (C).
Two new species o eyeless ca e nicolous Schae e ia 9
basal ee h; p e a sus wi h 1 lobe and 2 se ae (1 ex e nal and 1 in e nal) eaching base
o la e al ee h (Fig. 4D–F). Ra io o claw I leng h: wid h = 3.2.
Se al pa e n o e gi es. Do sal se al pa e n gene ally o ype II wi h well-
di e en ia ed se ae. Se ae on head and body sligh ly den a ed (Fig. 3G). Head.
Se al pa e n as in Fig. 3E. Se a a0 p esen ; d1–d5 (d2 and d4 as Mc); sd1–sd5;
1– 2 ( 2 as Mc); c1–c5; p1–p3, p5 (p3 and p5 as Mc, p4 absen ); oc1–oc3 (oc1
as Mc). Tho ax. Se al pa e n as in Fig. 5A. Th I wi h 3+3 se ae as m1, m3 and m4.
Th II–III wi h a1–a6 (a3 as Mc); m1, m4’–m6 (m6 as s), la e al mic osensillum
p esen on Th II in on o m6 (s); p1–p4 (p4 as s), p5–p6 (p2, p5 and p6 as Mc).
Abdomen. Se al pa e n as in Fig. 6A, B. Abd I–III wi h 6+6 se ae be ween p5 (s)
in ows ‘a’ and ‘m’: a1, a2, a4, m2, m3 and m4 (p2, p4, p6 and p7 as Mc). Abd IV
wi h 7+7 se ae be ween p5 (s) in ows ‘a’ and ‘m’: a1, a2, a4, a5, m1, m3 and m5
(a4, a5, p2, p4 and p6 as Mc). Abd V wi h 2+2 se ae be ween p3 (s) on ow ‘a’: a1
and a3 (a5, p1 and p5 as Mc).
E ymology. The species is named in he honou o p o . D . Penelope G eenslade,
an ou s anding Collembola axonomis and ecologis .
Taxonomic ema ks. Based on absence o eyes, S. g eensladeae sp. no . is simila
o i e ep esen a i es o he genus: S. coeca Cassagnau, 1959, S. gue e ense (Bone ,
1945), S. oaxacana Palacios-Va gas & Thibaud, 1985, S. p o undissima Jo dana &
Baque o, 2012 and S. subcoeca Deha eng & Thibaud, 1980. The new species is
cha ac e ized by 7 lobes in PAO, while he e a e 4 (6) lobes in PAO in S. coeca and
S. subcoeca, 4–5 lobes in S. oaxacana and 4 lobes in S. gue e ense and S. p o undis-
sima. Mo eo e , he new species is sepa a ed om S. coeca in numbe o sensilla
on An IV: 8–12 in S. coeca and 7–8 in S. g eensladeae. The bo h species di e also
in numbe o den al se ae: 3–5 se ae in S. coeca and 5 se ae in S. g eensladeae. Mu-
c o is absen in S. gue e ense and S. subcoeca, bu i is p esen in he new species.
Schae e ia g eensladeae sp. no . has 5 den al se ae, while S. oaxacana has 4 den al
se ae. Also, S. g eensladeae sp. no . in e ms o he se al pa e n o he do sal body
is close o S. oaxacana and S. p o undissima (see Table 1). Schae e ia g eensladeae
sp. no . di e s om S. p o undissima by he p esence o se a a1 on Abd IV, and
om S. oaxacana in ha ing se a m4 on Abd IV, whe eas S. oaxacana has m3. The
new species also ep esen s he i s eco d o his genus in he coun y (Meh a ooz
May an e al. 2023).
Schae e ia mon eneg ina Ko áč & Meh a ooz May an, sp. no .
h ps://zoobank.o g/13F15544-55EE-4D2A-AF61-80B6939D2D10
Figs 9–12, Table 1
Diagnosis. Eyes absen , PAO small, o ming a ose e o 5–6 egula o al esicles. An
IV wi h 7–8 sensilla and simple apical esicle. Th I wi h 2+2 se ae, Abd V wi hou se a
p2. Claw wi hou in e nal o la e al ee h. Re inaculum wi h 3–4 ee h, u ca well-
de eloped, wi h 5 den al se ae.
Ľubomí Ko áč e al. / Sub e anean Biology 54: 1–22 (2025)
16
Rema ks o phylogeny o he genus
Thibaud (1972) ou lined Schae e ia as a alid genus, ep esen a i es o which show mo -
phological eg essions in pigmen a ion, numbe o eyes and u cal complex. He ega ded
hese ea u es as mani es a ions o adap a ion o he eudaphic o ca e li es yle, ha ing
e olu iona y signi icance, and he genus as a phyle ic line o he species g oup Ce a o-
physella den icula a, o med as a esul o pos glacial coloniza ion o he unde g ound
en i onmen (deep soil, ca es). The genus Schae e ia, simila ly as gene a Typhlogas u a
Bone , 1930 and Bone ogas u a Thibaud, 1974, is appa en ly de i ed om Ce a ophysella
g oups o species, which ha e success ully in aded ca es and became mo e o less oglo-
mo phic (Ch is iansen and Wang 2006). In he con a y o Schae e ia, he bo h gen-
e a a e cha ac e ized by he main enance o a ully de eloped u ca. A e Ch is iansen
(1985), educ ion o u ca in Schae e ia may ep esen an edaphic phenomenon. I is
impo an o no e ha all h ee gene a ha e common Hola c ic dis ibu ion.
Howe e , any assump ions o phyle ic ela ionships wi hin he genus Schae e ia,
o p esumably ela ed gene a, emain o be cla i ied by a ho ough phylogene ic s udy
combining mo phological ai s wi h mi ochond ial and nuclea DNA da a. I is in-
deed no an easy ask when conside ing demanding ield collec ions o hese collem-
bolans wi h usually a e and dispe sed dis ibu ion in sub e anean spaces.
Rema ks on mo phological adap a ions o sub e anean en i onmen
In he p e ious sec ion, we b ie ly discussed oglomo phic and edaphomo phic adap a-
ions in Schae e ia ep esen a i es. Thus, blidness i sel doesn’ necessa ily imply asso-
cia ion wi h ca e li e, well documen ed in his genus by h ee edaphic species, all om
F ance: S. a iegica Cassagnau, 1959, S. coeca and S. subcoeca. Rega ding oglobio ic and
guanobio ic o ms, al oge he six species ha e been epo ed in his genus exclusi ely
om ca es (Lukić 2019): S. gue e ense and S. oaxacana om Mexico, S. p o undissima
om he Caucasus, S. ki akamiana Yosii, 1991 om Japan, S. baschki ica Kniss, 1985
om he U als, Russia, and S. hubba di Thibaud, 1995 om USA. Schae e ia mon e-
neg ina sp. no . has clea ly elonga ed claw and missing in e nal ee h, he combina ion
o bo h cha ac e s conside ed oglomo phic adap a ions (Ch is iansen 2012). I has
been no iced ha some guanobio ic species o he amily Hypogas u idae exhibi his
cha ac e (Deha eng and Bedos 2018) as an adap a ion o easie mo emen on esh
ba guano (Lukić 2019), such as in Ache on ides eleono ae Palacios-Va gas and Gnaspini-
Ne o, 1992 om B azil. Ne e heless, he e a e h ee blind species o Schae e ia wi h
elonga ed unguis: wo species desc ibed in his pape and S. p o undissima om K ube a-
Vo onya, a deep ca e sys em in he Caucasus Moun ains. Elonga ion o unguis (claw) in
hese species is an ob ious oglomo phic cha ac e ha e ol ed independen ly in h ee
geog aphically dis an ka s complexes (Dina ic Ka s , Albo z Moun ains and Caucasus
Moun ains) as an example o con e gen e olu ion, a well-known phenomenon in se -
e al in e eb a e g oups inhabi ing deep sub e anean ealm.
Two new species o eyeless ca e nicolous Schae e ia 17
Key o wo ld species o genus Schae e ia
The ollowing key was p epa ed based on he diagnos ic cha ac e s o wo ldwide
Schae e ia species (adap ed om Jo dana e al. 2020).
1 Muc o absen ..............................................................................................2
– Muc o educed o well-de eloped ...............................................................9
2 Wi h 7+7 eyes; dens wi h 6 se ae; Abd IV wi hou se a a1 ............................
.......................................................S. oculea Babenko, 1999 (Russia, soil)
– Wi h less han 7+7 eyes; dens wi h less han 6 se ae; Abd IV wi h se a a1 ...3
3 Wi h 5+5 eyes ....S. canigouensis Deha eng & Thibaud, 1980 (F ance, soil)
– Wi h less han 5+5 eyes...............................................................................4
4 Wi h 3+3 eyes .............................................................................................5
– Wi h less han 3+3 eyes...............................................................................6
5 Dens wi h 4 se ae; Abd II–III wi h 6+6 se ae in a- and m- ow oge he (m3
p esen ) ................... S. emuc ona a Absolon, 1900 (Czech Republic, ca e)
– Dens wi h 3 se ae; Abd II–III wi h 5+5 se ae in a- and m- ow oge he (m3
absen ) ........................................S. lindbe gi Gama, 1963 (Po ugal, ca e)
6 Wi h 2+2 eyes .............................................................................................7
– Wi h 0+0 eyes .............................................................................................8
7 Dens wi h 4 se ae; An IV wi h 8–9 sensilla; Abd II–III wi hou se ae in m-
ow; Abd IV wi hou se a a4 .........................................................................
........................................S. quad iocula a (S ach, 1939) (Ge many, ca e)
– Dens wi h 3 se ae; An IV wi h 7 sensilla; Abd II–III wi h 2 se ae in m- ow
(m2 and m4); Abd IV wi h se a a4................................................................
.................................................S. a iegica Cassagnau, 1959 (F ance, soil)
8 Dens wi h 4–5 se ae; Abd IV wi h se a a4; Abd IV wi h se ae a1, a2, a4, a5 .
.............................................. S. gue e ense (Bone , 1945) (Mexico, ca e)
– Dens wi h 3 se ae; Abd IV wi hou se a a4; Abd IV wi h se ae a1, a3, a5 ......
............................ S. subcoeca Deha eng & Thibaud, 1980 (F ance, soil)
9 Muc o educed .........................................................................................10
– Muc o well-de eloped ...............................................................................15
10 Wi h 5+5 eyes ...........................................................................................11
– Wi h less han 5+5 eyes.............................................................................12
11 Dens wi h 5 se ae; Abd II–III wi h 2+2 se ae in m- ow (m2, m4) .................
............................. S. maxima Deha eng & Thibaud, 1980 (F ance, soil)
– Dens wi h 3–4 se ae; Abd II–III wi h 3+3 se ae in m- ow (m2, m3, m4) ......
.........................................S. decemocula a (S ach, 1939) (Ge many, ca e)
12 Wi h 4+4 eyes .............................S. willemi (Bone , 1930) (Belgium, ca e)
– Wi h less han 4+4 eyes.............................................................................13
13 Wi h 3+3 eyes ...........................................................................................14
– Wi h 0–1 eye .................................S. coeca Cassagnau, 1959 (F ance, soil)
Ľubomí Ko áč e al. / Sub e anean Biology 54: 1–22 (2025)
18
14 Dens wi h 3 se ae; PAO wi h 4 esicles; Abd II–III wi hou se ae in m- ow;
Abd V wi hou se a a2 ..................................................................................
..................... S. pouadensis Delama e Debou e ille, 1945 (F ance, ca e)
– Dens wi h 6 se ae (some ime 5); PAO wi h 6–8 esicles; Abd II–III wi h 2+2
se ae in m- ow (m2, m4); Abd V wi h se a a2 ...............................................
.......................................S. scossi olii Dallai & Saba ini, 1981 (I aly, soil)
15 Wi h 6+6 eyes ...........................................................................................16
– Wi h less han 6+6 eyes.............................................................................24
16 An IV wi h 7 sensilla ...............................................................................17
– An IV wi h 6 sensilla ...............................................................................23
17 Dens wi h 3–4 se ae ............................... S. cheoha W ay, 1963 (USA, ca e)
– Dens wi h mo e han 4 se ae .....................................................................18
18 Abd V wi h se a a2 ........ S. jellbe gi Jo dana & Baque o, 2020 (USA, soil)
– Abd V wi hou se a a2 ..............................................................................19
19 Abd IV wi h 5+5 se ae in a- ow (a1–a5) .......................................................
.................................. S. andalica (Jo dana & A bea, 1990) (Spain, soil)
– Abd IV wi h less han 5+5 se ae in a- ow ..................................................20
20 Abd IV wi h 3+3 se ae in m- ow ...............................................................21
– Abd IV wi h 2+2 se ae in m- ow ...............................................................22
21 Abd II–III wi h se ae m2, m4, m5; dens wi h 5 se ae; a io Asp: dens as 4.5; a io
dens: muc o as 4 ........... S. babenkoi Jo dana & Baque o, 2020 (Russia, soil)
– Abd II–III wi h se ae m2, m3, m4; dens wi h 5–6 se ae; a io Asp: dens as
2.3; a io dens: muc o as 2.7 .....................................................................
..................................... S. duodecimocella a Bone , 1945 (Mexico, soil)
22 Dens wi h 5 se ae; Abd IV wi h se ae a1, a3, a5 in a- ow; Abd IV wi h se ae
m1, m4 in m- ow; a io Asp: dens as 1; a io dens: muc o as 2.3 ..................
................................................S. elegans (Cassagnau, 1959) (F ance, soil)
– Dens wi h 6 se ae; Abd IV wi h se ae a1, a2, a4, a5 in a- ow; Abd IV wi h
se ae m1, m5 in m- ow; a io Asp: dens as 2.2; a io dens: muc o as 2.5 .......
..................................... S. send ai Jo dana & Baque o, 2020 (Spain, soil)
23 Dens wi h 4 se ae; enaculum wi h 3+3 ee h; Abd IV wi h 2+2 se ae in m-
ow (m1, m3) ...........S. deha e gi (Jo dana & A bea, 1990) (Spain, soil)
– Dens wi h 5 se ae; enaculum wi h 4+4 ee h; Abd IV wi h 1+1 se a in m- ow
(m1) ..................................S. ja ae (Jo dana & A bea, 1990) (Spain, soil)
24 Wi h 5+5 eyes ...........................................................................................25
– Wi h less han 5+5 eyes.............................................................................29
25 An IV wi h 5 sensilla; dens wi h 3 se ae .......................................................
............................................S. biden a a (Cassagnau, 1954) (F ance, soil)
– An IV wi h mo e han 5 sensilla; dens wi h mo e han 3 se ae .................26
Two new species o eyeless ca e nicolous Schae e ia 19
26 PAO wi h 5 esicles; Abd V wi hou se a p1 .................................................
S. aulmugnizi Palacios-Va gas & Cas año-Meneses, 2008 (Mexico, soil)
– PAO wi h 4 esicles; Abd V wi h se ae p1 .................................................27
27 Dens wi h 4–5 se ae; Abd IV wi hou se a a1 ................................................
...................................................S. be mani Babenko, 2019 (Russia, soil)
– Dens wi h 5–6 se ae; Abd IV wi h se a a1 .................................................28
28 Labial papilla C absen ; Abd V wi hou se a a2’ ............................................
............................................. S. cze no i (Ma yno a, 1978) (Russia, soil)
– Labial papilla C p esen ; Abd V wi h se a a2’ ................................................
........................................... S. quinqueocula a (Yosii, 1956) (Japan, ca e)
29 Wi h 4+4 eyes ...........................................................................................30
– Wi h less han 4+4 eyes.............................................................................31
30 PAO wi h 4 esicles; An IV wi h 6–7 sensilla; Abd IV wi h 4+4 se ae in m-
ow (m1, m3, m4, m5) .......S. ukugakuchiana (Yosii, 1956) (Japan, ca e)
– PAO wi h 4–6 (7–8) esicles; An IV wi h 7–8 sensilla; Abd IV wi h 1+1 se a
in m- ow (m4) ............................S. baschki ica Kniss, 1985 (Russia, ca e)
31 Wi h 3+3 eyes ...........................................................................................32
– Wi h less han 3+3 eyes.............................................................................33
32 An IV wi h 6–7 sensilla; dens wi h 5 se ae; Abd II–III wi h 3+3 se ae in m-
ow (m2, m3, m5) ........................S. hubba di Thibaud, 1995 (USA, ca e)
– An IV wi h 9 (7–8) sensilla; dens wi h 4 se ae; Abd II–III wi hou se ae in
m- ow .................................S. sexocula a (Gisin, 1947) (Swi ze land, soil)
33 Wi h 2+2 eyes; An IV wi h 6 sensilla ...........................................................
...................................................S. ki akamiana Yosii, 1991 (Japan, ca e)
– Wi h less han 2+2 eyes; An IV wi h mo e han 6 sensilla ........................34
34 An IV wi h ilobed apical esicle; dens wi h 4 se ae; Ti I–III wi h 18, 18, 17
se ae, espec i ely ..........................................................................................
................. S. oaxacana Palacios-Va gas & Thibaud, 1985 (Mexico, ca e)
– An IV wi h simple apical esicle; dens wi h 5 se ae; Ti I–III wi h 19, 19, 18
se ae, espec i ely ......................................................................................35
35 PAO wi h 4 esicles; do sal head wi hou se a a0; Abd IV wi hou se a a1 in
a- ow ............S. p o undissima Jo dana & Baque o, 2012 (Geo gia, ca e)
– PAO wi h mo e han 4 esicles; do sal head wi h se a a0; Abd IV wi h se a a1
in a- ow ....................................................................................................36
36 PAO wi h 7 esicles; Th I wi h 3+3 se ae (m1, m3, m4); Abd V wi h se a p2;
Cl wi h a s ong inne oo h and pai o la e al ee h.....................................
............................................................S. g eensladeae sp. no . (I an, ca e)
– PAO wi h 5–6 esicles; Th I wi h 2+2 se ae (m1, m4); Abd V wi hou se a p2; Cl
wi hou inne o la e al ee h .....S. mon eneg ina sp. no . (Mon eneg o, ca e)
Ľubomí Ko áč e al. / Sub e anean Biology 54: 1–22 (2025)
20
Acknowledgemen s
The s udy was suppo ed om he Slo ak Resea ch and De elopmen Agency, p o-
jec APVV-21-0379, and om he Slo ak Scien i ic G an Agency, p ojec VEGA
1/0438/22. The hi d 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.
We would like o hank Mohammad Ali Ja i (head o he climbing boa d o Abbas
Abad coun y, I an), D . Saeid Esmaeilpou and Sahand Sa a i Nou i o assis ance
du ing he expedi ions o Danial Ca e. Also, we a e g a e ul o Ka oshga an Ca ing
Club in Mashhad o o help inc ease knowledge and p o ec I anian ca es.
Re e ences
Absolon K (1900) Vo läu ige Mi heilung übe einige neue Collembolen aus den Höhlen. Zo-
ologische Anzeige 23: 265–269.
Babenko AB (1999) New sp ing ail species o he amily Hypogas u idae om Eas e n Sibe ia.
Zoologichesky Zhu nal 78: 885–888.
Babenko AB (2019) A new species o Schae e ia Absolon (Collembola: Hypogas u idae) om
Chuko ka, Russia, wi h ema ks on he ela ed o ms. Zoo axa 4571(2): 247–254. h ps://
doi.o g/10.11646/zoo axa.4571.2.4
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
Bone F (1930) Rema ques su les Hypogas u iens ca e nicoles a ec desc ip ions d’espèces
nou elles (Collembola). EOS, Re is a Española de En omología 6(2): 113–139.
Bone F (1945) Nue os Gene os y Especies de Hipogas u idos de Mexico (Collembola). Re-
is a de la Sociedad Mexicana de His o ia Na u al 6(2): 13–45.
Bö ne C (1932) Ap e ygo a. In: B ohme P (Ed.) Fauna Von Deu schland. 4 h edn. Quelle &
Meye , Leipzig, 136–143.
Cassagnau P (1954) Faune ançaise des Collemboles. III. Podu omo phes de hau e mon agne.
Vie e Milieu 4(1): 65–74.
Cassagnau P (1959) Faune ançaise des Collemboles IX. Les Hypogas u a a. l. du Massi du
Néou ielle (Hau es-Py énées). Rema ques su che o axie des espèces. Vie e Milieu 9(4):
476–503.
Ch is iansen KA (1985) Reg essi e e olu ion in Collembola. Na ional Speleological Socie y
Bulle in 47(2): 89–100.
Ch is iansen KA (2012) Mo phological adap a ions. Pp. 517–528. In: Whi e WB, Cul e DC
(Eds) Encyclopedia o ca es. Else ie /Academic P ess, 945 pp. h ps://doi.o g/10.1016/
B978-0-12-383832-2.00075-X
Ch is iansen KA, Wang H (2006) A e ision o he genus Typhlogas u a in No h Ame ican
ca es wi h desc ip ion o i e new species. Jou nal o Ca e and Ka s S udies 68(2): 85–98.
Dallai R, Saba ini MA (1981) Rice che Sui Collemboli XXVIII. Su di una nuo a specie endo-
gea di Schae e ia. Redia 64: 195–204.
Two new species o eyeless ca e nicolous Schae e ia 21
Deha eng L, Bedos A (2018) Chap e 7. Di e si y o e es ial in e eb a es in sub e anean
habi a s. In: Moldo an OT, Ko áč Ľ, Halse S (Eds) Ca e Ecology. Ecological S udies Vol.
235. Sp inge , Cham, 107–172. h ps://doi.o g/10.1007/978-3-319-98852-8_7
Deha eng L, Thibaud JM (1980) T ois espèces nou elles du gen e Schae e ia Absolon, 1900
(Insec e, Collembole). Nou elle Re ue d’En omologie 10: 3–10.
Delama e-Debou e ille C (1945) Su quelques Collemboles de la égion de Banyuls (Py énées-
O ien ales) a ec la desc ip ion d’une espèce oglobie. Bulle in de la socié é en omologique
de F ance 6: 70–72. h ps://doi.o g/10.3406/bse .1945.15827
Dju o ic M, Dju o ic P (2021) Re iew o he mos signi ican ca es in Mon eneg o. Ac a Ca -
sologica 50(1): 49–65. h ps://doi.o g/10.3986/ac. i.7182
Fjellbe g A (1985) A c ic Collembola. I – Alaskan Collembola o he amilies Podu idae, Hy-
pogas u idae, Odon ellidae, B achys omellidae and Neanu idae. En omologica Scandi-
na ica 21: 1–126.
Gama MM da (1963) Qua e espèces nou elles de Collemboles d’Au iche e de Jougosla ie.
A chi es des Sciences (Genè e) 16: 43–50.
Ghelich Khani P, Qashqaei AT, Kolke H (2024) S a us o e eb a e species in Danial Ca e,
no he n I an. O yx 58(1): 104–109. h ps://doi.o g/10.1017/S003060532200165X
Gisin H (1947) Su les insec es ap é ygo es du Pa c Na ional Suisse. Espèces e g oupemen s
euédaphiques. Résul a s des echee ches scien i iques en e p ises au Pa c Na a ional Suisse
2: 77–91.
Jo dana R, A bea JI (1990) Especies ibé icas de Hypogas u a (Ce a ophysella) de seis ojos con
desc ipción de es nue as especies (Collembola, Hypogas u idae). G aellsia 46: 161–173.
Jo dana R, A bea JI, Simon JC, Lucianez MJ (1997) Collembola Podu omo pha. In: Ramos
MA, e al. (Eds) En: Fauna Ibe ica, 8. Museo Nacional de Ciencias Na u ales (CSIC),
Mad id, 807–807.
Jo dana R, Baque o E (2012) Re iews o he gene a Schae e ia Absolon, 1900, Deu e apho u a
Absolon, 1901, Plu omu us Yosii, 1956 and he Anu ida Laboulbène, 1865 species g oup
wi hou eyes, wi h he desc ip ion o ou new species o ca e sp ing ails (Collembola)
om K ube a-Vo onya ca e, A abika Massi , Abkhazia. Te es ial A h opod Re iews 5:
35–85. h ps://doi.o g/10.1163/187498312X622430
Jo dana R, Baque o E, Ledesma E, Send a A, O uño VM (2020) Podu omo pha (Collembola)
om a sampling in he meso oid shallow subs a um o he Sie a de Guada ama Na ional
Pa k (Mad id and Sego ia, Spain): Taxonomy and Biogeog aphy. Zoologische Anzeige
285: 81–96. h ps://doi.o g/10.1016/j.jcz.2020.02.001
Kniss VA (1985) Sp ing ails o he genus Schae e ia (Collembola, Hypogas u idae) om ca es
in he sou hwes e n pa o he Bashki ASSR. Zoologicheskii zhu nal 64(6): 934–937.
Lukić M (2019) Chap e 35 - Collembola. 308–319. In: Whi e WB, Cul e D, Pipan T (Eds)
Encyclopedia o Ca es, 3 d edn., Academic P ess, 1250 pp. h ps://doi.o g/10.1016/B978-
0-12-814124-3.00034-0
Ma yno a EF (1978) New species o Sp ing ails o he genus Hypogas u a s. l. in he No h-
Eas e n Asia Fauna. No ye I Maloiz es nye Vidy Fauny Sibi i 12: 27–47.
Meh a ooz May an M, G eenslade P, Sadeghi-Namaghi H (2023) An anno a ed check-
lis 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
Ľubomí Ko áč e al. / Sub e anean Biology 54: 1–22 (2025)
22
Meh a ooz May an M, Ko áč Ľ (2025) Two new species o he genus Ache on iella (Col-
lembola, 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
Palacios-Va gas JG, Cas año-Meneses G (2008) A new species o Schae e ia Absolon, 1900
(Collembola: Hypogas u idae). Dugesiana 15(2): 123–125.
Palacios-Va gas JG, Gnaspini-Ne o P (1992) A new B azilian species o Ache on ides (Col-
lembola: Hypogas u idae) wi h no es on i s ecology. Jou nal o The Kansas En omological
Socie y 65: 443–447.
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.
S ach J (1939) Die Höhlen auna des Gla ze Schneebe ges. II Die Collembolen auna de Sal-
zlöche bei Sei endo . Bei äge zu Biologie des Gla ze Schneebe ges 5: 395–415.
S eine W (1955) Bei äge zu Kenn nis de Collembolen auna Spaniens. Eos 34: 323–340.
Thibaud JM (1972) Re ision du gen e Schae e ia Absolon, 1900 (Insec e, Collembole). An-
nales de Spéléologie 27(2): 363–394.
Thibaud JM (1974) Subdi ision de Typhlogas u a, gen e oglobie de la amille des Hypogas-
u idae (Insec es Collemboles). Spelunca 8: 205–207.
Thibaud JM (1995) Une nou elle espèce de Schae e ia d’une g o e des E a s-Unis d’Amé ique
(Collembola, Hypogas u idae). Re ue F ançaise d’En omologie (Nou elle Sé ie) 17(2):
77–78.
Thibaud JM, Schulz HJ, Gama Assalino MM da (2004) Hypogas u idae. In: Dunge W (Ed.)
Synopses on Palaea c ic Collembola. Vol. 4, Abhandlungen und Be ich e des Na u kunde-
museums Gö li z 75(2): 1–287.
Vujačić D, Pe o ić Đ, G ozdanić G, Vujo ić F (2024) P o ec ed Speleological Objec s. In:
Ba o ic G (Eds) Speleology o Mon eneg o. Ca e and Ka s Sys ems o he Wo ld. Sp ing-
e , Cham. h ps://doi.o g/10.1007/978-3-031-49375-1_8
W ay DL (1963) Collembola o No h Ame ica; Pa I, Hypogas u inae. Jou nal o he Ten-
nessee Academy o Science 38(3): 101–104.
Yosii R (1991) Abou he p ose pinae g oup o Hypogas u a (Collembola) in he Ca es o P e .
Iwa e. Annals o he Speleological Resea ch Ins i u e o Japan (Iwaizumi) 9: 3–10.
Yosii R (1956) Monog aphie zu Höhlencollembolen japans. Con ibu ions om he Biologi-
cal Labo a o y Kyo o Uni e si y 3: 1–109.
Yosii R (1962) S udies on he collembolan genus Hypogas u a II. Nea c ic o ms collec ed by
P o . F. Bone . Con ibu ions om he Biological Labo a o y, Kyo o Uni e si y 13: 1–25.
Yuse inia AA (2017) Ancien Langa: Kela abad, Salmanshah and Abas Abad. Bolu Publishe ,
Rash , 61 pp. [In Pe sian]
