Te es ial isopods (C us acea, Isopoda,
Oniscidea) om Meso oid Shallow
Subs a um habi a s o Po ugal
Ri a P. Eusébio1, S e ano Tai i2, Ana So ia P. S. Rebolei a1,3
1Cen e o Ecology, E olu ion and En i onmen al Changes (CE3C) and CHANGE – Global Change and
Sus ainabili y Ins i u e, and Depa amen o de Biologia, Faculdade de Ciências, Uni e sidade de Lisboa, Campo
G ande, 1749–016, Lisboa, Po ugal 2Is i u o di Rice ca sugli Ecosis emi Te es i, Consiglio Nazionale delle
Rice che, Via Madonna del Piano 10, 50019 Ses o Fio en ino (Fi enze), I aly 3Museu Nacional de His ó ia
Na u al e da Ciência, Uni e sidade de Lisboa, Rua da Escola Poli écnica, Lisboa, Po ugal
Co esponding au ho : Ana So ia P. S. Rebolei ([email p o ec ed])
Academic edi o : Fabio S och|Recei ed 30 June 2025|Accep ed 25 Augus 2025|Published 8 Oc obe 2025
h ps://zoobank.o g/98FAD49E-ED52-4C9E-82B5-120452E46F19
Ci a ion: Eusébio RP, Tai i S, Rebolei a ASPS (2025) Te es ial isopods (C us acea, Isopoda, Oniscidea) om Meso oid
Shallow Subs a um habi a s o Po ugal. Sub e anean Biology 53: 129–153. h ps://doi.o g/10.3897/sub biol.53.163632
Abs ac
The Meso oid Shallow Subs a um (MSS) is a la gely unknown sub e anean habi a o med by he
accumula ion o ock agmen s in s eep slopes. In his s udy, we in es iga ed he di e si y, abundance
and seasonal dynamics o e es ial isopods in MSS habi a s ac oss Po ugal. We iden i ied 12 species
belonging o i e amilies, wi h Po cellionidae and A madillidiidae being he mos ep esen ed. The new
species, Eluma c is a a, endemic o he MSS o Alcobe as, is desc ibed. The e es ial isopod communi ies
included bo h su ace-dwelling and sub e anean-adap ed species, poin ing ou he connec i i y be ween
hese habi a s and highligh ing he adap abili y o e es ial isopods o di e se en i onmen s. Widesp ead
species occu ing in mul iple loca ions, and species con ined o single si es a e p esen . Seasonali y does
no seem o a ec he communi y composi ion. Ou indings highligh he ich di e si y o MSS habi a s,
hus hei impo ance o conse a ion.
Keywo ds
C us aceans, Ibe ian Peninsula, new species, shallow sub e anean habi a s, sub e anean ecosys ems
Sub e anean Biology 53: 129–153 (2025)
doi: 10.3897/sub biol.53.163632
h ps://sub biol.penso .ne
Copy igh Ri a P. Eusébio 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
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Ri a P. Eusébio e al. / Sub e anean Biology 53: 129–153 (2025)
130
In oduc ion
Sub e anean habi a s a e some o he leas explo ed ecosys ems in e ms o biodi e -
si y (Mammola e al. 2019, 2020). Among hese, he Meso oid Shallow Subs a um
(MSS), a ne wo k o oids o med by he accumula ion o ock agmen s in s eep
slopes, has ecei ed limi ed a en ion despi e i s ecological impo ance (Jube hie e al.
1980; Mammola e al. 2016).
While deep sub e anean habi a s such as ca es ha e inc easingly a ac ed scien-
i ic and public a en ion o hei biodi e si y and conse a ion alue (Mammola e al.
2019), MSS habi a s, despi e hei global dis ibu ion ac oss a ied geomo phological
se ings, emain poo ly s udied. Thei aunal composi ion and ecological dynamics a e
s ill poo ly unde s ood (Eusébio e al. 2021).
MSS habi a s a e cha ac e ized by ela i ely s able mic oclima ic condi ions, includ-
ing educed he mal luc ua ions compa ed wi h su ace habi a s. These ea u es p o-
ide bo h a e uge om su ace clima ic ex emes and a pe manen habi a o a di e se
ange o species (Ni zu e al. 2014; Mammola e al. 2016). These habi a s suppo ich
and specialized in e eb a e communi ies, including bo h sub e anean-adap ed and
su ace-dwelling species (Cul e and Pipan 2009, 2014; Mammola e al. 2016, 2017).
Te es ial isopods equen ly ound in sub e anean ecosys ems ha e limi ed dis-
pe sal abili ies and p e e ence o high mois u e (Ho nung 2011; Dixie e al. 2015).
Ve y ew pape s deal wi h e es ial isopods om he MSS (e.g., Ni zu e al. 2010, 2011;
Rebolei a e al. 2015, 2022; Ci uen es e al. 2021). In Po ugal, e es ial isopod auna
o sub e anean habi a s has been documen ed in ca es and spo adically in he MSS
(Vandel 1946; Rebolei a e al. 2015, 2022). Howe e , comp ehensi e da a on isopod di-
e si y, dis ibu ion and seasonal dynamics in MSS habi a s o Po ugal a e s ill lacking.
Sub e anean habi a s and hei specialized auna ace inc easing h ea s om
land-use change, global wa ming, and pollu ion (Mammola e al. 2019; Rebolei a
and Eusébio 2021, 2023; Rebolei a e al. 2022). While ca e in e eb a es ha e been
inc easingly included in conse a ion assessmen s such as he IUCN Red Lis (IUCN
2025) and he Po uguese Red Book o In e eb a es (ICNF 2023), species om shal-
low sub e anean habi a s emain o e looked, despi e hei ulne abili y and ecological
impo ance (Eusébio e al. 2021).
We s udied he di e si y, abundance, and seasonal dynamics o e es ial isopods in
MSS habi a s in Po ugal. We analysed new spa ial- empo al dis ibu ion pa e ns and de-
sc ibed a new species o he genus Eluma Budde-Lund, 1885. This in o ma ion is c ucial
o unde line he ele ance o he conse a ion o hese shallow sub e anean ecosys ems.
Ma e ial and me hods
Te es ial isopod sampling was conduc ed in i e MSS a eas ac oss Po ugal: Unhais
da Se a, Se a da Es ela (40°17'23.7"N, 7°35'20.1"W) a 1.330 m a.s.l, Vale dos
Poios, Sicó (39°58'49.3"N, 8°33'03.1"W) a 150 m, Alcobe as, Es emenho Ka s
Massi (39°25'46.8"N, 8°54'58.7"W) a 400 m, Cabanas de To es, Mon ejun o
Oniscidea (Isopoda) om MSS habi a s o Po ugal 131
(39°10'11.2"N, 9°04'14.1"W) a 420 m, and São Lou enço, A ábida (38°28'59.3"N,
9°00'09.9"W) a 300 m (Fig. 1).
Specimens we e collec ed using pi all aps modi ied o sample MSS habi a s
ollowing he me hod o Eusébio e al. (2021), wi h only one collec ing pi all cup
a a 50 cm dep h. In each loca ion i e aps we e ins alled, 10 m away om each
o he and 1 m away om he sh ub edge, using po k li e as bai and p opylene
glycol as a p ese ing liquid. Samples we e collec ed e e y h ee mon hs o one
yea (Ma ch 2022 – Ma ch 2023).
Te es ial isopod specimens we e iden i ied o species le el based on mo pho-
logical cha ac e s and he pe inen li e a u e, unde Wild M5 and M20 mic oscopes.
Specimens we e dissec ed and moun ed in mic o p epa a ions using Hoye ’s liquid.
Specimens we e d awn using a came a lucida, and pho og aphed using a
Leica DMC4500 came a, wi h a Leica Z6 APO mic oscope, and s acked wi h he
Leica LAS X so wa e. Backg ound o specimens’ images was cleaned using Adobe
Pho oshop (Ve sion 24.2.0).
Figu e 1. Dis ibu ion map. Sampling loca ions ma ked in colou ed iangles: blue – Se a da Es ela,
pu ple – Sicó, pink – Alcobe as, o ange – Mon ejun o, yellow – A ábida. The g een zones ep esen he
p o ec ed a eas o Po ugal.
Ri a P. Eusébio e al. / Sub e anean Biology 53: 129–153 (2025)
132
Da a analysis was done in R so wa e e sion 4.4.2 (R Co e Team 2024). Species
abundance was isualized using he ggplo unc ion (Wickham 2016). Non-me ic
mul idimensional scaling (NMDS) was used o isualize he spa ial and seasonal com-
muni y pa e ns, using he B ay-Cu is dissimila i y in he egan package (Oksanen e
al. 2025), only conside ing s ess alues below 0.2, which indica e a good da a ep e-
sen a ion o he wo-dimensional o dina ion space (Cla ke 1993).
Specimens a e deposi ed in he Na ional Museum o Na u al His o y and Science
(MNHNC), in he c us acean collec ion (MB11), Uni e si y o Lisbon, Po ugal, and
in he Museo di S o ia Na u ale “La Specola” (MZUF), Flo ence, I aly.
Resul s
Taxonomy
Class Malacos aca La eille, 1802
O de Isopoda La eille, 1817
Subo de Oniscidea La eille, 1802
Family T ichoniscidae Sa s, 1899
Genus T ichoniscoides Sa s, 1899
T ichoniscoides machadoi Vandel, 1946
Fig. 6A
T ichoniscoides machadoi Vandel, 1946: 168, igs 8–12; Vandel 1947: 270; Vandel
1948: 129, ig. 14;Vandel 1952: 262, igs 2, 9; Schmölze 1965: 37, igs 79–82;
Schmölze 1971: 4, 75, 139, map 4; Schmal uss 2003: 303; Rebolei a e al. 2015:
6, ig. 2; Ci uen es 2019: 48; Ci uen es and Da Sil a 2023: 7.
Ma e ial examined. Po ugal – Sicó • 2 ♂♂, 3 ♀♀; 39°58'51.8"N, 8°33'13.1"W;
19.XII.2022; leg. R.P. Eusébio; MSS pi all ap; MNHNC:MB11:001283, 001285,
001287 • 1 ♀; 20.III.2023; same da a as o p eceding; MNHNC:MB11:001379. –
A ábida • 1 ♂, 2 ♀♀; 38°28'59.2"N, 9°00'09.9"W; 21.VII.2022; leg. R.P. Eusébio;
MSS pi all ap; MNHNC:MB11:001141 • 3 ♂♂; 22.III.2023; same da a as o
p eceding; MNHNC:MB11:001313, 001323.
Dis ibu ion. Endemic o Po ugal. This species is widely dis ibu ed h oughou
he coun y.
Ecological no es. This species can be ound in ca es, MSS, and endogean habi a s.
Seasonal pa e ns in Po uguese MSS habi a s. This species was ound in wo
loca ions, Sicó and A ábida. In Sicó i was cap u ed du ing all and win e , wi h a
highe abundance in all (Fig. 7A). In A ábida i was cap u ed du ing sp ing and all
in equal abundances (Fig. 7A).
Oniscidea (Isopoda) om MSS habi a s o Po ugal 133
Family Philosciidae Kinahan, 1857
Genus C enoscia Ve hoe , 1928
C enoscia minima (Doll us, 1892)
Fig. 6B
Philoscia minima Doll us, 1892: 187, ig. 3.
Philoscia (Ben hana) minima Jackson, 1926: 196, pl. VIII igs 152–165.
C enoscia minima Vandel, 1946: 196, igs 42–49; Schmölze 1965: 141, ig. 605;
Schmölze 1971: 20; Schmal uss 2003: 87; Rebolei a e al. 2015: 40; Ci uen es
and Da Sil a 2023: 36.
C enoscia do salis; G ego y e al. 2012: 17.
Ma e ial examined. Po ugal – Sicó • 19 ♀♀; 39°58'51.8"N, 8°33'13.1"W;
20.VII.2022; leg. R.P. Eusébio; MSS pi all ap; MNHNC:MB11:001130,
001133, 001144, 001149, 001165 • 1 ♀; 19.IX.2022; same da a as o p eceding;
MNHNC:MB11:001256. – Alcobe as • 1 ♂, 3 ♀♀; 39°25'46.5"N, 8°54'59.0"W;
21.VII.2022; leg. R.P. Eusébio; MSS pi all ap; MNHNC:MB11:001185, 001187,
001202 • 6 ♀♀; 20.IX.2022; same da a as o p eceding; MNHNC:MB11:001205,
001212, 001218 • 3 ♀♀; 20.XII.2022; same da a as o p eceding;
MNHNC:MB11:001294, 001298 • 1 ♂, 2 ♀♀; 01.IV.2023; same da a as o
p eceding; MNHNC:MB11:001357, 001360. – Mon ejun o • 2 ♂♂, 13 ♀♀;
39°10'11.4"N, 9°04'14.8"W; 22.VII.2022; leg. R.P. Eusébio; MSS pi all ap;
MNHNC:MB11:001181, 001192 • 2 ♂♂, 3 ♀♀; 21.IX.2022; same da a as o p e-
ceding; MNHNC:MB11:001223, 001231 • 2 ♂♂; 21.XII.2022; same da a as o
p eceding; MNHNC:MB11:001340 • 2 ♀♀; 21.III.2023; same da a as o p eceding;
MNHNC:MB11:001385. – A ábida • 17 ♀♀, 1 ju .; 38°28'59.2"N, 9°00'09.9"W;
21.VII.2022; leg. R.P. Eusébio; MSS pi all ap; MNHNC:MB11:001137, 001151,
001154, 001172, 001176 • 4 ♂♂, 10 ♀♀, 2 ju s; 20.IX.2022; same da a as o p e-
ceding; MNHNC:MB11:001233, 001237, 001245 • 2 ♂♂, 11 ♀♀; 20.XII.2022;
same da a as o p eceding; MNHNC:MB11:001306, 001314, 001324.
Dis ibu ion. Po ugal, Spain, I aly, F ance (Co sica), Mal a, Tunisia, and Mo occo.
Ecological no es. This epigean species can also be ound in ca es and MSS.
Seasonal pa e ns in Po uguese MSS habi a s. This species was ound in ou
loca ions, Sicó, Alcobe as, Mon ejun o and A ábida. In Sicó i was cap u ed du ing
sp ing and summe , wi h a highe abundance in sp ing (Fig. 7B). In Alcobe as i was
cap u ed du ing all seasons, wi h abundance inc easing owa ds summe and hen de-
c easing ill win e (Fig. 7B). In Mon ejun o his species was also cap u ed du ing all
seasons, wi h abundance dec easing s eadily owa ds all and hen s aying he same in
win e (Fig. 7B). In A ábida i was cap u ed du ing sp ing, summe , and all, wi h a
s eady dec ease in abundance om sp ing o all (Fig. 7B).
Ri a P. Eusébio e al. / Sub e anean Biology 53: 129–153 (2025)
134
Family Po cellionidae B and , 1831
Genus Po cellionides Mie s, 1877
Po cellionides cingendus (Kinahan, 1857)
Fig. 6C
Po cellio cingendus Kinahan, 1857: 279, igs. 1, 2, 4, 6, 8, 9.
Po cellio (Po cellionides) lusi ano um A cangeli, 1935a: 21, igs 13–18.
Me opono hus (Lusi oniscus) cingendus Vandel, 1946: 274, igs 91–97; Schmölze
1965: 195, igs 698–699; Schmölze 1971: 33.
Po cellionides cingendus Schmal uss, 2003: 240; G ego y e al. 2012: 18; Rebolei a e
al. 2015; Ci uen es and Da Sil a 2024: 4, igs 1, 2A, 3A, 4A–D, 5, ab. 1.
Ma e ial examined. Po ugal – Sicó • 54 ♂♂, 58 ♀♀, 2 ju s; 39°58'51.8"N,
8°33'13.1"W; 20.VII.2022; leg. R.P. Eusébio; MSS pi all ap; MNHNC:MB11:001123,
001127, 001128, 001129, 001134, 001143, 001147, 001166 • 18 ♂♂, 38 ♀♀;
19.IX.2022; same da a as o p eceding; MNHNC:MB11:001255, 001257, 001258,
001259, 001262, 001263 • 32 ♂♂, 31 ♀♀; 19.XII.2022; same da a as o p eced-
ing; MNHNC:MB11:001275, 001278, 001282, 001284, 001288 • 30 ♂♂, 6 ♀♀;
20.III.2023; same da a as o p eceding; MNHNC:MB11:001372, 001374, 001376,
001381, 001383. – Alcobe as • 6 ♂♂, 21 ♀♀; 39°25'46.5"N, 8°54'59.0"W;
21.VII.2022; leg. R.P. Eusébio; MSS pi all ap; MNHNC:MB11:001188, 001197,
001199, 001201 • 13 ♂♂, 58 ♀♀; 20.IX.2022; same da a as o p ecedings;
MNHNC:MB11:001204, 001211, 001215, 001217 • 34 ♂♂, 46 ♀♀; 20.XII.2022;
same da a as o p eceding; MNHNC:MB11:001293, 001297, 001299, 001303 • 56
♂♂, 23 ♀♀; 01.IV.2023; same da a as o p eceding; MNHNC:MB11:001350, 001352,
001353, 001358. – Mon ejun o • 32 ♂♂, 44 ♀♀; 39°10'11.4"N, 9°04'14.8"W;
23.VII.2022; leg. R.P. Eusébio; MSS pi all ap; MNHNC:MB11:001161,
001178, 001180, 001190 • 3 ♂♂, 33 ♀♀; 21.IX.2022; same da a as o p eced-
ing; MNHNC:MB11:001220, 001222, 001226, 001230 • 55 ♂♂, 57 ♀♀, 2 ju s;
21.XII.2022; same da a as o p eceding; MNHNC:MB11:001329, 001338, 001339,
001346 • o e 500 specimens (♂ and ♀); 21.III.2023; same da a as o p eceding;
MNHNC:MB11:001387, 001393, 001397, 001400, 001403. – A ábida • 11 ♂♂,
17 ♀♀, 3 ju s; 38°28'59.2"N, 9°00'09.9"W; 21.VII.2022; leg. R.P. Eusébio; MSS
pi all ap; MNHNC:MB11:001136, 001142, 001155, 001171, 001175 • 18 ♂♂,
64 ♀♀, 46 ju s; 20.IX.2022; same da a as o p eceding; MNHNC:MB11:001234,
001236, 001242, 001246 • o e 200 specimens (♂ and ♀); 20.XII.2022; same da a
as o p eceding; MNHNC:MB11:001307, 001315, 001320, 001325, 001332 • 2
♂♂, 2 ♀♀, 1 ju .; 22.III.2023; same da a as o p eceding; MNHNC:MB11:001363,
001368.
Dis ibu ion. Uni ed Kingdom, F ance and Ibe ian Peninsula.
Ecological no es. This epigean species can also occu in ca es and MSS.
Oniscidea (Isopoda) om MSS habi a s o Po ugal 135
Seasonal pa e ns in Po uguese MSS habi a s. This species was ound in ou
loca ions, Sicó, Alcobe as, Mon ejun o and A ábida. Specimens we e ound du ing
all seasons in he ou loca ions. In Sicó he e was a s eady dec ease in abundance om
sp ing o win e , in Alcobe as he e was an inc ease om sp ing o win e , in Mon-
ejun o abundance dec eased om sp ing o summe and hen inc eased un il win e
(Fig. 7C). In A ábida abundance inc eased s eadily un il all and hen dec eased o
excep ionally low numbe s in win e (Fig. 7C).
Genus Po cellio La eille, 1804
Po cellio scabe La eille, 1804
Fig. 6D
Po cellio scabe La eille, 1804: 45; Doll us 1897: 206; Vandel 1946: 297; G ego y e
al. 2012: 20, ig. 6.
Po cellio (Po cellio) scabe A cangeli, 1935a: 9.
Po cellio (Eupo cellio) scabe a . scab io Ve hoe , 1907: 266.
Po cellio scabe scabe Schmölze , 1971: 39.
Ma e ial examined. Po ugal – Se a de Es ela • 1 ♂, 1 ♀; 40°17'22.9"N,
7°35'19.7"W; 20.VII.2022; leg. R.P. Eusébio; MSS pi all ap;
MNHNC:MB11:001167 • 1 ♂, 9 ♀♀; 19.IX.2022; same da a as o p eceding;
MNHNC:MB11:001240, 001248, 001249 • 1 ♂, 1 ♀; 19.XII.2022; same da a as
o p eceding; MNHNC:MB11:001269, 001273.
Dis ibu ion. Widesp ead in Eu ope and in oduced in o many pa s o he wo ld.
Ecological no es. This epigean species can also be ound in MSS.
Seasonal pa e ns in Po uguese MSS habi a s. This species was ound in one
loca ion, Se a da Es ela, du ing sp ing, summe , and all, and wi h abundance peak-
ing in summe (Fig. 7D).
Po cellio dila a us B and , 1831
Fig. 6E
Po cellio dila a us B and in B and and Ra zebu g 1831: 78, Pl. 12, ig. 6c, d; Vandel
1946: 316, ig. 135; Schmölze 1971: 44.
Po cellio (Po cellio) dila a us A cangeli, 1935a: 12.
Po cellio dila a us dila a us; Rebolei a e al. 2015: 46.
Ma e ial examined. Po ugal – Sicó • 1 ♀; 39°58'51.8"N, 8°33'13.1"W; 20.VII.2022;
leg. R.P. Eusébio; MSS pi all ap; MNHNC:MB11:001122 • 1 ♂; 19.XII.2022; same
da a as o p eceding; MNHNC:MB11:001274 • 1 ♂, 3 ♀♀; 20.III.2023; same da a as
o p eceding; MNHNC:MB11:001371, 001382. – Mon ejun o • 1 ♂; 39°10'11.4"N,
Ri a P. Eusébio e al. / Sub e anean Biology 53: 129–153 (2025)
136
9°04'14.8"W; 21.III.2023; leg. R.P. Eusébio; MSS pi all ap; MNHNC:MB11:001389.
– A ábida • 22 ♂♂, 41 ♀♀, 182 ju s; 38°28'59.2"N, 9°00'09.9"W; 21.VII.2022;
leg. R.P. Eusébio; MSS pi all ap; MNHNC:MB11:001135, 001138, 001140,
001150, 001152, 001158, 001169, 001170, 001173, 001174 • 2 ♂♂, 12 ♀♀, 20
ju s; 20.IX.2022; same da a as o p eceding; MNHNC:MB11:001235, 001238,
001243, 001247 • 28 ♂♂, 23 ♀♀, 32 ju s; 20.XII.2022; same da a as o p eced-
ing; MNHNC:MB11:001304, 001308, 001310, 001316, 001317, 001319, 001322,
001326, 001330, 001333 • 16 ♂♂, 13 ♀♀; 22.III.2023; same da a as o p eceding;
MNHNC:MB11:001361, 001364, 001365, 001366.
Dis ibu ion. Common ac oss Eu ope and in oduced in o many pa s o he wo ld.
Ecological no es. This species is mainly ound in ca es and MSS habi a s bu does
no show any adap a ion o sub e anean habi a s.
Seasonal pa e ns in Po uguese MSS habi a s. This species was ound in h ee
loca ions, Sicó, Mon ejun o and A ábida. In Sicó and Mon ejun o i was ound in
e y low numbe s (Fig. 7E). In A ábida i was cap u ed du ing all seasons, wi h abun-
dance dec easing om sp ing o summe , hen inc easing sligh ly in all and dec easing
again in win e (Fig. 7E).
Po cellio dispa Ve hoe , 1901
Fig. 6F
Po cellio dispa Ve hoe , 1901: 407; Vandel 1946: 289, igs 110–111; Schmölze 1965:
228; Schmölze 1971: 36; Schmal uss, 2003: 222; Ci uen es and Da Sil a 2023: 36.
Po cellio (Eupo cellio) dispa Ve hoe , 1907: 269.
Po cellionides dimo phus Jackson, 1926: 186, pl. I igs 1–22.
Po cellio (Po cellio) dispa A cangeli, 1935a: 10, igs 3–7.
Ma e ial examined. Po ugal – Sicó • 6 ♀♀; 39°58'51.8"N, 8°33'13.1"W;
20.VII.2022; leg. R.P. Eusébio; MSS pi all ap; MNHNC:MB11:001131, 001145,
001164 • 1 ♀; 19.IX.2022; same da a as o p eceding; MNHNC:MB11:001264 •
2 ♀♀; 19.XII.2022; same da a as o p eceding; MNHNC:MB11:001277 • 3 ♀♀;
20.III.2023; same da a as o p eceding; MNHNC:MB11:001373. – Mon ejun o • 2
♂♂, 4 ♀♀; 39°10'11.4"N, 9°04'14.8"W; 23.VII.2022; leg. R.P. Eusébio; MSS pi all
ap; MNHNC:MB11:001160, 001179 • 1 ♀; 21.XII.2022; same da a as o p eced-
ing; MNHNC:MB11:001343 • 3 ♂♂, 23 ♀♀; 21.III.2023; same da a as o p eced-
ing; MNHNC:MB11:001386, 001394.
Dis ibu ion. Po ugal, NW Spain, and F ance.
Ecological no es. This epigean species can also be ound in ca es and MSS.
Seasonal pa e ns in Po uguese MSS habi a s. This species was ound in wo
loca ions, Sicó and Mon ejun o. In Sicó i was cap u ed in all seasons, wi h abundance
dec easing om sp ing o summe , hen inc easing sligh ly un il win e (Fig. 7F). In
Mon ejun o i was cap u ed du ing sp ing, all and win e , wi h abundance dec easing
om sp ing o all and inc easing o i s highes in win e (Fig. 7F).
Oniscidea (Isopoda) om MSS habi a s o Po ugal 137
Po cellio no us A cangeli, 1935
Fig. 6G
Po cellio (Po cellio) no us A cangeli, 1935b: 40, igs 1–6.
Po cellio no us Vandel, 1946: 322; Schmölze 1955: 314; Schmölze 1965: 230, ig.
835; Schmölze 1971: 38; Schmal uss, 2003: 231.
Ma e ial examined. Po ugal – Alcobe as • 2 ♂♂, 3 ♀♀; 39°25'46.5"N, 8°54'59.0"W;
21.VII.2022; leg. R.P. Eusébio; MSS pi all ap; MNHNC:MB11:001196 • 6 ♂♂,
10 ♀♀, 1ju .; 20.IX.2022; same da a as o p eceding; MNHNC:MB11:001208,
001210, 001214, 001219 • 1 ♂, 3 ♀♀; 20.XII.2022; same da a as o p eceding;
MNHNC:MB11:001290, 001292 • 2 ♀♀; 1.IV.2023; same da a as o p eceding;
MNHNC:MB11:001355.
Dis ibu ion. Po ugal and wes e n Spain.
Ecological no es. This epigean species can also be ound in MSS.
Seasonal pa e ns in Po uguese MSS habi a s. This species was ound in one
loca ion, Alcobe as, du ing all seasons, wi h abundance inc easing om sp ing o
summe , hen dec easing s eadily owa ds win e (Fig. 7G).
Family A madillidiidae B and , 1833
Genus Eluma Budde-Lund, 1885
Eluma caela a (Mie s, 1877)
Fig. 6H
A madillidium caela um Mie s, 1877: 665, pl. 7 ig. 3.
Rhacodes insc ip us Ea on, 1882: 360 (no Rhacodes insc ip us C.L. Koch in Rosenhaue
1856: 422).
Eluma pu pu ascens Budde-Lund, 1885: 48, 294; P eudhomme de Bo e 1886: 112;
Doll us 1889a: 126; Doll us 1889b: 306; Doll us 1896a: 357; No man 1899: 69;
Ve hoe 1908: 370; Vandel 1946: 357, ig. 158.
Eluma caela um A cangeli, 1935a: 3; A cangeli 1948: 242; Schmölze 1965: 303, ig.
1213; Schmölze 1971: 58; Schmal uss 2003: 107.
Eluma caela a G ego y e al., 2012: 21, ab. 2, 3; Rebolei a e al. 2015: 47; Recue o e
al. 2022: 1315, ab. 1; Ci uen es and Da Sil a 2023: 29, igs 1A, 2A-C, 4G.
Ma e ial examined. Po ugal – Se a de Es ela • 1 ♀; 40°17'22.9"N, 7°35'19.7"W;
20.VII.2022; leg. R.P. Eusébio; MSS pi all ap; MNHNC:MB11:001121 • 1 ♂,
1 ♀; 19.IX.2022; same da a as o p eceding; MNHNC:MB11:001250 • 5 ♂♂, 7
♀♀; 19.XII.2022; same da a as o p eceding; MNHNC:MB11:001268, 001270,
001271, 001272 • 4 ♂♂, 5 ♀♀; 19.XII.2022; same da a as o p eceding; MZUF
10034 • 1 ♀; 20.III.2023; same da a as o p eceding; MNHNC:MB11:001347.
– Sicó • 3 ♀♀; 39°58'51.8"N, 8°33'13.1"W; 20.VII.2022; leg. R.P. Eusébio;
Ri a P. Eusébio e al. / Sub e anean Biology 53: 129–153 (2025)
144
Figu e 5. Eluma c is a a sp. no . Pa a ype ♂ om Alcobe as A pe eopod 1 B pe eopod 7 C geni al
papilla and pleopod 1 D pleopod 2 E pleopod 3 exopod F pleopod 4 exopod G pleopod 5 exopod.
sho e do sal scale-se ae o di e en shape (see ig. 4A–F in Ci uen es and Da
Sil a 2023).
Ecological no es. This species was only ound in his MSS habi a .
Seasonal pa e ns in Po uguese MSS habi a s. This species was only ound in
Alcobe as, du ing all seasons, and wi h a dec ease in abundance om sp ing o sum-
me , highes abundance du ing all and a new dec ease in win e (Fig. 7K).
Oniscidea (Isopoda) om MSS habi a s o Po ugal 145
Family A madillidae B and , 1831
Genus A madillo La eille, 1802
A madillo o icinalis Dumé il, 1816
Fig. 6L
A madillo o icinalis Dumé il, 1816: 117; Budde-Lund 1885: 16; P eudhomme de
Bo e 1886: 112; Doll us 1896a: 357; Doll us 1896b: 527; A cangeli 1935a: 2;
Vandel 1946: 371.
Cuba is globa o ; Ea on 1882: 361.
Cuba is o icinalis; Jackson 1926: 184.
Ma e ial examined. Po ugal – A ábida • 28 ♂♂, 63 ♀♀; 38°28'59.2"N, 9°00'09.9"W;
21.VII.2022; leg. R.P. Eusébio; MSS pi all ap; MNHNC:MB11:001139, 001153,
001156, 001168 • o e 1000 indi iduals (♂ and ♀); 20.IX.2022; same da a as o
p eceding; MNHNC:MB11:001232, 001239, 001241, 001244 • 84 ♂♂, 140 ♀♀,
2 ju s; 20.XII.2022; same da a as o p eceding; MNHNC:MB11:001305, 001309,
001312, 001318, 001321, 001331, 001334 • 1 ♀; 22.III.2023; same da a as o p e-
ceding; MNHNC:MB11:001362.
Dis ibu ion. Widely dis ibu ed in he Medi e anean and Black Sea a ea.
Ecological no es. This epigean species can also be ound in MSS.
Seasonal pa e ns in Po uguese MSS habi a s. This species was only ound in
A ábida, du ing all seasons, and abundance inc easing om sp ing o summe whe e
i eaches i s peak, hen dec easing in all (Fig. 7L).
Ecological analysis
Te es ial isopod communi ies di e ac oss loca ions and seasons (Fig. 8). All species
we e ound in ei he h ee o ou o he sampled seasons (Fig. 7). The spa ial dis i-
bu ion plo (Fig. 8A) shows ha hese communi ies a e ela i ely di e en om each
o he among he i e sampled loca ions. Se a da Es ela shows a dis inc ly di e en
isopod communi y composi ion compa ed wi h o he loca ions, la gely d i en by he
s ong in luence o Po cellio scabe (Fig. 8A). Alcobe as also has a dis inc communi y
composi ion, cha ac e ized p ima ily by Eluma c is a a and Po cellio no us (Fig. 8A).
In con as , he o he h ee loca ions (Sicó, A ábida, and Mon ejun o) show a g ea e
o e lap in species composi ion, sugges ing mo e simila isopod communi ies (Fig. 8A).
The seasonal si e-speci ic plo s e eal a ying deg ees o a ia ion in isopod
communi ies (Figs 8B–F). In Se a da Es ela (Fig. 8B), isopod communi ies a e
composed only by wo species wi h no clea seasonal pa e n. In Sicó (Fig. 8C),
Alcobe as (Fig. 8D) and Mon ejun o (Fig. 8E) he seasonal ellipses o e lap almos
comple ely, indica ing simila isopod communi ies h oughou he yea in each loca-
ion. In A ábida (Fig. 8F) he e is some dema ked seasonal a ia ion, wi h A madillo
o icinalis and Po cellionides cingendus in luencing summe and all, and he o he
ou species in luencing sp ing.
Ri a P. Eusébio e al. / Sub e anean Biology 53: 129–153 (2025)
146
Figu e 6. Specimen habi us A T ichoniscoides machadoi om Sicó B C enoscia minima om Mon ejun o
C Po cellionides cingendus om Sicó D Po cellio scabe om Se a da Es ela E Po cellio dila a us om
A ábida F Po cellio dispa om Sicó G Po cellio no us om Alcobe as H Eluma caela a om Mon ejun o
I Eluma ube cula a om Sicó J Eluma ma ae om Mon ejun o K Eluma c is a a sp. no . om Alcobe as
L A madillo o icinalis om A ábida.
Discussion
We ound 12 species o e es ial isopods in Po uguese MSS habi a s, belonging o
i e amilies, h ee mo e species and wo mo e amilies han hose eco ded in a simila
s udy done in Spain (Ci uen es e al. 2021). The mos well ep esen ed amily is Po -
cellionidae wi h i e species, ollowed by A madillidiidae wi h ou , and he emaining
amilies a e only ep esen ed by one species each. This s udy e ealed a new species,
Eluma c is a a sp. no ., ound exclusi ely in he MSS o Alcobe as.
Oniscidea (Isopoda) om MSS habi a s o Po ugal 147
Figu e 7. To al abundance pe season o each o he species. Each colou ep esen s one sampling loca-
ion: blue – Se a da Es ela, pu ple – Sicó, pink – Alcobe as, o ange – Mon ejun o, yellow – A ábida
A T ichoniscoides machadoi B C enoscia minima C Po cellionides cingendus D Po cellio scabe E Po cellio
dila a us F Po cellio dispa G Po cellio no us H Eluma caela a I Eluma ube cula a J Eluma ma ae K Eluma
c is a a sp. no . L A madillo o icinalis.
Ri a P. Eusébio e al. / Sub e anean Biology 53: 129–153 (2025)
148
Figu e 8. NMDS (Non-me ic mul idimensional scaling) o e es ial isopod communi ies A species
ound in each o he i e loca ions, wi h each poin ep esen ing a species and each a ow ep esen ing a
loca ion B–F loca ion speci ic seasonal NMDS B Se a da Es ela C Sicó D Alcobe as E Mon ejun o
F A ábida. In hese panels each poin ep esen s one in e eb a e ap ( i e aps pe season), ellipses
g oup he poin s by season, and he a ows ep esen he species.
Oniscidea (Isopoda) om MSS habi a s o Po ugal 149
The di e si y o e es ial isopods ound in MSS habi a s o Po ugal includes
bo h su ace-dwelling species, such as C enoscia minima, Po cellionides cingen-
dus, Po cellio scabe , P. dispa , P. no us, Eluma caela a, E. ma ae, and A madillo
o icinalis, and species mo e equen ly ound in he su ace-ca e eco one, such as
T ichoniscoides machadoi, Eluma ube cula a and Po cellio dila a us (Vandel 1946;
Rebolei a e al. 2015). This emphasises he connec i i y be ween su ace and sub-
e anean habi a s and e lec s he b oad ecological ole ance o e es ial isopods
and hei p e e ence o c yp ic and da k en i onmen s (Ho nung 2011; Dixie e
al. 2015). On he o he hand, he disco e y o Eluma c is a a sp. no ., highligh s
he ele ance o shallow sub e anean habi a s as habi a s o sho ange dis ib-
u ed species and high conse a ion alue.
The spa ial and seasonal dis ibu ion o e es ial isopod species ound in Po -
uguese MSS habi a s displayed dis inc pa e ns in abundance h oughou he yea .
Widesp ead species such as C enoscia minima and Po cellionides cingendus we e ound
in mul iple loca ions and h oughou all seasons, while Eluma c is a a sp. no . and Po -
cellio no us we e es ic ed o single MSS si es. The use o MSS habi a s by e es ial
isopod species di e s depending on he loca ion and species. Despi e se e al species
showing s ong seasonal abundance pa e ns, seasonali y does no seem o in luence
communi y composi ion in he majo i y o he s udied loca ions.
Gi en he obse ed high di e si y and ecological specializa ion, conse a ion
o MSS habi a s and hei isopod communi ies is c i ical, no only in Po ugal bu
wo ldwide. Despi e being legally p o ec ed (ICNB 2000) and acing inc easing h ea s
(Mammola e al. 2019; Rebolei a e al. 2022), MSS habi a s and hei auna emain
la gely o e looked in biodi e si y assessmen s. The e o e, hei s udy and p o ec ion
a e essen ial o sa egua d he unique in e eb a e di e si y hey ha bou .
Acknowledgmen s
This wo k was suppo ed by he Po uguese Na ional Funds h ough “Fundação pa a
a Ciência e a Tecnologia” wi hin he CE3C Uni unding ( e . UIDB/00329/2025),
CHANGE ( e . LA/P/0121/2020) (h ps://doi.o g/10.54499/LA/P/0121/2020) and
PhD g an ( e . 2021.04868.BD) (h ps://doi.o g/10.54499/2021.04868.BD), and
by he Chai in Sus ainabili y o Sub e anean Ecosys ems – Loulé, a he Facul y o
Sciences o he Uni e si y o Lisbon.
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