Rock sponges (li his id Demospongiae) o
he No heas A lan ic seamoun s, wi h
desc ip ion o en new species
F ancisca C. Ca alho
1
, Paco Cá denas
2
, Pila Ríos
3,4
, Ja ie C is obo
3,4
,
Hans To e Rapp
1
and Joana R. Xa ie
1,5
1Depa men o Biological Sciences and K.G. Jebsen Cen e o Deep-Sea Resea ch, Be gen
Uni e si y, Be gen, No way
2Pha macognosy, Depa men o Medicinal Chemis y, Uppsala Uni e si y, Uppsala, Sweden
3Cen o Oceanog áfico de Gijón, Ins i u o Español de Oceanog afia, Gijón, Spain
4Depa amen o de Zoología y An opología Física, Uni e sidad de Alcalá de Hena es, Mad id,
Spain
5CIIMAR-In e disciplina y Cen e o Ma ine and En i onmen al Resea ch, Uni e sidade do
Po o, Ma osinhos, Po ugal
ABSTRACT
Backg ound: Li his id demosponges, also known as ock sponges, a e a polyphyle ic
g oup o sponges which a e widely dis ibu ed. In he No heas A lan ic (NEA), 17
species a e known and he cu en knowledge on hei dis ibu ion is mainly
es ic ed o he Maca onesian islands. In he Medi e anean Sea, 14 species a e
eco ded and gene ally ound in ma ine ca es.
Me hods: Li his ids we e sampled in nine NEA seamoun s du ing he scien ific
expedi ions Seamoun 1 (1987) and Seamoun 2 (1993) o ganized by he MNHN o
Pa is. Collec ed specimens we e iden ified h ough he analyses o ex e nal and
in e nal mo phological cha ac e s using ligh and scanning elec on mic oscopy, and
compa ed wi h ma e ial om a ious museum collec ions as well as li e a u e
eco ds.
Resul s: A o al o 68 specimens we e analysed and a ibu ed o 17 species ac oss
wo o de s, se en amilies, and se en gene a, ep esen ing new eco ds o
dis ibu ion. Ten o hese species a e new o science, iz. Neosch ammeniella
inaequalis sp. no ., N. pise ai sp. no ., N. pomponiae sp. no ., Discode mia a bo sp.
no ., D. kellyae sp. no ., Macand ewia schus e ae sp. no ., M. minima sp. no .,
Exsupe an ia le ii sp. no ., Leiode ma ium uba sp. no . and Siphonidium elonga us
sp. no ., and a e he e desc ibed and illus a ed. New ba hyme ic eco ds we e also
ound o D. ami e a,D. e ucosa and M. obus a. The Me eo seamoun g oup
has a highe species ichness (15 species) compa ed o he Lusi anian seamoun
g oup (six species). The majo i y o he species had hei dis ibu ion es ic ed o one
seamoun , and en a e only known om a single locali y, bu his can be a esul o
sample bias.
Discussion: The numbe o species sha ed be ween he seamoun s and he
Maca onesian islands is e y educed. The same pa e n epea s be ween he
NEA and Medi e anean Sea. This s udy demons a es ha NEA seamoun s a e
ecosys ems wi h a highe di e si y o li his ids han p e iously hough , inc easing
he numbe o li his ids known o occu in he NEA and Medi e anean Sea om 26
o 36 species.
How o ci e his a icle Ca alho FC, Cá denas P, Ríos P, C is obo J, Rapp HT, Xa ie JR. 2020. Rock sponges (li his id Demospongiae) o
he No heas A lan ic seamoun s, wi h desc ip ion o en new species. Pee J 8:e8703 DOI 10.7717/pee j.8703
Submi ed 9 Oc obe 2019
Accep ed 6 Feb ua y 2020
Published 7 Ap il 2020
Co esponding au ho
F ancisca C. Ca alho,
[email p o ec ed]
Academic edi o
Joseph Pawlik
Addi ional In o ma ion and
Decla a ions can be ound on
page 79
DOI 10.7717/pee j.8703
Copy igh
2020 Ca alho e al.
Dis ibu ed unde
C ea i e Commons CC-BY 4.0
Subjec s Biodi e si y, Biogeog aphy, Ma ine Biology, Taxonomy, Zoology
Keywo ds Po i e a, Deep-sea, Li his ids, Biodi e si y, Te ac inellida, Buba ida, New species,
Biogeog aphy
INTRODUCTION
The class Demospongiae Sollas (1885) con ains se e al g oups o sponges a ificially
unified unde he name ‘li his id demosponges’o ‘ ock sponges.’Li his ids p oduce
hype silicified spicules (desmas) (Pise a & Lé i, 2002a) ha usually c ea es a e y igid
skele on. Fo a e y long ime, hey we e classified in o an o de , Li his ida (Schmid , 1870),
bu mo e ecen ly, se e al s udies ha e shown he polyphyle ic na u e o his g oup
(Cá denas e al., 2011;Kelly & Pomponi, 1994;Pise a & Lé i, 2002a;Schus e e al., 2015).
I is now acknowledge ha his ai , i.e., is he desmas, has e ol ed independen ly
mul iple imes (Schus e e al., 2015) and he 211 alid species cu en ly ecognized
wo ldwide a e dis ibu ed in h ee o de s-Te ac inellida Ma shall (1876), Sphae ocladina
Sch ammen (1924) and Buba ida Mo ow & Cá denas (2015), wi h he la ge majo i y
belonging o he o me o de (Mo ow & Cá denas, 2015;Pise a & Lé i, 2002a;Schus e
e al., 2015;Van Soes e al., 2019, WPD).
In he No heas A lan ic (NEA), he cu en s a e o knowledge on li his id sponges is
mainly es ic ed o he Maca onesian islands. So a , 17 species ha e been desc ibed
and eco ded om he Azo es (Ca alho & Pise a, 2019;G ay, 1859;Topsen , 1928,1904,
1898,1892), Madei a and Sel agens (Bowe bank, 1869;Ca e , 1873;Ca alho & Pise a,
2019;Johnson, 1863), Cana y Islands (Ca alho & Pise a, 2019;C uz, 2002;Topsen , 1892),
Po ugal mainland (Schmid , 1870) and Mo occo (Lenden eld, 1907), whe eas in he
Medi e anean Sea, 15 species ha e been epo ed (Maldonado e al., 2015;Manconi, Se usi
& Pise a, 2006;Manconi & Se usi, 2008;Pe ez e al., 2004;Pise a & Vacele , 2011;
Puli ze -Finali, 1972;Vacele , 1969). They a e commonly ound on ha d subs a e a
110–1,700 m dep h (Ca e , 1873;Ca alho, Pomponi & Xa ie , 2015;Topsen , 1928),
whe eas in he Medi e anean Sea hey usually occu in shallowe wa e s o in ca e
sys ems (Manconi & Se usi, 2008;Pise a & Vacele , 2011). Al hough he knowledge on
dis ibu ion o li his ids in he NEA has been inc easing, he e is no da a ega ding hei
occu ence on seamoun s in he a ea.
These opog aphic ea u es, which p o ide impo an habi a s o bo h ben hic and
pelagic o ganisms, a e e y nume ous and wo ldwide dis ibu ed (Yesson, 2011). In he
NEA, examples include he Lusi anian Seamoun s (Co al Pa ch, Ampe e, Go inge Bank,
Hi ondelle II, Josephine, Lion, D agon, Unico n and Seine), loca ed nea he Eu o-A ican
con inen al shel , app oxima ely 250 km om he Po uguese coas and he Me eo
Seamoun s (G ea Me eo , Hyè es, I ing, C uise , Pla o, Ty o and A lan is), si ua ed in
he cen al pa o he No h A lan ic, close o he Mid-A lan ic Ridge (MAR) and sou h o
he Azo es a chipelago. These seamoun s ha e e oked in e es o esea ch in he la e
19 h and ea ly 20 h Cen u y, and se e al scien ific expedi ions ook place, such as
Josephine (1869), Challenge (1873) and nume ous P ince Albe I o Monaco expedi ions.
La e in he 20 h and ea ly 21s Cen u ies, new e o s aiming o explo e he ben hic auna
o hese seamoun s we e unde aken. Two o hese expedi ions—Seamoun 1 and
Ca alho e al. (2020), Pee J, DOI 10.7717/pee j.8703 2/87
Seamoun 2—o ganized by he Na u al His o y Museum o Pa is (MNHN), su eyed
a ious o he Lusi anian and Me eo seamoun s a dep hs abo e 1,000 m (Bouche &
Mé i ie , 1988;Go as, 1993). These expedi ions esul ed in he disco e y and desc ip ion o
se e al species o a ious axonomic g oups, such as b achiopods (Logan, 1998), b yozoans
(Be ning, Ha melin & Bade , 2017;Sou o, Be ning & Os o sky, 2016), bi al es
(Dijks a & Go as, 2004), co als (Molod so a & Shi sho , 2011), ci ipeds (Young, 2001),
hyd ozoans (Ramil, Ve oo & Ansín, 1998), polychae es (Gille & Dau in, 2003;
Pax on & Gille , 2004) and gas opods (Go as, 2007) g ea ly ad ancing he unde s anding
o he biogeog aphic pa e ns and he biodi e si y o hese ecosys ems. Howe e , se e al
axonomic g oups, including sponges, emain sca cely documen ed in he li e a u e o
hese ecosys ems (Cá denas e al., 2018;C is obo e al., 2015;Lé i & Vacele , 1958;Topsen ,
1928;Xa ie & Van Soes , 2007).
In his s udy, we desc ibe he li his id demosponges collec ed du ing he F ench
expedi ions Seamoun 1 and Seamoun 2. New eco ds o geog aphic dis ibu ion a e
epo ed, en new species o science a e desc ibed and illus a ed, and he di e si y and
biogeog aphic pa e ns discussed. An iden ifica ion key o all li his id species epo ed o
he NEA and Medi e anean is also p o ided.
MATERIALS AND METHODS
The ma e ial examined in his s udy was collec ed du ing Seamoun 1 and Seamoun 2
scien ific expedi ions unde aken by he MNHN o Pa is o se e al NEA seamoun s
(Fig. 1;Supplemen al Ma e ial S1). The main aims o hese campaigns we e o s udy he
pa e ns o aunal di e si y and endemism ound on isola ed seamoun s in compa ison o
con inen al a eas and he ela ion wi h he dispe sal capaci y o he a ious axonomic
g oups. The Seamoun 1 campaign, coo dina ed by D . Philippe Bouche , ook place in
1987 onboa d o he esea ch essel L. No oî , and explo ed he Galicia Banks and he
Lusi anian Seamoun s (Go inge, Josephine, Ampè e, Lion and Seine) (Bouche &
Mé i ie , 1988). The second campaign, Seamoun 2, his ime lead by D . Se ge Go as,
explo ed he Me eo Seamoun s g oup (G ea Me eo , Hyè es, I ing, C uise , Pla o,
A lan is and Ty o) and he An ial ai Seamoun on boa d o he RV L. Su oî , sampling 165
s a ions also a dep hs abo e 1,000 m (Go as, 1993). Li his ids we e collec ed in 10 s a ions
on Seamoun 1 (11%) and in 42 s a ions on Seamoun 2 (32%) be ween 280 and 1,035 m
dep h using a ious sampling gea s (beam awl (CP), epiben hic d edge (DE) and Wa én
d edge (DW)), and p ese ed in o malin onboa d. The specimens examined a e deposi ed
in he ‘zoo hèque’o he MNHN in Pa is, and s o ed a oom empe a u e in e hanol 70%.
De ailed in o ma ion ega ding he collec ion o he specimens s udied he e, is deposi ed in
PANGAEA
Ò
Da a Publishe (www.pangaea.de) unde he digi al objec iden ifie (DOI):
h ps://doi.pangaea.de/10.1594/PANGAEA.896492.
The specimens we e analysed h ough he use o Ligh Mic oscopy (LM) and Scanning
Elec on Mic oscopy (SEM). Fo ligh mic oscopy, c oss sec ions and slides o loose
spicules we e moun ed in Canada Balsam
Ò
Sigma–Ald ich o Euki
Ò
Sigma–Ald ich
ollowing s anda ds p ocedu es (Bou y-Esnaul & Ru zle , 1997). In addi ion, a ew
specimens, ep esen a i e o each species, we e selec ed and p epa ed o SEM. Fo his
Ca alho e al. (2020), Pee J, DOI 10.7717/pee j.8703 3/87
pu pose, pieces o bo h he ec osome and choanosome o he sponge we e excised and hen
ei he di ec ly moun ed o diges ed in ni ic acid, washed se e al imes wi h dis illed wa e
and hen fixed in e hanol. The spicules we e hen placed on a s ub and co e ed wi h
gold-paladium. Thi y spicules o each spicule ype we e measu ed using he Leica
Applica ion Sui e (LAS . 4.5), o indi idual specimens. Minimum, mean and maximum
alues a e p esen ed o he measu emen s ob ained o each analysed specimen.
Fo he highe axa classifica ion, we ollowed he e ised Demospongiae classifica ion
(Mo ow & Cá denas, 2015).
Due o he o malin fixa ion, we we e no able o ex ac DNA o molecula analysis,
and any a emp s o ba code he mi ochond ial COI gene, including he mini-ba code
p o ocol used in o he e ac inellids (Cá denas & Moo e, 2017) we e unsuccess ul.
The elec onic e sion o his a icle in Po Table Documen Fo ma (PDF) will
ep esen a published wo k acco ding o he In e na ional Commission on Zoological
Nomencla u e (ICZN), and hence he new names con ained in he elec onic e sion a e
e ec i ely published unde ha Code om he elec onic edi ion alone. This published
wo k and he nomencla u al ac s i con ains ha e been egis e ed in ZooBank, he online
egis a ion sys em o he ICZN. The ZooBank Li e Science Iden ifie s (LSIDs) can be
esol ed and he associa ed in o ma ion iewed h ough any s anda d web b owse by
G ea Me eo
Hyè es
Pla o
Go inge Bank
Ty o
A z o e s
A lan is
Lion
Ampè e
M a d e i a
C a n a i e s
An ial ai
Figu e 1 Map o he s udy a ea. Seamoun s o he No heas A lan ic and s a ions o he Seamoun 1
and Seamoun 2 campaigns whe e li his id demosponges we e collec ed. Map p oduced wi h he so wa e
QGIS De elopmen Team (2019); ba hyme y ob ained om GEBCO Compila ion G oup (2019).
Full-size

DOI: 10.7717/pee j.8703/fig-1
Ca alho e al. (2020), Pee J, DOI 10.7717/pee j.8703 4/87
appending he LSID o he p efixh p://zoobank.o g/. The LSID o his publica ion is:
u n:lsid:zoobank.o g:pub:A0DA0236-4579-47A4-8BE4-E68803C2EC8F. The online
e sion o his wo k is a chi ed and a ailable om he ollowing digi al eposi o ies: Pee J,
PubMed Cen al and CLOCKSS.
RESULTS
In his s udy we analysed 68 specimens, collec ed be ween 280 and 1,035 m dep h on eigh
NEA seamoun s, and assigned hem o 17 species dis ibu ed ac oss wo o de s, se en
amilies, and se en gene a (Figs. 2–3). O hese, en species a e new o science—
Neosch ammeniella inaequalis sp. no ., N. pise ai sp. no ., N. pomponiae sp. no .,
Discode mia a bo sp. no ., D. kellyae sp. no ., Macand ewia schus e ae sp. no .,
M. minima sp. no ., Exsupe an ia le ii sp. no ., Leiode ma ium uba sp. no . and
Siphonidium elonga us sp. no (see below desc ip ions and illus a ions). All analysed
ma e ial is desc ibed and illus a ed below and compa ed wi h addi ional specimens om
a ious museum collec ions (MNHN, HBOI, RMNH and DOP). An iden ifica ion key o
all li his id species eco ded o da e o he NEA and MED is also p o ided. All new species
desc ibed he e ha e he axonomic au ho i y es ic ed o he fi s and las au ho .
SYSTEMATIC INDEX
Phylum Po i e a G an , 1836
Class Demospongiae Sollas, 1885
Subclass He e oscle omo pha Cá denas, Pé ez & Bou y-Esnaul , 2012
O de Te ac inellida Ma shall, 1876
Subo de As opho ina Sollas, 1887
Family Co allis idae Sollas, 1888
Genus Neosch ammeniella Pise a & Lé i, 2002b
Species Neosch ammeniella inaequalis sp. no .
Species Neosch ammeniella pise ai sp. no .
Species Neosch ammeniella pomponiae sp. no .
Family Theonellidae Lenden eld, 1903
Genus Discode mia du Bocage, 1869
Species Discode mia ami e a Topsen , 1892
Species Discode mia c . ami e a Topsen , 1892
Species Discode mia e ucosa Topsen , 1928
Species Discode mia a bo sp. no .
Species Discode mia kellyae sp. no .
Family Macand ewiidae Sch ammen, 1924
Genus Macand ewia G ay, 1859
Species Macand ewia c . azo ica G ay, 1859
Species Macand ewia obus a Topsen , 1904
Species Macand ewia schus e ae sp. no .
Species Macand ewia minima sp. no .
Family Phyma aphiniidae Sch ammen, 1924
Ca alho e al. (2020), Pee J, DOI 10.7717/pee j.8703 5/87

1 cm 1 cm
1 cm
1 cm
5 mm
1 cm
1 cm
1 cm
1 cm
1 cm
ABC
D
EF
G
H
I
J
K
1.5 cm
L
1 cm
Figu e 2 Specimens collec ed du ing Seamoun 1 and Seamoun 2 expedi ions. (A) Top iew o
Neosch ammeniella inaequalis sp. no ., holo ype MNHN-IP-2018-84, (B) bo om iew o N. inaequalis
sp. no ., holo ype MNHN-IP-2018-84. (C) N. pise ai sp. no ., holo ype MNHN-IP-2008-234.
(D) N. pomponiae sp. no ., holo ype MNHN-IP-2008-233. (E) Discode mia ami e a Topsen , 1892,
specimen MNHN-IP-2008-213. (F) D. e ucosa Topsen , 1928, specimen MNHN-IP-2008-205.
(G) D. a bo sp. no ., holo ype MNHN-IP-2008-211. (H) D. kellyae sp. no ., holo ype MNHN-IP-2008-
208. (I) Macand ewia c . azo ica, specimen MNHN-IP-2008-220. (J) M. obus a Topsen , 1904, speci-
mens MNHN-IP-2008-216. (K) M. schus e ae sp. no ., holo ype MNHN-IP-2018-87. (L) M. minima sp.
no ., holo ype MNHN-IP-2008-222. Full-size
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DOI: 10.7717/pee j.8703/fig-2
Ca alho e al. (2020), Pee J, DOI 10.7717/pee j.8703 6/87
Genus Exsupe an ia Özdikmen, 2009
Species Exsupe an ia a chipelagus Ca alho & Pise a, 2019
Species Exsupe an ia le ii sp. no .
Subo de Spi opho ina Be gquis & Hogg, 1969
Family Azo icidae Sollas, 1888
Genus Leiode ma ium Schmid , 1870
1 cm
1 cm
1 cm
1 cm
1 cm
AB
CD
EF
1 cm
Figu e 3 Specimens collec ed du ing Seamoun 1 and Seamoun 2 expedi ions. (A) Exsupe an ia
a chipelagus Ca alho & Pise a (2019), specimen MNHN-IP-2008-196. (B) E. le ii sp. no ., holo ype
MNHN-IP-2008-201. (C) Leiode ma ium lynceus Schmid (1870), specimen MNHN-IP-2008-239.
(D) L. uba sp. no ., holo ype MNHN-IP-2018-72. (E) Siphonidium elonga us sp. no ., holo ype MNHN-
IP-2008-236. (F) Pe omica (Pe omica) g imaldii Topsen , 1898, MNHN-IP-2018-92.
Full-size
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DOI: 10.7717/pee j.8703/fig-3
Ca alho e al. (2020), Pee J, DOI 10.7717/pee j.8703 7/87
Species Leiode ma ium lynceus Schmid , 1870
Species Leiode ma ium uba sp. no .
Family Siphonidiidae Lenden eld, 1903
Genus Siphonidium Schmid , 1879
Species Siphonidium elonga us sp. no .
O de Buba ida Mo ow & Cá denas, 2015
Family Desman hidae Topsen , 1893
Genus Pe omica Topsen , 1898
Subgenus Pe omica (Pe omica)Topsen , 1898
Species Pe omica (Pe omica)g imaldii Topsen , 1898
Species desc ip ions
O de TETRACTINELLIDA Ma shall, 1876
Subo de ASTROPHORINA Sollas, 1887
Family CORALLISTIDAE Sollas, 1888
Genus Neosch ammeniella Pise a & Lé i, 2002b
Synonymy. Iouea sensu Lé i & Lé i, 1988: 248.
Diagnosis. Co allis idae wi h smoo h dicho iaenes and wo o h ee ypes o mic oscle es:
me as e s, amphias e s/s ep as e s and/o spi as e s (emended a e Kelly, 2007;Pise a &
Lé i, 2002b;Pise a & Vacele , 2011;Schlache -Hoenlinge , Pise a & Hoope , 2005).
Defini ion. Polymo phic Co allis idae, shallow cup-shaped o deep ase-shaped; su ace
can be smoo h o ugose; ec osomal megascle es a e smoo h dicho iaenes; choanosomal
megascle es a e dic anoclone desmas wi h di e en ypes o o namen a ion, a ying
om poo ly o ex emely ube cula ed in di e en species; diac ines a e equen ly p esen
in he ec osome and iaenes a e a e; mic oscle es a e me as e s, amphias e /s ep as e
and/o acan hose spi as e s ( ype I co e ed by sho blun ays, and ype II i egula wi h
sho blun ays only on he edges), bu he numbe and ype o mic oscle es a ies
be ween species (emended a e Kelly, 2007;Pise a & Lé i, 2002b;Pise a & Vacele , 2011;
Schlache -Hoenlinge , Pise a & Hoope , 2005).
Type species. Neosch ammeniella mo e i Lé i & Lé i, 1988 ( ype by mono ypy).
Neosch ammeniella inaequalis sp. no .
Figu es 2A–2B,4–5and Table 1
U n:lsid:zoobank.o g:ac :8A516D9B-5351-47AF-8EC2-7EBC44166D35
Holo ype. MNHN-IP-2018-84 (1988-09-26, Go inge Seamoun , beam awl, CP28,
3638′N, 1129.8′W, 605–675 m, Seamoun 1 campaign).
Pa a ype. MNHN-IP-2018-85 (1988-09-26, Go inge Seamoun , beam awl, CP28,
3638′N, 1129.8′W, 605–675 m, Seamoun 1 campaign).
Ca alho e al. (2020), Pee J, DOI 10.7717/pee j.8703 8/87
O he ma e ial. MNHN IP-2018-86 (1988-09-24, Go inge Seamoun , beam awl,
DW21, 3634.9′N, 1128.4′W, 460–480 m, Seamoun 1 campaign).
Compa a i e ma e ial examined. Neosch ammeniella bowe bankii (Johnson, 1863)
(HBOM 003:00592, Madei a), N. bowe bankii (HBOM 003:00810, Madei a), N. pise ai sp.
no . (MNHN-IP-2008-234, Pla o Seamoun ), N. pomponiae sp. no . (MNHN-IP-2008-
233, Hyè es Seamoun ).
Diagnosis. Cup-shaped Neosch ammeniella wi h ounded edges and smoo h su aces;
dic anoclone desmas o ine-like appea ance; i egula dicho iaenes.
Desc ip ion (holo ype MNHN-IP-2018-84). Massi e, fla ened cup-shaped, wi h a
conca e cen e, 73 mm leng h, 29 mm high and 64 mm wide (Fig. 2A); op su ace is
smoo h wi h some oxeas pe o a ing he su ace and se e al small openings e enly
dis ibu ed; walls a e ounded and hick, 14–17 mm wide; bo om su ace is also smoo h,
ull o li le openings dispe sed h oughou he en i e su ace, 31–56 mm in diame e ,
and some oxeas (Fig. 2B); colou is ligh b own in e hanol; he smoo h su aces could
indica e ha hese specimens we e no a ached o any subs a e, and he e o e had a ee
li ing mode (Fig. 2B).
Skele on. Ec osomal skele on composed o smoo h dicho iaenes o a iable shape and
size, along wi h a dense laye o mic oscle es (Figs. 4A and 4B); long-sha ed iaenes o
unde -de eloped dicho iaenes, can also be obse ed (Fig. 4E); choanosomal skele on is
made o an i egula and loose ne wo k o dic anoclone desmas (Figs. 4C and 4D),
spi as e s and me as e s; oxeas can be obse ed c ossing he skele on and p ojec ing he
su ace.
Spicules (holo ype MNHN-IP-2018-84).
1. Dic anoclones, smoo h, i egula , slende , o ine-like appea ance, 354–576–975 ×
12–25–39 mm in size; clones can ha e ew o se e al ube cles, ha a e smoo h o sligh ly
ugose (Figs. 4C–4F);
2. Oxeas, la ge, hin, cu ed, 670–1,144 × 5.2–7.8–13.4 mm in size (Figs. 4A and 4B);
3. Dicho iaenes, ha e a smoo h cladome, ha can be e y i egula , ha ing ounded o
poin ed ips, o clades o unequal size, 118–233–406 mm in diame e (Figs. 5A–5D);
habdome is ei he sho o long, and has a ounded ip, 136–432–1,211 × 9–18–31 mm
in size (Fig. 5A); small b anches o p o ube ances can be obse ed on he habdome, bu
hey a e uncommon (Fig. 5B);
4. Spi as e s, wi h sho and hick a ms, mainly spiny on he a ms, 12.1–18.5–26.6 mmin
size (Figs. 5E–5H);
5. Me as e s, less abundan , co e ed by spines, wi h long and hin a ms, 14.6–31.6–
47.9 mm in size (Fig. 5I).
Dis ibu ion. N. inaequalis sp. no . was ound in he Go inge Seamoun be ween 460 and
675 m dep h.
Ca alho e al. (2020), Pee J, DOI 10.7717/pee j.8703 9/87
Desc ip ion (holo ype MNHN-IP-2008-233). La ge sponge, 54 mm heigh and 81 mm in
diame e , wi h a small pedicel 23 mm wide; i s ex e nal mo phology esembles a bowl;
walls a e abou 11 mm hick; he su aces o he sponge a e ugose, and hispid due o oxeas
p o uding he su ace; openings a e small and e enly sp ead on bo h su aces, 40–87 mm
in diame e ; colou is b own in e hanol (Fig. 2D).
Skele on. Ec osome is composed o a laye o dicho iaenes pe pendicula o he su ace
ha is co e ed by a ious mic oscle es (Figs. 8A and 8B); choanosome composed o a
dense mesh o dic anoclone desmas, oxeas c ossing he choanosome p o uding he
su ace (Fig. 8A), and se e al mic oscle es sp ead h ough he skele on.
Spicules (holo ype MNHN-IP-2008-233).
1. Dic anoclones, compac , i egula and wi h he clones e y ube cula ed,
185–427–666 × 18–39–88 mm in size; ays o desmas a e co e ed by nume ous and
o namen ed ube cles ha ha e a ugose appea ance (Figs. 8C–8E); clones a icula ed
in o complex and in ica e zygoses (Fig. 8F);
2. Oxeas, long, wi h sha p ips, 1455–1643 × 17–18 mm in size (Fig. 8A);
3. Dicho iaenes, wi h a smoo h cladome, 157–274–374 mm in diame e and a long
habdome wi h a blun ip, 239–478–684 × 11–21–37 mm in size (Fig. 9A);
4. Spi as e s, e y abundan , i egula , spiny, wi h sho and hick a ms, 10.7–
18.9–35.8 mm in size (Figs. 9B–9E).
5. Me as e s, less abundan , spiky, wi h long and hin a ms, 16.2–27.6–39.3 mm in size
(Figs. 9F–9I).
E ymology. Named a e D . Shi ley Pomponi om he Ha bou B anch Oceanog aphic
Ins i u e (HBOI) in ecogni ion o he aluable con ibu ions o he knowledge o deep-sea
sponges (including li his ids) o he No h-wes e n A lan ic Ocean and Ca ibbean.
Dis ibu ion. N. pomponiae sp. no . is known om i s ype locali y, Hyè es Seamoun ,
whe e i was collec ed a 480 m dep h.
Rema ks. The genus Neosch ammeniella was e ec ed by Pise a & Lé i (2002b) o
accommoda e Co allis idae wi h smoo h dicho iaenes and wo o h ee ypes o
mic oscle es. This genus is widely dis ibu ed, wi h eco ds spanning he Sou he n Ocean,
SW Pacific, Medi e anean Sea and NEA. Un il now, six species we e desc ibed and
only one, N. bowe bankii (Johnson, 1863), was known o occu in he Medi e anean Sea
(Pise a & Vacele , 2011) and he NEA in he Madei a a chipelago (Ca alho & Pise a,
2019;Johnson, 1863). In he p esen wo k, we desc ibed and illus a e h ee new species o
Neosch ammeniella, ha can mainly be dis inguished by hei habi us, sculp u e o
he desmas, p esence o absence o oxeas, and, shape and size o he dicho iaenes.
The ex e nal mo phology o N. pomponiae sp. no . esembling a bowl, con as s wi h he
cup-shaped o con o ed lamella e masses wi h hick walls in N. bowe bankii, he fla ened
cup-shaped wi h a conca e cen e in N. inaequalis sp. no . and he la ge cup- ec angula
Ca alho e al. (2020), Pee J, DOI 10.7717/pee j.8703 16/87

shape in N. pise ai sp. no . The sculp u e o he desmas is also e y dis inc among all
hese species, while N. bowe bankii has e y ube cula ed dic anoclones di ided in o
smalle and i egula lobes/ ube cles ( edesc ip ion in Pise a & Vacele , 2011),
N. inaequalis sp. no . p esen s a dis inc shape o desmas wi h ine-like appea ance and
ew o se e al ube cles, N. pise ai sp. no . has i egula and compac dic anoclones
A
BF
C
D
E
G
H
I
2 um
10 um
2 um
2 um
2 um
2 um
2 um
2 um
100 um
Figu e 9 Spicules o Neosch ammeniella pomponiae sp. no ., holo ype MNHN-IP-2008-233.
(A) Smoo h dicho iaene, (B)–(E) spi as e s, (F)–(I) me as e .
Full-size

DOI: 10.7717/pee j.8703/fig-9
Ca alho e al. (2020), Pee J, DOI 10.7717/pee j.8703 17/87
Table 1 Compa a i e able o ex e nal mo phology and spicula mic ome ies o all Neosch ammeniella species eco ded in he No h A lan ic Ocean. Spicule
measu emen s (n= 30 unless s a ed o he wise) a e p esen ed as minimum–mean–maximum. Da a compiled om he o iginal desc ip ions, o subsequen e-desc ip ions o
ype ma e ial (ma ked wi h numbe s).
Habi us Size Dic anoclones Dicho iaenes Oxeas Spi as e s Me as e s Locali y
1
N. bowe bankii
Holo ype
BMNH
69.11.60.1
(PZS 1862)
–––Cladome:
319–397 µm
in diame e ;
habdome
487–939 µm
leng h
–20.2–23.7 ×
7.0–11.7 µm
(as spi as e ype I)
28.3–39.3 ×
19.6–32.7 µm
(as spi as e
ype II)
Madei a (dep h
unknown)
2
N.bowe bankii
(Johnson,
1863)
Cup-shaped o
con o na ed lamella e
masses wi h hick
walls; colou whi e
80 × 60 ×
60 mm in size
290–402 µm in
size
Cladome:
176–323 µm;
habdome:
223–513 µm
340–820 ×
1.5–2.5 µm
Sho a ms, 17–24 ×
7.06–11.1 µm in size
(as spi as e ype I)
Long a ms,
26.2–39.2 ×
18.5–23.9 µm
(as spi as e
ype II)
Medi e anean
Sea
(20–22 m)
N. inaequalis sp.
no . (Holo ype
MNHN-2018-
84)
Fla ened cup-shape,
wi h a conca e cen e ;
bo h su aces a e
smoo h; colou ligh
b own
73 × 64 ×
29 mm in
size; walls,
14–17 mm
hick
354–576–975
×12–25–39
µm (n= 12)
Cladome: smoo h,
e y i egula ,
118–233–406 µm;
habdome: long
wi h a ound ip,
136–432–1211 ×
9–18–31 µm
La ge, hin,
cu ed,
670–1144 ×
5.2–7.8–13.4
µm (n=5)
Sho wi h hick a ms,
e y abundan ,
12.1–18.5–26.6 µm
Long and hin
a ms,
14.6–31.6–
47.9 µm
Go inge
Seamoun
(605–675 m
dep h)
N. inaequalis sp.
no . (Pa a ype
MNHN-2018-
85)
Small, ball shaped wi h
a conca e op; bo h
su aces a e smoo h;
colou ligh b own
34 mm
diame e ,
20 mm
heigh
308–431–575
×21–34–49
µm (n= 15)
Cladome:
158–298–463 µm;
habdome:
221–550–1228 ×
13–23–38 µm
La ge, hin,
cu ed
449–1034 ×
5–7–10 µm
(n=8)
10.4–20.3–26.1 µm 15.1–32.7–
47.6 µm
(n= 17)
Go inge
Seamoun
(605–675 m
dep h)
N. pise ai sp.
no .
(Holo ype
MNHN-IP-
2008-234)
La ge cup- ec angula
sponge a ached o
he subs a e by he
en i e lowe base;
bo h su aces smoo h;
colou beige
69 mm in
diame e a
he op, and
43 mm a he
base, 98 mm
heigh ; walls,
11 mm hick
280–428–522
×16–25–37
µm (n=6)
Cladome: smoo h,
153–244–389 µm;
habdome: long
wi h a ound ip,
198–366–535 ×
10–19–33 µm
No p esen Sho wi h hick a ms,
e y abundan ,
14.7–18.7–23.7 µm;
some e y i egula ,
hab-like,
13.5–17.8–23.1 µm
Long and hin
a ms,
18.9–30.7–
41.5 µm
Pla o
Seamoun
(695 m
dep h)
N.pomponiae
sp. no .
(Holo ype
MNHN-IP-
2008-233)
Cup- ounded shape 54 × 81 mm in
size wi h a
small pedicel,
23 mm in
size; walls,
11 mm hick
185–427–666
×18–39–88
µm (n= 13)
Cladome:
157–274–374 µm;
habdome:
239–478–684 ×
11–21–37 µm
(n= 17)
La ge, hick,
1,455–1,643
×17–18 µm
(n=2)
Ve y abundan ,
10.7–18.9–35.8 µm
16.2–27.6–
39.3 µm
Hyè es
Seamoun
(480 m
dep h)
No es:
1
In o ma ion p o ided by P o . A. Pise a, 2019, pe sonal communica ion.
2
Re-desc ip ion in Pise a & Vacele (2011).
‘–’ no in o ma ion/no men ioned.
Ca alho e al. (2020), Pee J, DOI 10.7717/pee j.8703 18/87
ha a e usually smoo h, and N. pomponiae sp. no . has desmas densely co e ed by
nume ous and o namen ed ube cles wi h a ugose appea ance. Finally, N. inaequalis sp.
no . is he only one wi h e y a iable dicho iaenes ei he in size and shape, while N. pise ai
sp. no . does no ha e oxeas, a ype o megascle e p esen in he o he h ee species.
Family Theonellidae Lenden eld, 1903
Genus Discode mia du Bocage, 1869
Synonymy. Collinella Schmid , 1879 (junio synonym); Desmahabana Alcolado & Go e a,
1986 (junio synonym).
Diagnosis. Theonellidae wi h disco iaenes exclusi ely as ec osomal megascle es and
choanosomal e aclone desmas; mic oscle es a e acan hoxeas and acan ho habds.
Defini ion. Polymo phic sponges, om massi e i egula o cup-shaped, b anched o
cylind ical; ec osomal megascle es a e smoo h disco iaenes; choanosomal megascle es a e
e aclone desmas ( egula o i egula ) ha can be smoo h o ube cula ed, and oxeo es o
s ylo es; mic oscle es a e acan hoxeas and acan ho habds (Kelly, 2007;Pise a & Lé i,
2002c;Pise a & Vacele , 2011).
Type species. Dac ylocalyx polydiscus Bowe bank, 1869.
Discode mia ami e a Topsen , 1892
Figu es 2E,10–11 and Table 2
Ma e ial examined. MNHN-IP-2008-204 (1993-01-09, Me eo Seamoun , beam awl,
CP138, 3002′N, 2829′W, 300 m), MNHN-IP-2008-207 (1993-01-10, G ea Me eo
Seamoun , epiben hic d edge, DE140, 3001′N, 2828′W, 308 m), MNHN-IP-2008-213
(1993-01-11, G ea Me eo Seamoun , beam awl, CP156, 2956′N, 2824′W, 320 m),
MNHN-IP-2008-214 (1993-01-10, G ea Me eo Seamoun , beam awl, CP144, 3010′N,
2829′W, 335 m). All om he Seamoun 2 campaign.
Compa a i e ma e ial examined. Discode mia e ucosa Topsen , 1928 (MNHN-IP-
2008-205, A lan is Seamoun ; MNHN-IP-2008-206, Pla o Seamoun ; HBOM 003:00869,
Madei a; HBOM 003:00870, Madei a; HBOM 003:00868, Sel agens; HBOM 003:00640,
Cana y Islands; RMNH6237, Sel agens), D. kellyae sp. no . (holo ype MNHN-IP-2008-
208, Pla o Seamoun ), D. a bo sp. no . (holo ype MNHN-IP-2008-211, G ea Me eo
Seamoun ).
Diagnosis. Small Discode mia, elonga ed o b anching in shape, wi h smoo h e aclone
desmas.
Desc ip ion (MNHN-IP-2008-213). Elonga ed and b anched, small sponge, 15–29 mm
high and 3–10 mm hick (Fig. 2E); su ace is smoo h and anspa en , whe e i is possible
o see he subde mal wa e canals, ha gi es a s ia ed appea ance o he sponge when
obse ed unde a magnifie ; openings o m a small ele a ion on he sponges’su ace;
colou is beige o ligh yellow in e hanol.
Ca alho e al. (2020), Pee J, DOI 10.7717/pee j.8703 19/87
AB
CD
100 um 200 um
100 um 20 um
Figu e 10 Skele on o Discode mia ami e a Topsen , 1892, specimen MNHN-IP-2008-213.
(A) O e iew o choanosomal desmas, (B) e aclone desmas and some disco iaenes, (C) de ail o he
smoo h e aclone desmas wi h ube cles in he zygome, (D) zygosis.
Full-size

DOI: 10.7717/pee j.8703/fig-10
ABC
DEF
100 um 100 um 100 um
100 um
10 um
10 um
Figu e 11 Spicules o Discode mia ami e a Topsen , 1892, MNHN-IP-2008-213. (A)–(C) Lowe iew
o disco iaenes, (D) op iew o disco iaene, (E) acan homic oxeas, (F) acan ho habds.
Full-size

DOI: 10.7717/pee j.8703/fig-11
Ca alho e al. (2020), Pee J, DOI 10.7717/pee j.8703 20/87
Skele on. Ec osome is composed o a laye o o e lapping disco iaenes and abundan
mic oscle es such as acan homic oxeas and acan ho habds, sp ead h ough his pa o
he skele on; choanosomal skele on has e aclone desmas (Fig. 10), smoo h oxeas and
some mic oscle es sp ead h ough he en i e sponge; desmas o m an i egula and
compac ne on he choanosome bu a loose mesh nea he ec osome wi h big spaces
be ween hem; oxeas can be obse ed c ossing he in e io o he skele on.
Spicules (MNHN-IP-2008-213).
1. Te aclone desmas, wi h smoo h ays (Figs. 10A–10C) and ube cula ed zygoses
(Fig. 10D); ube cles a e gene ally smoo h bu in some cases one ube cle may be di ided
in o a ious smalle ube cles; e aclones a e 182–328–470 × 24–32–48 µm in size;
2. Disco iaenes, e y a iable in shape, om ound/o al o i egula and inden ed
cladome; cladome can be fla o sligh ly conca e, 124–160–213 µm diame e ; habdome,
sho and conical, 23–32–40 µm × 8–10–14 µm in size (Figs. 11A–11D).
3. Oxeas, long, smoo h wi h ounded ex emi ies (Fig. 10C); he as majo i y o oxeas
we e b oken, hus measu emen s o hese megascle es a e no p esen ed he e.
4. Acan homic oxeas, sligh ly cu ed wi h poin ed ends, a ely cen o ylo es, 23–28–33 ×
1.0–1.5–1.8 µm in size (Fig. 11E).
5. Acan ho habds, simila o mic oxeas wi h he excep ion hey a e smalle and ha e
ounded ips, 3.9–10.3–13.9 × 1.1–1.4–1.9 µm in size (Fig. 11F).
Dis ibu ion. Specimens we e collec ed a he G ea Me eo Seamoun be ween 300 and
335 m dep h.
Rema ks. D. ami e a was desc ibed by Topsen (1892) om ma e ial collec ed in he
Azo es (318 m dep h), and la e e-collec ed in he same a chipelago a 98 m dep h
(Topsen , 1904). So a , hese we e he only eco ds in he No h A lan ic. He e we disco e
o he fi s ime he p esence o his species in he G ea Me eo seamoun (be ween
300 and 335 m dep h). The specimens analysed in his wo k ha e a simila ex e nal
mo phology compa ed o he ones desc ibed by Topsen (i.e., small, elonga ed o
b anching sponge wi h finge -like ex ensions), and simila spicule composi ion. Howe e ,
he spicules’sizes a e in gene al smalle om hose p esen ed in he o iginal desc ip ion
(Table 2). Disco iaenes ha e a smalle cladome, 124–213 µm in he analysed ma e ial
e sus he 300 µm in diame e in he o iginal desc ip ion; acan homic oxeas (22.8–32.6 µm
s 40–45 µm long) and acan ho habds a e also smalle (3.9–13.9 µm s 20–25 µm long),
bu see Discussion o mo e de ails on hese di e ences.
Discode mia c . ami e a Topsen , 1892
Ma e ial. MNHN-IP-2008-210 (1993-02-02, A lan is Seamoun , epiben hic Wa én
d edge, DW258, 3400′N, 3012′W, 420 m, Seamoun 2 campaign).
Compa a i e ma e ial examined. D. ami e a (MNHN-IP-2008-204, G ea Me eo
Seamoun ; MNHN-IP-2008-207, G ea Me eo Seamoun ; MNHN-IP-2008-213, G ea
Ca alho e al. (2020), Pee J, DOI 10.7717/pee j.8703 21/87

Me eo Seamoun ; MNHN-IP-2008-214, G ea Me eo Seamoun ), Discode mia e ucosa
Topsen , 1928 (MNHN-IP-2008-205, A lan is Seamoun ; MNHN-IP-2008-206, Pla o
Seamoun ; HBOM 003:00869, Madei a; HBOM 003:00870, Madei a; HBOM 003:00868,
Sel agens; HBOM 003:00640, Cana y Islands; RMNH6237, Sel agens), D. kellyae sp. no .
(holo ype MNHN-IP-2008-208, Pla o Seamoun ), D. a bo sp. no . (holo ype MNHN-IP-
2008-211, G ea Me eo Seamoun ).
Desc ip ion (MNHN-IP-2008-210). Small agmen , 20 × 10 mm in size, o elonga ed
shape, wi h a smoo h su ace; subde mal wa e canals a e isible, gi ing a s ia ed
appea ance o he sponge; colou is beige in e hanol.
Skele on. Ec osomal skele on is o med by a laye o o e lapped disco iaenes, and se e al
mic oscle es sp ead h ough he su ace; choanosome is o med by i egula e aclone
desmas, oxeas c ossing he in e io o he sponge and nume ous mic oscle es sp ead
h ough he in e io o he sponge.
Spicules (MNHN-IP-2008-210).
1. Te aclone desmas, i egula , wi h smoo h clones and e y ube cula ed on he
ex emi ies, 400–455–534 × 30–51–82 µm in size; ube cles a e smoo h;
2. Disco iaenes, cladome a ies om o al o inden ed in shape, usually fla , 195–328–
560 µm in diame e ; habdome is sho , conical, wi h a blun ip, 20–42–68 ×
9.5–20.3–37.9 µm in size;
3. Oxeas, a e p esen , bu all o hem we e b oken;
4. Acan homic oxeas, e y abundan , spinous, wi h sha p ips, 24.6–39.0–59.8 ×
1.8–3.3–5.4 µm in size;
5. Acan ho habds, small, abundan , spinous, wi h ounded ex emi ies, 15.2–20.2–24.2 ×
2.1–2.9–4.4 µm.
Dis ibu ion. This specimen was collec ed in he A lan is Seamoun a 420 m dep h.
Rema ks. Al hough he ex e nal mo phology, ype o spicules and desma o namen a ion
a e in ag eemen wi h he desc ip ion o D. ami e a, he spicules sizes o his specimen a e
significan ly la ge when compa ed o he ones ound in he G ea Me eo (Table 2).
Fo his eason, we conside his species as D. c . ami e a.
Discode mia e ucosa Topsen , 1928
Figu es 2F,12–13 and Table 2
Ma e ial examined. MNHN-IP-2008-205 (1993-02-02, A lan is Seamoun , beam awl,
CP257, 3404′N, 3015′W, 338 m), MNHN-IP-2008-206 (1993-02-01, Pla o Seamoun ,
epiben hic Wa én d edge, DW246, 3314′N, 2936′W, 520 m). All om Seamoun 2
campaign.
Compa a i e ma e ial examined. D. ami e a (MNHN-IP-2008-204, G ea Me eo
Seamoun ; MNHN-IP-2008-207, G ea Me eo Seamoun ; MNHN-IP-2008-213, G ea
Ca alho e al. (2020), Pee J, DOI 10.7717/pee j.8703 22/87
200 um 100 um
200 um 30 um
AB
CD
Figu e 12 Skele on o Discode mia e ucosa Topsen , 1928, specimen MNHN-IP-2008-205.
(A) O e iew o e aclone desmas, (B) and (C) i egula and compac ne o e aclone desmas,
(D) de ail o he s ongly ube cula ed zygosis. Full-size

DOI: 10.7717/pee j.8703/fig-12
A
BC
D
E
F
20 um
20 um
20 um
2 um
30 um
10 um
Figu e 13 Spicules o Discode mia e ucosa Topsen , 1928, specimen MNHN-IP-2008-205.
(A)–(D) Uppe and lowe iew o disco iaenes, (E) acan homic oxeas, (F) acan ho habds.
Full-size

DOI: 10.7717/pee j.8703/fig-13
Ca alho e al. (2020), Pee J, DOI 10.7717/pee j.8703 23/87
Me eo Seamoun ; MNHN-IP-2008-214, G ea Me eo Seamoun ), D. kellyae sp. no .
(holo ype MNHN-IP-2008-208, Pla o Seamoun ), D. a bo sp. no . (holo ype MNHN-IP-
2008-211, G ea Me eo Seamoun ).
Diagnosis. Cup-shaped o sphe ical sponges wi h nume ous wa s/p o ube ances, and
ex emely ube cula ed e aclone desmas (emended a e Topsen , 1928).
Desc ip ion (MNHN-IP-2008-205). Sphe ical polymo phic sponge wi h se e al ound
p o ube ances, 15–20 mm high and 12–13 mm wide, wi h a ough su ace (Fig. 2F); po es
canno be seen wi h naked eye; colou a ies om whi ish o ligh b own in e hanol.
Skele on. Ec osome composed o a compac laye o disco iaenes, usually o e lapping
each o he , nume ous mic oscle es (acan homic oxeas and acan ho habds) sp ead
h ough he su ace, and oxeas pe o a ing he sponges’su ace; occasionally, bundles o
oxeas can be obse ed; choanosome wi h s ongly ube cula ed and compac e aclone
desmas (Fig. 12), o ming an i egula ne wi h dispe sed mic oscle es in he in e io o he
sponge.
Spicules (MNHN-IP-2008-205).
1. Te aclone desmas, la ge, obus , mos ly wi h ube cles sp ead h ough he en i e
clone, al hough some pa s can be smoo h, 106–170–278 × 19–34–46 µm in size
(Figs. 12A–12C); zygoses e y obus and ex emely ube cula e (Fig. 12D);
2. Disco iaenes, i egula in shape, om ound o o al, o en inden ed (Figs. 13A–13D);
cladome smoo h, sligh ly conca e, 102–153–222 µm in diame e ; habdome is sho
wi h a conical shape, 15–25–47 × 5–8–13 µm (Fig. 13D);
3. Oxeas, long, smoo h wi h ounded ends; leng h no p esen ed he e because hey we e all
b oken due o hei la ge size.
4. Acan homic oxeas, spinous, sligh ly cu ed wi h poin ed ends, 22.8–35.2–53.5 ×
1.3–2.2–3.9 µm (Fig. 13E).
5. Acan ho habds, cylind ical, spinous, wi h blun ips, 7.5–12.9–19.0 × 1.2–1.6–3.0 µm in
size (Fig. 13F).
Dis ibu ion. Specimens o D. e ucosa we e ound in A lan is and Pla o Seamoun s
be ween 338 and 580 m dep h.
Rema ks. Discode mia e ucosa was fi s ound in he Cana y Islands and desc ibed by
Topsen (1928). The species di e s om he D. ami e a on he habi us and sculp u e o
desmas. D. e ucosa has a cup o sphe ical shape wi h se e al ounded p o ube ances/wa s
and s ongly ube cula ed e aclones. On he o he hand, D. ami e a has an elonga ed o
b anching shape and smoo h e aclone desmas only ube cula ed in he ex emi ies.
The specimens analysed in his s udy o e all ma ch he desc ip ion o D. e ucosa,apa
om wo di e ences: (1) he disco iaenes a e much smalle and (2) he mic oscle es
p esen a wide size ange when compa ed o he o iginal desc ip ion (see Table 2).
Ca alho e al. (2020), Pee J, DOI 10.7717/pee j.8703 24/87
Discode mia a bo sp. no .
Figu es 2G,14–15 and Table 2
U n:lsid:zoobank.o g:ac :7A732A92-8D8B-4D73-97B1-CD53E9494121
Holo ype. MHNH-IP-2008-211 (1993-01-11, G ea Me eo Seamoun , beam awl,
DW159, 2944′N, 2820′W, 330 m, Seamoun 2 campaign).
Compa a i e ma e ial examined. D. ami e a (MNHN-IP-2008-204, G ea Me eo
Seamoun ; MNHN-IP-2008-207, G ea Me eo Seamoun ; MNHN-IP-2008-213, G ea
Me eo Seamoun ; MNHN-IP-2008-214, G ea Me eo Seamoun ), D. e ucosa (MNHN-
IP-2008-205, A lan is Seamoun ; MNHN-IP-2008-206, Pla o Seamoun ; HBOM
200 um 30 um
200 um 200 um
20 um 20 um
AB
CD
EF
Figu e 14 Su ace and skele on o Discode mia a bo sp. no ., holo ype MNHN-IP-2008-211.
(A) O e iew o he su ace, (B) de ail o he su ace showing he o e lapped disco iaenes, (C) o e -
iew o choanosomal e aclone desmas, (D) de ail o e aclone desmas, (E) complex zygoses be ween
se e al desmas, (F) de ail o he desmas o namen a ion, showing smoo h ube cles.
Full-size

DOI: 10.7717/pee j.8703/fig-14
Ca alho e al. (2020), Pee J, DOI 10.7717/pee j.8703 25/87
Table 2 (con inued).
Habi us Size Te aclones Disco iaenes Oxeas Acan homic oxeas Acan ho habds Locali y
D. a bo sp. no .
(Holo ype
MNHN-IP-
2008-211)
Massi e discode mia o
ee like appea ance,
wi h a long s em and
h ee b anches; su ace
is smoo h; colou is
beige in e hanol
Full sponge
leng h is 58
mm; s em
is 15 mm
high and
7.5–12 mm
wide and
b anches
a e
13–28 mm
long
Usually wi h he
a ms
ube cula ed,
bu can be
smoo h; e y
s ong
zygoses;181–
392–567 ×
15–36–56 µm in
size
Cladome:
148–256–396
µm in diame e ;
habdome:
34–53–71 ×
15–21–24 µm
(n=9)
No p esen Sligh ly cu ed,
spinous, wi h
sha p ips,
24.1–35.1–50.1 ×
1.4–2.3–3.5 µm
Co e ed by nume ous
spines, wi h unequal
ips (blun o sha p)
6.7–16.1–25.9 ×
1.1–2.2–4.3 µm
G ea
Me eo
Seamoun
(330 m
dep h)
D. kellyae sp.
no .
(Holo ype
MNHN-IP-
2008-208)
Massi e sponge,
polymo phic o bulb
appea ance, wi h la ge
p o ube ances o
ound shape; su ace is
i egula wi h a
c umble/ ugose
appea ance; colou is
beige o ligh b own
53 mm high
and 31 mm
wide
La ge, compac ,
hick,
112–338–589 ×
20–42–76 µm
(n= 20)
Cladome: e y
a iable in
shape and size,
121–289–425
µm in diame e ;
habdome
36–81–142 ×
13–31–44 µm
S ongyles, one
ip ounded
and he o he
one sha p,
418–444 ×
6.0–7.9 µm in
size (n=2)
S aigh o cu ed,
wi h sha p ips,
spinous,
16.7–43.2–66.5 ×
1.5–2.5–3.7 µm
Spinous, wi h blun
ips, 5.3–13.3–24.9 ×
1.2–2.1–3.7 µm
Pla o
Seamoun
(580 m
dep h)
No es:
1
Redesc ip ion in Pise a & Lé i (2002c).
2
Pise a & Lé i (2002d).
3
This desc ip ion was aken om Pise a & Pomponi (2015) since he species was poo ly desc ibed in he o iginal and no in o ma ion on he spicules measu emen s was gi en.
4
Topsen (1892).
5
Topsen (1928).
6
Pise a & Vacele (2011).
7
Van Soes , Mees e s & Becking (2014).
Ca alho e al. (2020), Pee J, DOI 10.7717/pee j.8703 32/87

ec osome; he ays o he e aclones a icula e in o a complex zygosis; se e al mic oscle es
and some s ongyles a e sp ead loosely in he choanosome.
Spicules (holo ype MNHN-IP-2008-208).
1. Te aclone desmas, compac , i egula , wi h smoo h and hick clones, 112–338–589 ×
20–42–76 µm in size (Figs. 16C and 16D); he e mina ion o he clones has se e al
ube cles, esul ing in e y complex and la ge zygoses (Figs. 16D–16F); ube cles o he
clones a e smoo h (Fig. 16F).
2. Disco iaenes, i egula , wi h di e se shapes and sizes; cladomes a y om o al o
inden ed discs, and hey a e ei he fla o conca e, 121–289–425 µm in diame e
(Figs. 16A,16B and 17A–17G); habdome is also e y a iable in size, 36–78–119 ×
13–30–44 µm, wi h a blun o sha p ip.
3. S ongyles, wi h one o he ips ounded and he o he one sha p, some imes esembling
a c oche needle, 418–444 × 6.0–7.9 µm in size (Figs. 17H and 17I);
4. Acan homic oxeas, e y abundan , long, s aigh o cu ed, co e ed by nume ous
spines, wi h sha p ips, 16.7–43.2–66.5 × 1.5–2.5–3.7 µm in size (Figs. 17J and 17K);
5. Acan ho habds, e y abundan , wi h blun ips, co e ed by nume ous spines, e y
a iable in size, 5.3–13.3–24.9 × 1.2–2.1–3.7 µm (Figs. 17L and 17M).
E ymology. Named a e D . Michelle Kelly om he Na ional Ins i u e o Wa e and
A mosphe ic Resea ch (NIWA) in ecogni ion o he wo k on axonomy and sys ema ics
o Po i e a, pa icula ly on li his id demosponges o New Zealand.
Dis ibu ion. D. kellyae sp. no . is only known om i s ype locali y, he Pla o Seamoun
a 580 m dep h.
Rema ks. The iden ifica ion o species belonging o he genus Discode mia is pa icula ly
challenging due o he ew and e y a iable mo phological cha ac e s used o he
dis inc ion o species (Pise a & Vacele , 2011). Mo eo e , o some species we a e limi ed o
he o iginal desc ip ions whe e de ailed in o ma ion o skele al composi ion and spicule
sizes, o images a e lacking.
In he No h A lan ic and Medi e anean Sea, a o al o nine species ha e been desc ibed,
including he wo desc ibed species in his s udy (Table 2). Despi e he high plas ici y o
mo phological cha ac e s, he main di e ences be ween species a e (1) habi us, (2) he
sculp u e and size o he desmas, (3) size and shape o he disco iaenes, and (4) size and
shape o he mic oscle es. We p opose D. kellyae sp. no . as a new species based on (1) he
habi us o his sponge: he polymo phic sponge o bulb appea ance con as s wi h he
massi ely enc us ing shape o D. adhae ens, he sphe ical o i egula masses in
D. polymo pha, he cup-shaped wi h nume ous wa s/p o ube ances in D. e ucosa, he
elonga ed wi h se e al finge -like ex ensions in D. ami e a, he ee-like shape o D. a bo ,
he clus e o knobby finge s in D. dissolu a and he i egula mush oom shape o
D. polydiscus; (2) e aclones o D. kellyae sp. no . ha e simila o namen a ion o he ones
ound in D. ami e a ( e aclones wi h smoo h clones ha a e ube cula ed in he zygomes),
Ca alho e al. (2020), Pee J, DOI 10.7717/pee j.8703 33/87
howe e , hey a e mo e compac and hicke (24–32–48 µm s 20–42–76 µm) esembling he
ones p esen in D. e ucosa; he o he species ha e slende and smoo h desmas wi hou
s ong/complex zygoses; (3) he in aspecific size ange o disco iaenes is usually wide, and
simila be ween he di e en species, bu in D. kellyae sp. no . he size ange o he cladomes
is e y la ge, 121–425 µm, and his can only be obse ed in D. e ucosa (200–560 µm) and
D. a bo sp. no . (148–396 µm); besides ha , he shape o he habdome is also a iable in
D. kellyae sp. no ., whe e he ips o he habdomes can be blun o sha p; (4) he size o he
acan homic oxeas in D. kellyae sp. no . is la ge (16.7–43.2–66.5 µm) compa ed o he o he
species, excep when compa ed o D. dissolu a (41.6–68.0 µm; howe e , hese alues we e
aken om Pise a & Pomponi, 2015 whe e he au ho s p esen ed a de ailed desc ip ion o he
species, since in he o iginal desc ip ion, he species was poo ly desc ibed and no
measu emen s we e gi en); (5) D. kellyae sp. no ., along wi h D. a bo sp. no ., a e he only
species wi h a wide acan ho habds size ange (5.3–13.3–24.9 µm and 6.7–16.1–25.9 µm,
espec i ely) while he o he species ha e a conside ably na owe ange (Table 2).
The species D. insc ip a (Schmid , 1879) was no included he e o compa ison because
he ype ma e ial was deciduous and he species is he e o e conside ed ince ae sedis
(Pise a & Lé i, 2002d).
Family Macand ewiidae Sch ammen, 1924
Genus Macand ewia G ay, 1859
Diagnosis. Macand ewiidae wi h phyllo iaenes/disco iaenes as ec osomal megascle es;
choanosmal megascle es a e oxeas and desmas wi h a iaenose c epsis; mic oscle es a e
mic oxeas (emended a e Pise a & Lé i, 2002e).
Defini ion. Polymo phic Macand ewiidae; ec osomal spicules a e den a e phyllo iaenes
and/o disco iaenes; desmas a e smoo h wi h a iaenose ( a ely monaxial) c epsis, and a
e minal zygosis; oxeas a e smoo h; mic oscle es a e mic oxeas (emended a e Pise a &
Lé i, 2002b).
Type species. Macand ewia azo ica G ay, 1859 ( ype by mono ypy).
Macand ewia c . azo ica G ay, 1859
Figu es 2I,18–19 and Table 3
Ma e ial. MNHN-IP-2008-217 (1993-02-03, A lan is Seamoun , beam awl, DW263,
3426′N, 3032′W, 610 m), MNHN-IP-2008-220 (1993-02-03, A lan is Seamoun ,
epiben hic Wa én d edge, DW258, 3400′N, 3012′W, 1,000 m), MNHN-IP-2008-225
(1993-02-06, Ty o Seamoun , epiben hic Wa én d edge, DW277, 3400′N, 2821′W, 1,000
m), MNHN-IP-2008-226 (1993-01, no da a abou s a ion, 500 m), MNHN-IP-2008-229
(1993-01-06, G an Cana ia, epiben hic Wa én d edge, DW129, 2808′N, 1552′W, 480
m), MNHN-IP-2008-249a (1993-01-06, Hyè es Seamoun , epiben hic Wa én d edge,
DW202, 3116′N, 2843′W, 640 m). All om Seamoun 2 campaign.
Compa a i e ma e ial examined. M. azo ica (holo ype BMNH 1851.7.28.16, S. Miguel
island, Azo es; HBOM 003:00784, Sel agens), M. obus a (MNHN-IP-2008-216, Hyè es
Ca alho e al. (2020), Pee J, DOI 10.7717/pee j.8703 34/87
Seamoun ; MNHN-IP-2008-224, Hyè es Seamoun ), M. schus e ae sp. no . (holo ype
MNHN-IP-2018-87 and pa a ype MNHN-IP-2018-90, Go inge Seamoun ), M. minima
sp. no . (MNHN-IP-2008-222, G ea Me eo Seamoun ).
Desc ip ion (MNHN-IP-2008-220). Polymo phic sponges a ached o he subs a e by a
hick pedicel/s em, 67 × 50 mm in size; lamellas a e hin, ounded and undula e, 3–5mm
hick (Fig. 2I); inne su ace ( op) has openings isible o he naked eye, a ound 224 µm
in size (Fig. 18A); ou e su ace is smoo h wi h se e al li le openings sp ead andomly
h ough he en i e sponge, 40–83 µm in size (Fig. 18B); colou is beige o ligh b own in
e hanol.
AB
CD
EF
200 um 200 um
100 um 200 um
200 um 20 um
Figu e 18 Su ace and skele on o Macand ewia c . azo ica G ay, 1859, specimen MNHN-IP-2008-
220. (A) Uppe /inne su ace wi h la ge openings, (B) lowe /ou e su ace wi h se e al small openings,
(C) di ision be ween ec osome and choanosome: op o he image showing he ec osome o med by
phyllo iaenes and mic oxeas, and he bo om showing he desmas, (D) choanosomal desmas,
(E) choanosomal desmas esembling e aclones, (F) de ail o he sculp u e o desmas and zygoses.
Full-size

DOI: 10.7717/pee j.8703/fig-18
Ca alho e al. (2020), Pee J, DOI 10.7717/pee j.8703 35/87
Skele on. Ec osome o med by a laye o o e lapped phyllo iaenes co e ed by nume ous
mic oxeas (Figs. 18A–18C); small openings a e su ounded by mic oxeas (Fig. 18A)
whe eas la ge openings a e delimi ed by bo h phyllo iaenes and mic oxeas (Fig. 18B);
choanosomal skele on o med by a egula and solid ne wo k o desmas wi h a iaenose
c epsis, esembling e aclone desmas (Figs. 18D and 18E), some oxeas and mic oxeas a e
sp ead in he in e io o he sponge.
Spicules (MNHN-IP-2008-220).
1. Desmas, wi h a iaenose c epsis, compac , o ming a egula mesh, esembling
e aclones; ays a e smoo h wi h b anches, especially on he e mina ion o he clone,
measu ing 212–281–343 × 16–34–51 µm in size; b anches ha e blun ends and hei size
is e y a iable, 34–54–74 × 5.9–8.3–11.5 µm in size (Figs. 18D–18F);
2. Phyllo iaenes, wi h pa icula ly incised cladome wi h 194–267–333 µm in diame e ,
and a sho conical-shaped habdome, 62–99–129 × 11.6–14.4–17.8 µm in size;
cladomes a e e y a iable, om a simple (Fig. 19A) o a e y complex and incised shape
(Figs. 19B and 19C);
3. Oxeas, smoo h, sligh ly cu ed wi h poin ed ends, 215–246–301 × 6.8–7.8–9.1 µm in
size (Fig. 19D);
4. Mic oxeas, smoo h, usi o m wi h blun ips, sligh ly cu ed, e y abundan ,
33.3–55.0–83.6 × 2.5–3.9–5.1 µm in size (Fig. 19E).
AB
C
D
E
20 um
20 um
20 um
mu
01
mu
0
1
mu
0
0
1
Figu e 19 Spicules o Macand ewia c . azo ica G ay, 1859, specimen MNHN-IP-2008-220.
(A)–(C) Phyllo iaenes wi h a e y incised cladome, (D) oxeas, (E) mic oxeas.
Full-size

DOI: 10.7717/pee j.8703/fig-19
Ca alho e al. (2020), Pee J, DOI 10.7717/pee j.8703 36/87
Dis ibu ion. The specimens we e ound on he A lan is Seamoun be ween 420 and
610 m dep h, and one specimen was collec ed in G an Cana ia a 480 m dep h.
Rema ks.Pise a & Lé i (2002d) e-desc ibed and illus a ed he holo ype o M. azo ica,a
specimen collec ed in he Azo es a chipelago. Since we also had access o he holo ype o
M. azo ica we ha e made new measu emen s o he spicules, in o de o fill he gaps o
some spicule’s measu emen s missing in he edesc ip ion. The compa ison o he
holo ype o M. azo ica wi h he specimens collec ed du ing he campaigns Seamoun 1 and
2, lead us o conside hese specimens as M. c . azo ica. Al hough e y simila in he
habi us hey di e om he holo ype in wo ea u es: (1) desmas a e conside ably mo e
obus and hicke , esembling e aclones (MNHN-IP-2008-220: 16–34–51 µm wid h s
holo ype BMNH 1851.7.28.16: 8.5–19.0–30.8 µm wid h), o ming compac ne wo k,
while in he edesc ip ion o he holo ype, he desmas ha e a “ a iable mo phology”
esembling e aclones o hizoclones, wi h s ongly b anched clones a he ip, o ming a
complex and loose a icula ion (Pise a & Lé i, 2002e); (2) he size o he cladome o he
phyllo iaenes (MNHN-IP-2008-220: 194–267–333 µm in diame e s holo ype BMNH
1851.7.28.16: 297–363–456 µm in diame e ) and oxeas (MNHN-IP-2008-220:
215–246–301 µm leng h s holo ype BMNH 1851.7.28.16: 532–652–780 µm leng h) is
conside ably smalle (Table 3).
Nine een la ge specimens we e ound in he same s a ion in he Hyè es seamoun
(s a ion DW202), sugges ing ha he species may be o ming a sponge g ound in his a ea
o he seamoun .
Macand ewia obus a Topsen , 1904
Figu es 2J,20–21 and Table 3
Ma e ial. MNHN-IP-2008-216, wo specimens (1993-01-16, Hyè es Seamoun , epiben hic
Wa én d edge, DW184, 3124′N, 2852′W, 705 m), MNHN-IP-2008-224 wo specimens
(1993-01-16, Hyè es Seamoun , epiben hic Wa én d edge, DW184, 3124′N, 2852′W,
705 m). All om Seamoun 2 campaign.
Compa a i e ma e ial examined. M. azo ica (holo ype BMNH 1851.7.28.16, S. Miguel
island, Azo es; HBOM 003:00784, Sel agens), M. c . azo ica (MNHN-IP-2008-217,
A lan is Seamoun ; MNHN-IP-2008-220, A lan is Seamoun ; MNHN-IP-2008-225, Ty o
Seamoun ; MNHN-IP-2008-226, no da a; MNHN-IP-2008-229, G an Cana ia; MNHN-
IP-2008-249a, Hyè es Seamoun ), M. schus e ae sp. no . (holo ype MNHN-IP-2018-87
and pa a ype MNHN-IP-2018-90, Go inge Seamoun ), M. minima sp. no . (MNHN-IP-
2008-222, G ea Me eo Seamoun ).
Diagnosis. Small fici o m o globula shape Macand ewia wi h a fla ened op and a sho
and hick pedicel.
Desc ip ion (MNHN-IP-2008-216). Small sponges wi h a fici o m o globula shape,
18–20 × 14–22 mm in size, a ached o he subs a e by a sho and hick pedicel (8 mm in
heigh and 16 mm wid h) (Fig. 2J); op o he sponge is fla ened, smoo h, whe e openings
Ca alho e al. (2020), Pee J, DOI 10.7717/pee j.8703 37/87

can be obse ed in small clus e s leading o wa e canals gi ing a s ia ed appea ance
o he sponge; openings and he subde mal wa e canals isible o he naked eye; la e al
walls o he sponge a e smoo h wi h small openings sp ead e enly h ough his su ace;
in some indi iduals, he op o uppe su ace has a sligh dep ession; colou a ies om
beige o ligh b own in alcohol.
Skele on. Ec osome is composed o a laye o o e lapped phyllo iaenes and nume ous
mic oxeas; hese mic oxeas su ound he openings adially; choanosomal skele on o med by
desmas, oxeas and dispe sed mic oxeas; desmas o m an i egula and e y dense mesh (Fig. 20).
Spicules (MNHN-IP-2008-216).
1. Desmas, wi h a iaenose c epsis, compac , obus , wi h smoo h clones ha a e e y
b anched, 248–362 µm in size and 17–22–31 µm hick (Figs. 20A and 20B); clones ha e
se e al sho (18–41–75 µm), hick (7–10–12 µm) and blun b anches (Figs. 20D and 20E);
he zygosis, ha can be o med by nume ous clones, is s ong and complex (Fig. 20D).
2. Phyllo iaenes, e y a iable in shape, wi h a cladome pa icula ly inden ed on he
edges, 15–228–309 µm in diame e , wi h a conical habdome 46–91–141 × 13–19–
25 µm in size (Figs. 21A–21D);
3. Oxeas, smoo h wi h ounded ips, 203–329 × 7.2–8.2 µm hick (Fig. 21F).
4. Mic oxeas, smoo h, wi h ounded ex emi ies, sligh ly cu ed, 34.6–57.4–79.2 ×
3.1–4.7–6.9 µm wide (Fig. 21E).
AB
CD
200 um
20 um
100 um
20 um
Figu e 20 Skele on o Macand ewia obus a Topsen , 1904, specimen MNHN-IP-2008-216.
(A) O e iew o choanosomal desmas, (B) desmas, (C) zygoses, (D) sculp u e o he desmas.
Full-size

DOI: 10.7717/pee j.8703/fig-20
Ca alho e al. (2020), Pee J, DOI 10.7717/pee j.8703 38/87
Dis ibu ion. These specimens we e ound on Hyè es seamoun a 705 m dep h.
Rema ks. In he specimens he e examined, phyllo iaenes (165–230 µm s 154–309;
Table 3) and oxeas (330–400 s 203–309; Table 3) a e smalle when compa ed o p e ious
eco ds o he species (Topsen , 1904). Howe e , M. obus a has a e y dis inc habi us in
ela ion o he o he Macand ewia desc ibed o he No h A lan ic Ocean (Table 3).
I s fici o m o globula shape, wi h a sho and hick pedicel, con as s wi h he cya hi o m
o flabella e shape wi h undula ing ounded ma gins in M. azo ica, he enc us ing wi h
s anding unks o M. amosa, he olia e wi h hick lamellas in M. schus e ae sp. no .,
o he globula shape wi h a small pedicel as in M. minima sp. no . (desc ip ions o he
la e wo below). Di e ences in spicule sizes we e obse ed in ano he species analysed
in his wo k as well as in o he s udies (see ‘Spicules dimensions’sec ion in he Discussion
o u he in o ma ion ega ding his opic).
Two specimens om he Seamoun 2 collec ion could no be confiden ly iden ified
down o species le el (MNHN-IP-2008-228 and MNHN-IP-2018-94). They a e e y small
agmen s, seemingly enc us ing, and mos likely i is a Macand ewia a an ea ly s age o
AB
C
D
F
E
20 um
20 um
20 um
20 um
mu 00
1
mu
0
1
Figu e 21 Spicules o Macand ewia obus a Topsen , 1904, specimen MNHN-IP-2008-216.
(A)–(D) Phyllo iaenes, (E) mic oxeas, (F) oxea. Full-size
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DOI: 10.7717/pee j.8703/fig-21
Ca alho e al. (2020), Pee J, DOI 10.7717/pee j.8703 39/87
de elopmen . The spicules we e measu ed and hey all wi hin he size ange ound in
M. obus a.
Macand ewia schus e ae sp. no .
Figu es 2K,22–23 and Table 3
u n:lsid:zoobank.o g:ac :2BA2C1EF-8FAB-4C91-89CB-DCB59DDA61EB
Holo ype. MNHN-IP-2018-87 (1988-09-26, Go inge Seamoun , beam awl, CP28,
3628′N, 1129’W, 605–675 m, Seamoun 1 campaign).
Pa a ype. MNHN-IP-2018-88 (1988-09-26, Go inge Seamoun , beam awl, CP28,
3628’N, 1129′W, 605–675 m, Seamoun 1 campaign).
O he ma e ial. MNHN-IP-2018-90, six specimens (1988-09-26, Go inge Seamoun ,
beam awl, CP28, 3628′N, 1129′W, 605–675 m, Seamoun 1 campaign),
MNHN-IP-2018-89 (1988-09-26, Go inge Seamoun , beam awl, CP28, 3628′N, 1129′
W, 605–675 m, Seamoun 1 campaign), MNHN-IP-2018-91 (1988-09-26, Go inge
Seamoun , beam awl, CP28, 3628′N, 1129′W, 605–675 m, Seamoun 1 campaign),
MNHN-IP-2008-219 (1993-02-06, Ty o Seamoun , epiben hic Wa én d edge, DW279,
3356′N, 2824′W, 805 m, Seamoun 2 campaign), MNHN-IP-2008-230 (1993-02-01,
Pla o Seamoun , epiben hic Wa én d edge, DW246, 3314′N, 2936′W, 520 m,
Seamoun 2 campaign).
Compa a i e ma e ial examined. M. azo ica (holo ype BMNH 1851.7.28.16, S. Miguel
island, Azo es; HBOM 003:00784, Sel agens), M. c . azo ica (MNHN-IP-2008-217,
A lan is Seamoun ; MNHN-IP-2008-220, A lan is Seamoun ; MNHN-IP-2008-225, Ty o
Seamoun ; MNHN-IP-2008-226, no da a; MNHN-IP-2008-229, G an Cana ia; MNHN-
IP-2008-249a, Hyè es Seamoun ), M. obus a (MNHN-IP-2008-216, Hyé es Seamoun ;
MNHN-IP-2008-224, Hyè es Seamoun ), M. minima sp. no . MNHN-IP-2008-222 (G ea
Me eo Seamoun ).
Diagnosis. Folia e o ase shaped Macand ewia wi h hick, i egula and undula ed
lamellas, wi h a small pedicel.
Desc ip ion (holo ype MNHN-IP-2018-87). Massi e, olia e o ase shape wi h undula e
lamellas, 94 mm high and 142 mm wide a he op and 45 mm wide a he base,
usually a ached o he subs a e by a la ge pedicel; lamellas a e gene ally i egula and
con o ed; walls a e hick, 7–10 mm (Fig. 2K); in e io su ace wi h openings sligh ly
ele a ed and e enly dis ibu ed, 278–378 µm in diame e (Fig. 22A); subde mal wa e
canals a e isible on he inne su ace; ex e nal su ace is smoo h and co e ed by small
openings, 29–98 µm in size (Fig. 22B); colou ligh b own o whi e in alcohol.
Skele on. Ec osome has phyllo iaenes ha a e co e ed by nume ous mic oxeas,
su ounding he openings adially (Figs. 22A and 22B); choanosome has desmas, oxeas
and dispe sed mic oxeas; desmas a e compac , i egula and c ea e a dense ne wo k
(Figs. 22C and 22D).
Ca alho e al. (2020), Pee J, DOI 10.7717/pee j.8703 40/87
Spicules (holo ype MNHN-IP-2018-87).
1. Desmas, wi h a iaenose c epsis, smoo h, i egula , o ming an in ica e and complex
ne , 301–386–463 × 10.2–19.9–39.2 µm in size (Figs. 22C–22E); clones ha e he
e mina ions spli ing in se e al b anches ha a e usually sho , and blun , 17–37–78 ×
5–9–15 µm in size; zygoses is complex and solid (Fig. 22F).
2. Phyllo iaenes, cladome i is pa icula ly incised on he edges, 177–304–420 µm in
diame e ; sho habdome 67–119–178 × 13–21–26 µm in size (Figs. 23A–23E).
3. Oxeas, smoo h wi h ounded ips, 263–437–620 × 8.1–12.4–16.0 µm in size (Fig. 23F).
4. Mic oxeas, smoo h, wi h ound edges, 43.8–67.9–95.2 × 2.5–4.3–7.7 µm size (Fig. 23G).
BA
CD
EF
100 um 100 um
200 um 200 um
20 um 20 um
Figu e 22 Su ace and skele on o Macand ewia schus e ae sp. no ., holo ype MNHN-IP-2018-87.
(A) In e nal su ace wi h la ge openings, (B) ex e io su ace wi h small openings, (C) o e iew o
choanosomal skele on, (D) choanosomal desmas, (E) de ail o he sculp u e o desmas, (F) zygoses.
Full-size
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DOI: 10.7717/pee j.8703/fig-22
Ca alho e al. (2020), Pee J, DOI 10.7717/pee j.8703 41/87
ABC
E
FD G
H
mu 2
mu 2
100 um
2 um
30 um
30 um
100 um
Figu e 27 Spicules o Exsupe an ia a chipelagus Ca alho & Pise a, 2019, specimen MNHN-IP-
2008-196. (A)–(D) Phyllo iaenes, (E) acan homic oxeas, (F) acan ho habds, (G) and (H) s ep as e s/
amphias e s. Full-size

DOI: 10.7717/pee j.8703/fig-27
100 um 200 um
100 um 20 um
AB
CD
Figu e 26 Skele on o Exsupe an ia a chipelagus Ca alho & Pise a, 2019, specimen MNHN-IP-
2008-196. (A) O e iew o choanosomal ide s, (B) sub ylos yles c ossing he skele on, (C) de ail o
ide - ype desmas, (D) zygosis and close up o a ide showing he desma o namen a ion.
Full-size
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DOI: 10.7717/pee j.8703/fig-26
Ca alho e al. (2020), Pee J, DOI 10.7717/pee j.8703 48/87

Seamoun , epiben hic Wa én d edge, s . DW258, 3400′N, 3012′W, 420 m), MNHN-IP-
2008-200 (1993-01-31, Pla o Seamoun , epiben hic Wa én d edge, s . DW242, 3312′N,
2857′W, 710 m), MNHN-IP-2008-202 (1993-02-02, A lan is Seamoun , epiben hic
Wa én d edge, s . DW254, 3405′N, 3013′W, 480 m), MNHN-IP-2008-240 (1993-02-01,
Pla o Seamoun , epiben hic Wa én d edge, s . DW246, 3314′N, 2936′W, 520 m),
MNHN-IP-2008-243 (1993-02-02, A lan is Seamoun , epiben hic Wa én d edge, s .
DW258, 3400′N, 3012′W, 420 m). All om he Seamoun 2 campaign.
Compa a i e ma e ial examined. E. a chipelagus (holo ype MNHN DT 782/1 Azo es;
pa a ype MNHN DT 782/2, Azo es; pa a ype DOP 1976, Azo es); E. le ii sp. no .
(holo ype MNHN-IP-2008-201, Hyè es Seamoun ).
Diagnosis. Columna o fici o m Exsupe an ia wi h ide - ype desmas ha ha e smoo h
ube cles ( ew p esen ing ugosi ies).
Desc ip ion (MNHN-IP-2008-196). Small phyma apiniid 22–23 × 8–18 mm in size,
columna o fici o m in habi us, wi h o wi hou la e al p o ube ances (Fig. 3A); some
specimens ha e a “V”shape mo phology; su ace is smoo h wi h conspicuous subde mal
wa e canals gi ing a s iped appea ance o he sponge; oscula o po es a e no isible;
colou beige in e hanol.
Skele on. Ec osome is o med by a laye o phyllo iaenes co e ed by la ge amoun s o
mic oscle es: openings a e su ounded by hese mic oscle es; choanosomal skele on is
mainly buil o ide - ype desmas, ha o m a egula ne wo k wi h la ge spaces in
be ween (Fig. 26); some sub ylos yles (Fig. 26B) and mic oscle es a e also p esen and
sp ead h ough he skele on.
Spicules (MNHN-IP-2008-196).
1. T ide - ype desmas, smoo h, e y ube cula ed, 261–342–419 × 23–30–44 µm in size
(Figs. 26A–26C); ube cles a e smoo h, some imes wi h ugosi ies, 7–10–11 µm
diame e ; ip o he ide is smoo h and has a conical shape (Fig. 26D).
2. Phyllo iaenes, i egula , smoo h cladome 412–450–493 µm in diame e
(Figs. 27A–27D), long habdome, 43–75–126 µm in size, wi h poin ed ip (Fig. 27A).
3. Sub ylos yles o ylo es, smoo h, 401–542–629 × 6.0–9.9–12.8 µm in size (Fig. 26B).
4. Acan homic oxeas, slende , wi h sha p ips, 16.8–22.1–28.1 × 1.6–2.2–3.1 µm (Fig. 27E).
5. Acan ho habds, hick wi h blun ends, 9.8–12.7–17.5 × 1.2–2.0–2.8 µm (Fig. 27F).
6. Amphias e s, wi h se e al a ms co e ed by spines, 5.0–6.6–8.6 µm long (Figs. 27G
and 27H).
Dis ibu ion. E. a chipelagus was ound in Ty o, Hyè es, A lan is, and Pla o
Seamoun s be ween 280 and 1,000 m dep h and also in G an Cana ia island a 660 m
dep h.
Ca alho e al. (2020), Pee J, DOI 10.7717/pee j.8703 49/87
Rema ks. The size o he spicules measu ed in hese specimens a e conside able
smalle when compa ed o he ype ma e ial (Ca alho & Pise a, 2019)(Table 4), bu see
Discussion o mo e in o ma ion ega ding his opic.
Exsupe an ia le ii sp. no .
Figu es 3B,28–29 and Table 4
u n:lsid:zoobank.o g:ac :24B5934A-4767-4429-B172-A649C4CE0D83
Holo ype. MNHN-IP-2008-201 (1993-01-16, Hyè es Seamoun , epiben hic Wa én
d edge, s . DW182, 3123′N, 2854′W, 480 m, Seamoun 2 campaign).
Compa a i e ma e ial examined. E. a chipelagus (holo ype MNHN DT 782/1 Azo es;
pa a ype MNHN DT 782/2, Azo es; pa a ype DOP 1976, Azo es).
Diagnosis. Clus e s o globula o fici o m knob-like sho finge s wi h apical osculum;
phyllo- o disco iaenes as ec osomal megascle es.
Desc ip ion (holo ype MNHN-IP-2008-201). Clus e s o globula o fici o m knob-like
sho finge s, 30 mm in leng h and 29 mm wide; oscula, app oxima ely 2 mm in diame e ,
a e loca ed on he op o he knobs (Fig. 3B); su ace is ugose wi h a s ia ed appea ance
due o he isible subde mal wa e canals; colou is b own in e hanol.
Skele on. Ec osome is composed by phyllo- o disco iaenes ha a e e y a iable in shape,
and se e al mic oscle es; choanosomal skele on has egula and a icula ed ide s,
200 um
100 um 100 um
30 um
AB
CD
Figu e 28 Skele on o Exsupe an ia le ii sp. no ., specimen MNHN-IP-2008-201. (A) Ou line o
ide - ype desmas, (B) ide s, (C) zygosis, (D) de ail o a ide desma.
Full-size
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Ca alho e al. (2020), Pee J, DOI 10.7717/pee j.8703 50/87
o ming an i egula and ela i ely loose ne wo k (Fig. 28); sub ylos yles a e p esen
c ossing he skele on (Figs. 28A and 28B); mic oscle es a e p esen and e y abundan ,
excep o s ep as e s ha a e less nume ous.
30 um
30 um
100 um
100 um
30 um
30 um
2 um
2 um
mu 2
mu
2
m
u 2
2 um
2 um
ABC
DEF
GH I
JK
L
M
Figu e 29 Spicules o Exsupe an ia le ii sp. no ., specimen IP-2008-201. (A)–(C) Top iew o cla-
domes o phyllo- o disco iaenes, (D) and (E) bo om iew o cladomes, (G)–(I) acan homic oxeas,
(J) and (K) acan ho habds, (L) and (M) s ep as e s/amphias e s.
Full-size
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DOI: 10.7717/pee j.8703/fig-29
Ca alho e al. (2020), Pee J, DOI 10.7717/pee j.8703 51/87
Table 4 Compa a i e able o ex e nal mo phology and spicula mic ome ies o all Exsupe an ia species eco ded in he No h A lan ic Ocean. Spicule measu emen s
(n= 30 unless s a ed o he wise) a e p esen ed as minimum–mean–maximum. Da a compiled om he o iginal desc ip ions, o subsequen e-desc ip ions o ype ma e ial
(ma ked wi h numbe s).
Species Habi us Size T ide s Phylo- o
disco iaenes
Sub ylos yles
o ylo es
Acan homic oxeas Acan ho habds Amphias e s Locali y
1
E. cla a
(Schmid ,
1879) (Syn ype
MZUS
PO146)
Cylind ical o cla a e Up o 30 mm
long, 10 mm
hick
230–320 µm
in size
–P esen Fusi o m,
occasionally
cen o ylo es
P esen Slende ays Cuba (dep h
unknown)
2
E. a chipelagus
Ca alho &
Pise a, 2019
(Holo ype
MNHN DT-
782/1)
Columna o fici o m,
wi h o wi hou
la e al p o ube ances
o b anches; su ace
is smoo h, wi h
ma ked wa e canals;
colou beige o
whi ish
20–30 mm ×
10–20 mm
409–693 ×
52–98 µm in
size
Phyllo iaenes.
Cladome:
640–890 µm
in diame e ;
habdome:
229–320 ×
71 µm
Sub ylos yles
o ylos yles:
260–1114 ×
3–38 µm in
size
31–47 ×
2.9–4.2 µm
18–24 ×
2.3–4.1 µm
15–19 ×
1.2–1.7 µm
Azo es
(168–594 m
dep h)
E. a chipelagus
(MNHN-IP-
2008-196).
Columna o fici o m
in habi us, wi h o
wi hou la e al
p o ube ance; wa e
canals isible on he
su ace; colou beige
22–23 ×
8–18 mm
260–362–464
(n=7)×
15–29–
44 µm hick
(n= 15)
Phyllo iaenes.
Cladome:
199–358–
470 µm in
diame e
(=11);
habdome:
140 × 34.4 µm
(n=1)
296–515–
618 × 6.1–
9.7–13.4 µm
(n= 11)
18.6–25.0–44.0 ×
1.4–2.2–3.7 µm
7.2–12.0–15.9 ×
1.2–1.8–
3.2 µm
5.3–8.6–
15.0 µm
(n= 15)
Hyè es
Seamoun
(310 m
dep h)
E. le ii sp. no .
(Holo ype
MNHN-IP-
2008-201)
Clus e s o globula o
fici o m knob-like
sho finge s
293–346–
503 µm in
size, 28–45–
67 µm hick
(n= 12)
Phyllo- o
disco iaenes.
Cladome:
143–299–486
diame e
(n= 20);
habdome:
25–73–130 ×
10 –28–44 µm
(n= 10)
234–307–
436 µm ×
8.6–9.8–
11.3 µm
(n=6)
21.6–28.2–35.6 ×
1.7–2.6–3.8 µm
10.3–14.1–19.3
× 1.9–2.7–
3.5 µm
(n= 25)
7.4–10.0–
14.8 µm
(n= 20).
Hyè es
Seamoun
(480 m
dep h)
No es:
1
Pise a & Lé i (2002d).
2
Ca alho & Pise a (2019).
‘–’ no in o ma ion/no men ioned.
Ca alho e al. (2020), Pee J, DOI 10.7717/pee j.8703 52/87
Spicules (holo ype MNHN-IP-2008-201).
1. T ide - ype desmas, egula , smoo h, 293–346–503 × 28–45–67 µm in size, wi h la ge
and fla ened ube cles ha can be smoo h o e y ube cula ed, 15.4–21.2–29.9 µm in
diame e (Figs. 28A–28D).
2. Phyllo- o disco iaenes, smoo h, cladome e y a iable in shape, 143–299–486 µm in
diame e ; habdome has a conical shape and a sha p ip, 25–73–130 µm × 10–28–44 µm
in size (Figs. 29A–29F).
3. Sub ylos yles, smoo h, la ge, sligh ly cu ed, 234–307–436 × 8.6–9.8–11.3 µm in size
(Figs. 28A and 28B).
4. Acan homic oxeas, hin, sligh ly cu ed, wi h sha p ips, 21.5–26.2–31.6 × 1.8–2.9–4.1
µm (Figs. 29G and 29H); occasionally, hese spicules a e i egula , and exhibi one sha p
and one blun ip, esembling an in e media e s age be ween an acan homic oxea and an
acan ho habd (Fig. 29I);
5. Acan ho habds, small, obus , 9.3–15.1–22.5 × 1.6–2.8–3.8µminsize(Figs. 29J and 29K).
6. Amphias e s, hin wi h spiny a ms, 5.9–8.2–11.5 µm long (Figs. 29L and 29M).
Dis ibu ion. Exsupe an ia le ii sp. no . is known om i s ype locali y, he Hyè es
Seamoun a 480 m dep h.
E ymology. Named a e P o esso Claude Lé i om he Muséum Na ional d’His oi e
Na u elle Pa is (MNHN) o his li elong con ibu ion o he axonomy and sys ema ics o
Po i e a, including li his id sponges.
Rema ks. Recen ly, a e ision o he genus Exsupe an ia allowed o cla i y some
axonomic p oblems by es ablishing wo species, E. cla a (NWA) and E. a chipelagus
(NEA), ha we e p e iously conside ed a single species (Ca alho & Pise a, 2019).
Acco ding o he au ho s, he main di e ences be ween hese wo species a e he desmas
mo phology and o namen a ion.
He e we p opose E. le ii sp.no .asanewspecies, hi do hegenus,basedno onlyon
desmas mo phology and o namen a ion, bu also on he habi us o his new species.
The ide - ype desmas on E. le ii sp. no . esemble he ones ound in E. cla a, i.e., he ube cles
a e o namen ed and he ip o he ide has a ube cle, while in E. a chipelagus i usually
has a conical shape. In gene al, he size o he spicules o E. le ii sp. no . is smalle when
compa ed o he holo ype E. a chipelagus (un o una ely he size o spicules o he E. cla a is
no known, wi h excep ion o he desmas, since he ype ma e ial was deciduous and
mic oscle es we e no p esen (Pise a & Lé i, 2002 )), howe e , he mos dis inc ea u e is he
habi us o E. le ii sp. no .: a clus e o globula knob-like finge s wi h la ge apical oscula on op,
con as ing wi h he columna o fici o m mo phology o he o he wo species.
Subo de Spi opho ina Be gquis & Hogg, 1969
Family Azo icidae Sollas, 1888
Genus Leiode ma ium Schmid , 1870
Ca alho e al. (2020), Pee J, DOI 10.7717/pee j.8703 53/87

Diagnosis. Azo icidae wi h spiny hizoclones and diac ines as megascle es; ec osomal
spicules and mic oscle es a e absen (Pise a & Lé i, 2002i).
Defini ion. Lamella e, pla e-like, oliose, ase- o ea -shaped Azo icidae; oscules a e
isible; choanosomal desmas a e spiny hizoclones; megascle es a e diac ines; mic oscle es
a e absen (Kelly, 2007; modified om Pise a & Lé i, 2002h).
Type species. Leiode ma ium lynceus Schmid , 1870.
Leiode ma ium lynceus Schmid , 1870
Figu es 3C,30–31 and Table 5
AB
CD
EF
200 um 200 um
200 um 200 um
100 um 100 um
Figu e 30 Su ace and skele on o Leiode ma ium lynceus Schmid , 1870, specimen MNHN-IP-2018-
93. (A) O e iew o he ou e su ace wi h la ge and ele a ed oscula (su ace no diges ed in ni ic acid),
(B) o e iew o he inne su ace wi h smalle and dep essed po es (su ace no diges ed), (C) o e iew o
he ou e su ace wi h la ge and ele a ed po es (diges ed su ace in ni ic acid), (D) o e iew o he inne
su ace wi h smalle po es (diges ed su ace), (E) de ail o he oscula, (F) de ail o he po es.
Full-size
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DOI: 10.7717/pee j.8703/fig-30
Ca alho e al. (2020), Pee J, DOI 10.7717/pee j.8703 54/87
Synonym. Azo ica p ei e ae a . enuilamina is Sollas, 1888 (genus ans e and junio
synonym).
Ma e ial examined. MNHN-IP-2018-93 (1988-09-24, Go inge Seamoun , beam awl,
s . CP20, 3633.7′N, 1130.1′W, 305–320 m, Seamoun 1 campaign), MNHN-IP-2008-239
(1993-01-16, Hyè es Seamoun , epiben hic Wa én d edge, s . DW182, 3123′N, 2854′W,
480 m, Seamoun 2 campaign).
Compa a i e ma e ial examined. L. uba sp. no . (holo ype MNHN-IP-2018-72,
Go inge Seamoun ; pa a ype MNHN-IP-2018-73, Go inge Seamoun ).
100 um 20 um
20 um 2 um
10 um 2 um
AB
CD
EF
Figu e 31 Skele on o Leiode ma ium lynceus Schmid , 1870, specimen MNHN-IP-2018-93.
(A) O e iew o choanosomal desmas, (B) hizoclone desmas o ming a e y compac mesh,
(C) de ail o hizoclones, (D) o namen a ion o hizoclone desmas, (E) zygosis be ween se e al ays,
(F) de ail o he sculp u e o he desma. Full-size
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Ca alho e al. (2020), Pee J, DOI 10.7717/pee j.8703 55/87
Diagnosis. Folia e o undula e polymo phic masses, wi h la ge openings in he ou e
su ace o he sponge and small openings in he inne su ace.
Desc ip ion (MNHN-IP-2018-93). La ge olia e o undula e i egula masses, wi h
hick lamellas, 5–12 mm, ha in some cases can o m cups/ unnels (Fig. 3C); inne and
ou e su aces a e di e en om each o he , and i is possible o dis inguished a naked eye;
ou e su ace has la ge openings sligh ly ele a ed om he su ace, 243–269 µm in
diame e , (Figs. 3C,30A and 30C) while he inne su ace is smoo h wi h small openings,
68–145 µm in diame e , e enly dis ibu ed (Figs. 3C,30B and 30D); bo h su aces a e
hea ily p o uded by long oxeas; colou a ies om beige o b own in e hanol.
Skele on. A e y in ica e, i egula and dense mesh o hizoclones desmas ex emely
b anched and spiny (Figs. 31A–31C); he body o his sponge is mainly buil o desmas,
gi ing hem a s ony consis ency; nea he openings and wa e canals, he a ms o he desmas
a e mo e elonga ed; la ge oxeas c oss he skele on and pe o a e he su ace; no mic oscle es.
Spicules (MNHN-IP-2018-93).
1. Rhizoclones, e y spiny a ms wi h mul i u ca ing spines ips, 156–179–223 µm long
and 8.4–19.4–49.9 µm hick (Figs. 31A–31F); zygosis is complex and obus (Fig. 31E).
2. Oxeas, smoo h, s aigh o cu ed, up o 1 mm long and 8.5–9.6–10.7 µm hick.
Dis ibu ion. These specimens we e ound on he Go inge and Hyè es Seamoun s,
be ween 305 and 480 m dep h.
Rema ks. Wi hin Te ac inellida, he genus Leiode ma ium is pa icula ly di ficul om a
axonomic s andpoin , gi en he ew cha ac e s a ailable o dis inguish and desc ibe he
di e en species. In he No h A lan ic, only wo species ha e been desc ibed o da e—
L. lynceus Schmid , 1870 and L. p ei e ae (Ca e , 1873); he o me om specimens
collec ed o he coas o Po ugal, and he la e om Madei a island i.e. bo h om he
NEA bu unknown dep hs. La e , Ca e (1876) o mally explained he di e ences be ween
hese wo species: (1) L. lynceus has la ge oscula loca ed on ou e su ace while in
L. p ei e ae hey a e on he inne su ace; (2) L. p ei e ae has nume ous usi o m oxeas
on he edge o he laminae, while L. lynceus has “isola ed ace a es”(Schmid , 1870)
(howe e hey we e no ound in he edesc ip ion o he holo ype L. lynceus (Pise a &
Lé i, 2002i)). Ano he impo an de ail, is he di e ence be ween he hickness o he
laminae on bo h species, L. lynceus has hinne (3–4 mm) laminae compa ed o
L. p ei e ae (6–17 mm; see Table 5).
In addi ion o hese wo cu en ly ecognized species, Po i ella deciduum (Schmid ,
1879), was also assigned o his genus (Lenden eld, 1903) bu his alloca ion is conside ed
ques ionable (Pise a & Lé i, 2002i). Also, Sollas (1888) epo ed a numbe o a ie ies o
L. p ei e ae om he ma e ial collec ed in he cou se o he Challenge expedi ion in
he A lan ic, iz. A. p ei e ae enuilamina is (Bahia, B azil, unknown dep h) and
A. p ei e ae enuilamina is osculis disjunc is (Be muda, 795–1965 m dep h). Howe e ,
Ca alho e al. (2020), Pee J, DOI 10.7717/pee j.8703 56/87
he ma e ial was deciduous and he e o e he desc ip ions a e incomple e (see also
e iew in Kelly, 2007). Reco ds o L. lynceus and L. p ei e ae o he wes e n A lan ic
(e.g. Van Soes & S en o , 1988) need o be ca e ully e-assessed, as hey may ep esen
di e en and likely undesc ibed species gi en ha se e al pu a i ely new Leiode ma ium
species ha e been epo ed o he opical wes e n A lan ic (Schus e e al., 2019) bu
s ill lack o mal desc ip ion. Topsen (1892) epo s one specimen o Azo ica p ei e ae o
he Azo es (s . 234, 454 m dep h) wi h a s ong blue colo a ion. Howe e , om he
illus a ion p o ided, i appea s ha he specimen has ele a ed openings on he ex e nal
su ace, he eby con o ming o L. lynceus.
The specimens analysed in his s udy a e e y simila o he holo ype o L. lynceus
ega ding he mo phology, su aces and he o namen a ion o he desmas. The only
di e ence lays on he size o he openings: he holo ype has la ge oscula on he ou e
su ace, 500–750 µm in diame e , while in ou specimen oscula a e 243–269 µm in
diame e ; he same happens in ela ion o he po es o he inne su ace o he holo ype,
which a e 156–188 µm in diame e , agains 68–145 µm in ou specimen (Table 5).
Leiode ma ium uba sp. no .
Figu es 3D,32–33 and Table 5
u n:lsid:zoobank.o g:ac :041DAB82-B538-4EB9-A43A-1E3E79B67CF8
Holo ype. MNHN-IP-2018-72 (1988-09-23, Go inge Seamoun , beam awl, s . CP11,
3626.4′N, 1140.2′W, 805–830 m, Seamoun 1 campaign).
Pa a ype. MNHN-IP-2018-73 (1988-09-23, Go inge Seamoun , beam awl, s . CP11,
3626.4′N, 1140.2′W, 805–830 m, Seamoun 1 campaign).
O he ma e ial. MNHN-IP-2018-74 (1988-09-25, Go inge Seamoun , epiben hic Wa én
d edge, s . DW25, 3649.7′N, 1103.3′W, 970–1,035 m, Seamoun 1 campaign); MNHN-IP-
2018-75 (1988-09-23, Go inge Seamoun , beam awl, s . CP11, 3626.4′N, 1140.2′W,
805–830 m, Seamoun 1 campaign); MNHN-IP-2018-76 (1988-09-23, Go inge Seamoun ,
beam awl, s . CP11, 3626.4′N, 1140.2′W, 805–830 m, Seamoun 1 campaign); MNHN-
IP-2008-235 (1993-01-31, Pla o Seamoun , epiben hic Wa én d edge, s . DW242, 3312′N,
2857′W, 710 m, Seamoun 2 campaign); MNHN-IP-2008-237 (1993-02-03, A lan is
Seamoun , epiben hic Wa én d edge, s . DW265, 3429′N, 3036′W, 545 m, Seamoun 2
campaign); MNHN-IP-2008-249b (1993-01-19, Hyè es Seamoun , epiben hic Wa én
d edge, s . DW202, 3116′N, 2843′W, 640 m, Seamoun 2 campaign); MNHN-IP-2008-253
(1993-01-11, G ea Me eo Seamoun , epiben hic Wa én d edge, s . DW159, 2944′N,
2820′W, 330 m, Seamoun 2 campaign); MNHN-IP-2008-255 (1993-01-06, G an Cana ia,
epiben hic Wa én d edge, s . DW130, 2809′N, 1553′W, 660 m, Seamoun 2 campaign).
Compa a i e ma e ial examined. L. lynceus (MNHN-IP-2018-93, Go inge Seamoun ;
MNHN-IP-2008-239, Hyè es Seamoun ).
Diagnosis. Massi e lamella e ase o con o ed walls, some imes o ming a cone, wi h
smoo h and simila su aces.
Ca alho e al. (2020), Pee J, DOI 10.7717/pee j.8703 57/87
Dis ibu ion. Siphonidium elonga us sp. no . was ound in he A lan is, Hyé es, Lion, and
Go inge seamoun s, and in G an Cana ia, be ween 470 and 675 m dep h
E ymology. F om he la in elonga us = elonga ed, due o an elonga ed shape o he desmas,
especially hose composing he fis ules.
Rema ks. Th ee species o Siphonidium ha e been desc ibed in he A lan ic Ocean, and
only one, S. amosum, has been epo ed o bo h sides o he No h A lan ic (Schmid ,
Table 6 Compa a i e able o ex e nal mo phology and spicula mic ome ies o all Siphonidium species eco ded in he No h A lan ic Ocean
and Medi e anean Sea. Spicule measu emen s (n= 30 unless s a ed o he wise) a e p esen ed as minimum–mean–maximum. Da a compiled om
he o iginal desc ip ions, o subsequen e-desc ip ions o ype ma e ial (ma ked wi h numbe s).
Habi us Size Rhizoclones Exo ylos yles S ongyles Locali y
1
Siphonidium
amosum
(Schmid , 1879)
(Holo ype MCZ
6321, 6322)
Small, i egula massi e o
cylind ical, wi h nume ous small
fis ules
20–55 mm
high, 10 mm
wide; fis ules
a e 1–2mm
in diame e
Massi e
hizoclones,
s ongly
ube cula ed,
180–220 µm in
size
Wi h spinose
heads:
160–220 µm
×2–3µm
No p esen Flo ida (dep h
unkonw)
2
Siphonidium
amosum
(Schmid , 1879)
–––800–1,000 µm
long,
4–6µm
hick
–Azo es
(349–793 m)
3
Siphonidium
dubium Lé i,
1959 (Holo ype)
Massi e and ha d sponge wi h a
la ge base, ha i s subdi ided in o
h ee lobes 2 cm long ba ely
sepa a ed, ha ended on a fla
su ace; su ace e icula ed and
co e ed by nume ous po es;
os ia, 1–1.5 cm in diame e ;
colou ligh beige when ali e and
da k b own in e hanol
–Compac No p esen Abundan , g ouped
pe pendicula o he
su ace, ip sligh ly
ugose and ounded,
600–800 µm
P incipe, Gul
o Guinea
(50 m dep h)
Siphonidium
geminum
(Schmid , 1879)
(Holo ype
MNHN DT
2194)
Fla and i egula inc us ing basis
bea ing simple o bi u ca e cone
shaped p olonga ions wi h ound
ends; su ace is co e ed by a
finely co uga ed cu icula
–I egula , like
“ h ee- oo s”
and la e hey
a e bumpy and
ha d
–No p esen Gul o Mexico
(240 m dep h)
Siphonidium
elonga us sp.
no .
(Holo ype
MNHN-IP-
2008-236)
Cylind ical o a bo escen ,
some imes bulb shape; su ace is
smoo h and exhibi s fis ules ha
a e o en closed bu may be open;
colou is beige o b own
33–49 ×
2–9mmin
size
123–197–267 ×
10.4–23.5–40.3
µm; desmas
b anches,
13.8–30.9–88.2
µm long
173–363–504
×
2.9–5.1–6.6
µm (n=4)
No p esen A lan is
seamoun
(545 m dep h)
Siphonidium
elonga us sp.
no .
(Pa a ype
MNHN-IP-
2018-79)
Cylind ical, elonga ed wi h se e al
fis ules; colou is b own
54 × 4–5mm
in size
129–210–326 ×
12.1–22.1–34.0
µm; desmas
b anches,
16.8–45.4–83.7
µm long
248–393 ×
6.5–12.4 µm
(n=3)
No p esen Go inge
seamoun
(605–675 m
dep h)
No es:
1
Redesc ip ion in Pise a & Lé i (2002g).
2
Topsen (1904).
3
Schmid (1879).
Ca alho e al. (2020), Pee J, DOI 10.7717/pee j.8703 64/87

1879;Topsen , 1928,1904,1892;Van Soes , 2017;Van Soes & S en o , 1988) and
Medi e anean Sea (Longo, Mas o o a o & Co ie o, 2005;Vacele , 1969;Zib owius &
Ta iani, 2005). Wi h he edesc ip ion o S. amosum in (Pise a & Lé i, 2002g), a de ailed
accoun o he ex e nal mo phology and spicules was gi en, allowing a be e defini ion o
he species. Despi e he ela i ely simila habi us o S. amosum and S. elonga us sp. no .,
he main di e ence be ween hese wo species elies on he desmas mo phology and
o namen a ion: S. elonga us sp. no . has e y spiny hizoclones wi h slim a ms
o namen ed wi h mic ospines on he edges, con as ing wi h he ube cula ed hizoclones
o S. amosum. Ano he dis inc ea u e, is he p esence o s yles in S. elonga us sp. no .
(e en hough hey a e a e) ha we e ne e men ioned in he edesc ip ion o S. amosum.
Fu he mo e, when S. elonga us sp. no . is compa ed wi h he o he No h A lan ic
species, i s ex e nal mo phology and spicules di e : S. dubium Lé i, 1959 is a massi e
sponge wi h a la ge base, subdi ided in o h ee lobes and he only one wi hin he genus
wi h s ongyles; S. geminum (Schmid , 1879) has a fla and i egula inc us ing base wi h
simple o bi u ca ed cone shape.
Topsen (1904) p esen ed a small desc ip ion o S. amosum om se e al specimens
ound in he Azo es. In his accoun , he shape and he o namen a ion o he desmas a e no
explici ly desc ibed o illus a ed, bu he spicules sizes a e gi en and a e much la ge han
he ones desc ibed by Schmid (1879) om ma e ial collec ed in he Gul o Mexico
(Table 6). The spicules sizes in S. elonga us sp. no . a e mo e simila o he ones in
S. amosum desc ibed by Schmid han o he one desc ibed by Topsen . I was p e iously
s a ed by Van Soes (2017), ha he S. amosum epo ed om he Azo es, is mos likely a
di e en species due o he di e ence in he spicules sizes when compa ed o he ype
ma e ial. A e ision o Topsen ’s ma e ial would be equi ed o cla i y his ques ion.
O de BUBARIDA Mo ow & Cá denas, 2015
Family DESMANTHIDAE Topsen , 1893
Genus Pe omica Topsen , 1898
Synonymy. Monan hus Ki kpa ick, 1903 (junio synonym).
Diagnosis. Massi e, enc us ing o globula shape Desman hidae wi h desmas b anching in
a ious planes o ming a loosely a icula ed o non-a icula ed choanosomal skele al
s uc u e. (Lis -A mi age & Hoope , 2002;Pise a & Lé i, 2002h).
Defini ion. Massi e, enc us ing o globula in shape, wi h o wi hou fis ule-like papillae.
Su ace smoo h, hispid, conules can be p esen . Comp essible o igid, o so o agile
sponges. Ac epid o monoc epid smoo h desmas, b anched in se e al planes. Desmas can
be isola ed, non-a icula ed, used, o dispe sed in he ec osome and choanosome; zygomes
a y om simple o complex; zygosis when p esen , is a ely ully a icula ed in he
skele on u ning in o a loose skele on. O he megascle es a e oxeas, whe e he ips can a y
om sha p o blun . Mic oscle es no p esen (Lis -A mi age & Hoope , 2002;Mu icy
e al., 2001;Pise a & Lé i, 2002h)
Ca alho e al. (2020), Pee J, DOI 10.7717/pee j.8703 65/87
Type species. Pe omica (Pe omica)g imaldii Topsen , 1898 ( ype by mono ype).
Subgenus Pe omica Topsen , 1898
Diagnosis. Fi m and igid sponge, wi h o wi hou papillae, wi h ac epid o monoc epid
desmas ha can o m a loose o well- o med skele on. Oxeas p esen and a iable in size
(Lis -A mi age & Hoope , 2002).
Pe omica (Pe omica)g imaldii Topsen , 1898
Figu es 3F,36–37 and Table 7
Synonym. Pe omica g imaldii Topsen , 1898 accep ed, al e na e ep esen a ion (subgenus
assignmen ).
200 um
100 um
200 um
100 um
10 um
20 um
AB
CD
EF
Figu e 36 Skele on o Pe omica (Pe omica) g imaldii Topsen , 1898, specimen MNHN-IP-2008-92.
(A) O e iew o monoc epid desmas, (B) bundles o anisoxeas and s ongyloxeas, (C) monoc epid
desmas, (D) de ail o anisoxeas and s ongyloxeas c ossing he desmas, (E) zygosis, (F) zygosis be ween a
desma and an oxea. Full-size

DOI: 10.7717/pee j.8703/fig-36
Ca alho e al. (2020), Pee J, DOI 10.7717/pee j.8703 66/87
Ma e ial. MNHN-IP-2018-92 (1998-09-24, Go inge Seamoun , epiben hic Wa én
d edge, s . DW16, 3631.1′N, 1132.5′W, 255–265 m, Seamoun 1 campaign).
Diagnosis. Small sponge wi h a conulose su ace and no papillae; desmas a e monoc epid.
20 um
30 um
mu
0
1
mu 2
mu 00
1
mu
0
01
mu
0
01
A
B
CD
E
Figu e 37 Spicules o Pe omica (Pe omica) g imaldii Topsen , 1898, specimen MNHN-IP-2008-92.
(A) Monoc epid desma poin ing a de ail on he o namen a ion o he ips o he desma, (B) monoc epid
desma showing a close up o he zygome, (C) Anisoxea, (D) and (E) s ongyloxeas.
Full-size

DOI: 10.7717/pee j.8703/fig-37
Ca alho e al. (2020), Pee J, DOI 10.7717/pee j.8703 67/87
Desc ip ion (MNHN-IP-2018-92). F agile, so , massi e sponge wi h a so and conulose
su ace; small, 21 mm heigh and 14 mm wid h; colou whi e in e hanol (Fig. 3F); specimen
in poo condi ion.
Skele on. No clea dis inc ion be ween ec osome and choanosome; skele on is composed
o smoo h monoc epid desmas (Figs. 36A–36D), poo ly a icula ed (Fig. 36E), o ming
a loose, con use and i egula skele on; o he megascle es a e anisoxeas and s ongyloxeas,
a ely ylos yles, usually a anged in bundles (Fig. 36B); i is also possible o obse e a
zygosis be ween he desmas and he oxeas (Fig. 36F); mic oscle es a e absen .
Spicules (MNHN-IP-2018-92).
1. Monoc epid desmas, smoo h, wi h b anches in se e al planes, excep o he ips
which can ha e some o namen a ion wi h a spiny appea ance (Figs. 36 and 37A),
347–499–652 × 8–22–80 µm in size; ips a e 45–82–147 × 10.6–15.2–30.8 µm in
Table 7 Compa a i e able o ex e nal mo phology and spicula mic ome ies o all Pe omica species eco ded in he No h A lan ic Ocean.
Spicule measu emen s (n= 30 unless s a ed o he wise) a e p esen ed as minimum–mean–maximum. Da a compiled om he o iginal desc ip ions,
o subsequen e-desc ip ions o ype ma e ial (ma ked wi h numbe s).
Habi us Size Monoc epid
desmas
Anisoxeas S ongyloxeas Locali y
1
Pe omica
(Chaladesma)
ciocalyp oides (Van
Soes & Zea, 1986)
(Holo ype RMNH
1309)
Basal mass bu ied in he
sand, wi h p oeminen
ape ing fis ules; colou
pale yellow-o ange ali e
and whi e in alcohol
1 cm hick;
fis ules a e
la ge,
100 mm
long,
3–9mm
hick
Smoo h, long
clads, 600–700
µm, clads up o
300 µm, epi habd
180–300 × 10–28
µm; no zygosis
–Some imes modified o
s yles, 378–592 ×
5–22 µm
Saba Bank,
Colombia
(34 m
dep h)
2
Pe omica
(Chaladesma)
ci ina Mu icy,
Hajdu, Mine ino,
Madei a &
Peixinho, 2001
(Holo ype MNRJ
580)
Thickly enc us ing o
massi e, i egula , wi h
small cone-shaped o
digi io m su ace
p ojec ions and la ge
papillae (1–9); colou ali e
b igh o ange-yellow, pale
yellow in e hanol
Base: 30–90 ×
16–60 mm
wide,
4–20 mm
hick
180–337–620 µm,
epi hab
40–87.4–190 ×
9.8–32.6 µm;
cladii
50–126–300 µm.
long
–Usually bo h
ex emi ies a e
ace a e, bu
occasionally s ylo e
o s ongylo e can be
p esen ,
320–527–780 ×
3.2–26 µm
São Sebas ião
island,
B azil
(25 m
dep h)
3
Pe omica
(Pe omica)
g imaldii Topsen ,
1898 (Holo ype
MNHN DT
850)
Massi e sponge, la ge in he
base and slime on op,
co e ed wi h conules
Ve y a iable,
la ge
specimens
up o
40 mm high
and 30 mm
diame e
Up o 570 µm in
size
956–1250 ×
23–30 µm
700–1280 × 23 µm Azo es
(200–599 m
dep h)
Pe omica
(Pe omica)
g imaldii (MNHN-
IP-2008-92)
Massi e sponge wi h a so
and ugose su ace; papillae
absen
Small, 21 mm
heigh and
14 mm
wid h
347–499–652 µm
×8–22–80 µm
890–1,213–1,376
×
14.5–22.5–28.9
µm
541–1,122–1,561 ×
13.1–21.1–33.2 µm
(n= 29)
Go inge
Seamoun
(255–265 m
dep h)
No es:
1
Van Soes & Zea (1986).
2
Mu icy e al. (2001).
3
Redesc ip ion in Pise a & Lé i (2002e).
Ca alho e al. (2020), Pee J, DOI 10.7717/pee j.8703 68/87
size; zygomes a e spiny, mainly in he inne pa , abou 57–118–207 µm in size
(Fig. 37B);
2. Anisoxeas, e y abundan , usi o m, smoo h, wi h ace a e ips, 890–1,213–1,376 µm
long and 14.5–22.5–28.9 µm hick (Fig. 37C);
3. S ongyloxeas, smoo h, can be s aigh o cu ed, wi h one ace a e ip and one blun ip,
541–1,122–1561 µm long and 13.1–21.1–33.2 µm wide (Figs. 37D and 37E).
Dis ibu ion. This specimen was ound on he Go inge seamoun be ween 255 and 265 m
dep h.
Rema ks. Pe omica is a widely dis ibu ed genus, and so a , eigh species ha e been
desc ibed. In he No h A lan ic, h ee species ha e been epo ed, P. (Chaladesma)
ciocalyp oides and P. (Chaladesma) ci ina o he NWA and P.(Pe omica)g imaldii om
he NEA and MED (Table 7). P.(P.) g imaldii was fi s desc ibed om he Azo es
a chipelago by Topsen (1898) whe e i was ound o be a e y common sponge, collec ed
h oughou he a chipelago be ween 200 and 914 m dep h (Topsen , 1928,1904, 1898).
This species has been also epo ed om he MED (Bou y-Esnaul , Pansini & U iz, 1994;
Puli ze -Finali, 1972) and since mic ospine desmas’ e mina ions we e absen , P.(P.)
massalis Dendy, 1905 (a species om he Indian Ocean) and P.(P.) g imaldii we e
synonymized (Puli ze -Finali, 1972). Acco ding o Mu icy e al. (2001), hese mic ospines
a e no p esen in all desmas in he same specimen and hey can be a e. The e o e, he
absence o mic ospines in he desmas is no enough o dis inguish one species om
ano he . A mo e de ailed examina ion o he specimens om he MED would be necessa y
o allow o cla i y his unce ain y (Mu icy e al., 2001) and make su e he Pe omica ound
in MED a e in ac P.(P.) g imaldii. In he specimen examined in his s udy spicules
sizes a e e y simila o hose o he holo ype ( om he Azo es) and he mic ospines in he
e mina ion o he desmas a e p esen and e y e iden (Fig. 37).
DIVERSITY
The specimens desc ibed in he p esen wo k cons i u e he fi s eco ds o li his id
demosponges o hese wo g oups o NEA seamoun s, excep o Exsupe an ia
a chipelagus. The Me eo seamoun g oup ha bou s a mo e di e se li his id auna, 15
species, compa ed o he Lusi anian seamoun g oup, whe e six species a e eco ded
(Table 8). A a smalle scale, he Hyè es seamoun is he mos di e se whe e eigh species,
namely N. pomponiae sp. no ., M. c . azo ica,M. obus a., E. a chipelagus,E. le ii sp. no .,
L. lynceus,L. uba sp. no . and S. elonga us sp. no . we e ound, ollowed by he
Go inge and A lan is (six species), Pla o and G ea Me eo (fi e species), Ty o
( h ee species) and Lion seamoun (one species). Two specimens we e ound on he
An ial ai and Ampè e seamoun (one on each) bu i was no possible o iden i y hem
down o species le el because hey we e small and inc us ing specimens, possibly young
indi iduals o M. obus a. The majo i y o he species ha e a es ic ed dis ibu ion
Ca alho e al. (2020), Pee J, DOI 10.7717/pee j.8703 69/87

Table 8 O e all dis ibu ion o li his id demosponges in he No heas A lan ic and Medi e anean Sea. Species ound in his s udy () and
eco ds om he li e a u e (○). Newly desc ibed species a e highligh ed in bold.
Seamoun s Oceanic a chipelagos Con inen al shel /
slope
Species ATR ATL TYR PLT HYR MET LIO GOR AMP AZO MAD SEL CAN PT MED MOR
CORALLISTIDAE Sollas, 1888
Co allis es elegan io Schmid , 1870 ○
Co allis es masoni (Bowe bank, 1869)○○
Isabella ha bo b anchi Ca alho, Pomponi
& Xa ie , 2015
○
Neoph issospongia endoumensis Pise a &
Vacele , 2011
○
Neoph issospongia nana Manconi &
Se usi, 2008
○
Neoph issospongia noli ange e (Schmid ,
1870)
○○ ○○ ○
Neoph issospongia adjae Pise a & Vacele ,
2011
○
Neosch ammeniella bowe bankii (Johnson,
1863)
○○
Neosch ammeniella inaequalis sp. no . 
Neosch ammeniella pise ai sp. no . 
Neosch ammeniella pomponiae sp. no . 
Neosch ammeniella sp. 
THEONELLIDAE Lenden eld, 1903
Discode mia a bo sp. no . 
Discode mia kellyae sp. no . 
Discode mia polydiscus (Bowe bank, 1869)○○
Discode mia polymo pha Pise a & Vacele ,
2011
○
Discode mia ami e a Topsen , 1892 ○
Discode mia c . ami e a 
Discode mia sp. 
Discode mia e ucosa Topsen , 1928  ○○ ○
Theonella annula a Lenden eld, 1907 ○
MACANDREWIIDAE Sch ammen, 1924
Macand ewia azo ica G ay, 1859 ○○○
Macand ewia c . azo ica   
Macand ewia schus e ae sp. no .  
Macand ewia minima sp. no . 
Macand ewia amosa Topsen , 1904 ○
Macand ewia obus a Topsen , 1904 ○
Macand ewia sp. 
PHYMARAPHINIIDAE Sch ammen,
1924
Ca alho e al. (2020), Pee J, DOI 10.7717/pee j.8703 70/87
and a e ound only in one o wo seamoun s, excep E. a chipelagus, M. c . azo ica,
M. schus e ae sp. no ., L. uba sp. no . and S. elonga us sp. no ., ha a e dis ibu ed
be ween h ee o fi e di e en seamoun s. Fou species we e also sampled in G an
Cana ia, M.c .azo ica (480 m dep h), E. a chipelagus (660 m dep h), L. uba sp. no .
(660 m dep h) and S. elonga us sp.no .(470mdep h), he wola e ep esen ing
he fi s eco ds o he Cana y Islands. New ba hyme ic eco ds we e also epo ed
o h ee species, iz. M. obus a (705 m), D. ami e a (300–420 m), D. e ucosa
(338–520 m) (Fig. 38).
Some o he examined ma e ial was o e y small size and/o in poo condi ion, which
hampe ed i s iden ifica ion o lowe axonomic le els. These specimens we e he e o e no
iden ified and a e no included in his manusc ip (see Supplemen al Ma e ial).
Table 8 (con inued).
Seamoun s Oceanic a chipelagos Con inen al shel /
slope
Species ATR ATL TYR PLT HYR MET LIO GOR AMP AZO MAD SEL CAN PT MED MOR
Exsupe an ia a chipelagus Ca alho and
Pise a, 2018
  ○○ 
Exsupe an ia le ii sp. no . 
AZORICIDAE Sollas, 1888
Leiode ma ium lynceus Schmid , 1870 ○○ ○ ○
*
Leiode ma ium p ei e ae (Ca e , 1876)○○ ○
Leiode ma ium uba sp. no .  
SCLERITODERMIDAE Sollas, 1888
Aciculi es medi e anea Manconi, Se usi &
Pise a, 2006
○
Mic oscle ode ma lamina Pe ez e al., 2004 ○
SIPHONIDIIDAE Lenden eld, 1903
Gas ophanella phoeniciensis Pe ez e al.,
2004
○
Siphonidium elonga us sp. no .  
Siphonidium amosum (Schmid , 1870)?○
DESMANTHIDAE Topsen , 1893
Desman hus inc us ans (Topsen , 1889)○
Pe omica (Pe omica)g imaldii Topsen ,
1898
○○
Sulcas ella enens (Vacele , 1969)○
To al o species/ axa 36 1 6 3 5 8 5 1 6 1 11 7 2 10 2 15 1
No es:
AMP, Ampe e seamoun ; ATL, A lan is seamoun ; ATR, An ial ai seamoun ; AZO, Azo es; CAN, Cana ies; GOR, Go inge seamoun ; HYR, Hyè es seamoun ; LIO,
Lion; MAD, Madei a; MED, Medi e anean Sea; MET, G ea Me eo seamoun ; MOR, Mo occo; PLT, Pla o seamoun ; PT, Po ugal; SEL, Sel agens; TYR, Ty o
seamoun .
Sou ces o he li e a u e eco ds: Bowe bank (1869);Ca e (1876);Ca alho, Pomponi & Xa ie (2015);Ca alho & Pise a (2019);C uz (2002);du Bocage (1869);G ay
(1859);Johnson (1863);Lenden eld (1907);Longo, Mas o o a o & Co ie o (2005);Magnino e al. (1999);Maldonado e al. (2015);Manconi, Se usi & Pise a (2006);
Manconi & Se usi (2008);Pe ez e al. (2004);Pise a & Vacele (2011);Puli ze -Finali (1972);Schmid (1870);Topsen (1889,1892,1898,1904,1928); Vacele (1969).
*
Va enuilamina e (Topsen , 1928).
?The assignmen o he specimens examined by Topsen in Azo es need o be e ised in o de o cla i y i i is in ac S. amosum.
Ca alho e al. (2020), Pee J, DOI 10.7717/pee j.8703 71/87
DISCUSSION
Di e si y and biogeog aphic pa e ns
Wi h he p esen wo k, we desc ibe o he fi s ime he li his id auna o wo seamoun
g oups o he NEA, he G ea Me eo and he Lusi anian seamoun s. All o he 17 species
he e epo ed cons i u e new eco ds o hese seamoun s and en a e new o science.
The only excep ion is E. a chipelagus p e iously epo ed o he G ea Me eo Seamoun
as Exsupe an ia sp. (Cá denas e al., 2011). These 10 newly desc ibed species add o
he 17 species p e iously epo ed o he NEA, ep esen ing an inc ease o app oxima ely
60% o he li his id di e si y o his a ea. These findings show how unde s udied he
auna o hese ecosys ems is and sugges s ha addi ional species a e likely o be ound as
su ey e o s inc ease. I also concu s wi h p e ious s udies made o o he in e eb a e
g oups based on ma e ial collec ed om he same seamoun s whe e se e al new
species we e desc ibed (Be ning, Ha melin & Bade , 2017;Cá denas e al., 2018;Geo ge
& Schminke, 2002;Go as, 2007;Sou o, Be ning & Os o sky, 2016). The G ea Me eo
g oup, appea s o ha bou a mo e di e se li his id auna, wi h a o al o 15 species
(nine new o science), whe eas in he Lusi anian g oup, six species we e eco ded ( ou
new o science). In e es ingly, only a ela i ely small p opo ion o he li his id species
known om he NEA (7 ou o 17) we e ound du ing he p esen s udy. Finally, he
finding o 19 la ge specimens o M.c .azo ica in he same s a ion in he Hyè es
Seamoun (s . DW202), sugges s ha his species may occu in ela i ely la ge densi ies,
possibly o ming a sponge g ound in his a ea. Howe e , his would equi e e ifica ion
wi h o he sampling and obse a ion ools such as emo ely ope a ed o au onomous
unde wa e ehicles (ROV/AUV). Such finding would add on o he agg ega ions domina ed
by Leiode ma ium p ei e ae, ecen ly epo ed on h ee seamoun s in he Wes e n
Figu e 38 Ba hyme ic dis ibu ion o he li his id demosponges collec ed du ing he Seamoun 1
and Seamoun 2 expedi ions on he No heas A lan ic seamoun s.
Full-size

DOI: 10.7717/pee j.8703/fig-38
Ca alho e al. (2020), Pee J, DOI 10.7717/pee j.8703 72/87
Medi e anean Sea (Maldonado e al., 2015), which sugges s ha some ex an li his ids may
s ill o m highly s uc u ed habi a s compa able o he Mesozoic ee s (Maldonado e al.,
2015;Reid, 1967),
Se e al pa adigms in seamoun ecology, including he seamoun endemism hypo hesis,
ha e been hea ily deba ed in ecen yea s, wi h some au ho s conside ing seamoun s
as places o high endemism (de Fo ges, Koslow & Poo e, 2000), while o he s a ibu ed
he obse ed pa e ns o sample bias (Samadi e al., 2006;seealsoMcClain, 2007;
Rowden e al., 2010). In ou s udy, he majo i y o he species (Neosch ammeniella
inaequalis sp. no ., N. pise ai sp. no ., N. pomponiae sp. no ., Discode mia. a bo sp.
no ., D. kellyae sp. no ., D. ami e a, Macand ewia minima sp. no ., M. obus a,
Leiode ma ium lynceus and Exsupe an ia le ii sp. no .) we e only ound on one o he
seamoun s. These findings concu wi h a s udy on li his ids o he No olk Ridge
(New Caledonia) whe e he au ho s epo ed 16 species (se en new o science, including
a new genus) wi h he hal o he species (eigh ) es ic ed o one seamoun (Schlache -
Hoenlinge , Pise a & Hoope , 2005). On he o he hand, fi e species, M.c .azo ica,
M. schus e ae sp. no ., S. elonga us sp. no ., L. uba sp. no .andE. a chipelagus,ha ea
wide dis ibu ion ( ound in h ee o fi e seamoun s), and he la e h ee a e sha ed
be ween he wo seamoun g oups. The di e ences in di e si y and dis ibu ion ound in
ou s udy may be a esul o une en sampling e o be ween he di e en seamoun s
(be ween 2 and 35 s a ions) and he wo seamoun g oups (92 s a ions in Seamoun 1 s
131 s a ions in Seamoun 2).
When examined a a la ge scale, seamoun s sha e mos species wi h he Azo es and
Cana y a chipelagos, wi h se en (D. ami e a,D. e ucosa,M. azo ica,M. obus a,
E. a chipelagus,L. lynceus,P.(P.) g imaldii) and six species (D. e ucosa,M. azo ica,
E. a chipelagus,L. lynceus,L. uba sp. no . and S. elonga us sp. no .) sha ed, espec i ely.
Gi en he ela i e p oximi y be ween locali ies and he oceanog aphic se ing, i would
be expec ed ha he Azo es would sha e mo e species wi h he G ea Me eo g oup,
ins ead o he Cana ies, Madei a, Sel agens and he con inen al shel o he Lusi anian
g oup (Fig. 1). Howe e , his is no obse ed in ou s udy as only wo species (D. ami e a
and M. obus a) a e exclusi ely sha ed be ween he Azo es and he Me eo Seamoun
g oup. One species (L. lynceus) is common o Azo es, Madei a, Cana ies and he wo
g oups o seamoun s, and wo species (M. azo ica and D. e ucosa) a e sha ed be ween
he Me eo g oup and he oceanic islands. All he species ound in he Lusi anian g oup a e
sha ed wi h he a chipelagos and/o he Me eo Seamoun , wi h only one excep ion,
N. inaequalis sp. no . ha is exclusi ely known om he Go inge Seamoun . Howe e ,
none o he species epo ed om he Po uguese (Co allis es elegan io Schmid , 1870) and
Mo occan con inen al shel es (Theonella annula a Lenden eld, 1907) we e ound o
occu in he Lusi anian seamoun s g oup. I should be no ed ha he desc ip ion o
C. elegan io is ague and does no p o ide a de ailed cha ac e iza ion o all spicules.
Mo eo e , his species was ne e obse ed since i s desc ip ion by Schmid (1870) in
Po ugal o in he su ounded a eas, hus i should be conside ed a axon inqui endum.
Ca alho e al. (2020), Pee J, DOI 10.7717/pee j.8703 73/87
Pila Ríos pe o med he expe imen s, au ho ed o e iewed d a s o he pape , and
app o ed he final d a .
Ja ie C is obo pe o med he expe imen s, au ho ed o e iewed d a s o he pape ,
and app o ed he final d a .
Hans To e Rapp pe o med he expe imen s, au ho ed o e iewed d a s o he pape ,
and app o ed he final d a .
Joana R. Xa ie concei ed and designed he expe imen s, pe o med he
expe imen s, analyzed he da a, au ho ed o e iewed d a s o he pape , and app o ed
he final d a .
Da a A ailabili y
The ollowing in o ma ion was supplied ega ding da a a ailabili y:
Da a is a ailable a PANGAEA: Ca alho, F ancisca C; Cá denas, Paco; Rios, Pila ;
Rapp, Hans To e; Xa ie , Joana R (2018): Li his ids om he Seamoun s o he No heas
A lan ic Ocean. PANGAEA, DOI 10.1594/PANGAEA.896492.
Specimens examined in his s udy a e s o ed in he ’zoo hèque’o he Muséum Na ional
d’His oi e Na u elle o Pa is, F ance (MNHN).
Specimen numbe s and deposi ion in o ma ion a e a ailable as a Supplemen al File.
New Species Regis a ion
The ollowing in o ma ion was supplied ega ding he egis a ion o a newly desc ibed
species:
Publica ion LSID: u n:lsid:zoobank.o g:pub:A0DA0236-4579-47A4-8BE4-E68803C2EC8F.
Neosch ammeniella inaequalis sp. no .: U n:lsid:zoobank.o g:ac :8A516D9B-5351-
47AF-8EC2-7EBC44166D35.
Neosch ammeniella pise ai sp. no .: U n:lsid:zoobank.o g:ac :77F1F52E-28C9-43C0-
A501-1ADAD03241A5.
Neosch ammeniella pomponiae sp. no .: u n:lsid:zoobank.o g:ac :2AA76193-B27E-
491E-8E50-FE591786FA26.
Discode mia a bo sp. no .: U n:lsid:zoobank.o g:ac :7A732A92-8D8B-4D73-97B1-
CD53E9494121.
Discode mia kellyae sp. no .: u n:lsid:zoobank.o g:ac :E7A06142-4AF7-404E-B369-
B30240ADE5F4.
Macand ewia schus e ae sp. no .: u n:lsid:zoobank.o g:ac :2BA2C1EF-8FAB-4C91-
89CB-DCB59DDA61EB.
Macand ewia minima sp. no .: u n:lsid:zoobank.o g:ac :E405AE49-5636-4778-9B07-
ED39E9EBB7BE.
Exsupe an ia le ii sp. no .: u n:lsid:zoobank.o g:ac :24B5934A-4767-4429-B172-
A649C4CE0D83.
Leiode ma ium uba sp. no .: u n:lsid:zoobank.o g:ac :041DAB82-B538-4EB9-A43A-
1E3E79B67CF8.
Siphonidium elonga us sp. no .: u n:lsid:zoobank.o g:ac :26B193F9-2588-4479-ACB2-
27AD1945DEE4.
Ca alho e al. (2020), Pee J, DOI 10.7717/pee j.8703 80/87

Supplemen al In o ma ion
Supplemen al in o ma ion o his a icle can be ound online a h p://dx.doi.o g/10.7717/
pee j.8703#supplemen al-in o ma ion.
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