123
A new ee -dwelling co al, Pa ona giannii sp. no . (Scle ac inia,
Aga iciidae), wi h an o e iew o he skele al mo phology o he
ype specimens o he genus Pa ona
F ancesca Benzoni1
1 Biological and En i onmen al Science and Enginee ing Di ision, Ma ine Science P og am, HaBB Lab, King Abdullah Uni e si y o Science and Technology,
Thuwal, Saudi A abia
Co esponding au ho : F ancesca Benzoni ( ancesca.benz[email p o ec ed])
Copy igh : © F ancesca Benzoni.
This is an open access a icle dis ibu ed unde
e ms o he C ea i e Commons A ibu ion
License (A ibu ion 4.0 In e na ional – CC BY 4.0).
Resea ch A icle
Abs ac
Ha d co al species in he aga iciid genus Pa ona a e common in shallow and meso-
pho ic co al ee communi ies ac oss he Indo-Paci ic, bu hei axonomy has long been
o e looked. Pa ona giannii sp. no . is he e desc ibed based on newly collec ed ma e ial
ac oss he Indian Ocean, and his o ical museum specimens. In i o and skele al mo -
phological ea u es a e desc ibed, diagnos ic cha ac e s measu ed, and compa ison
wi h congene s pe o med. The new species o ms an enc us ing co allum de oid o
aised idges on i s su ace. I s co alli es a e lush wi h he su ace, no inclined, and less
han hal a co alli e diame e apa . Co alli es a angemen is hamnas e oid and se ies
can o m locally. Whe e hey occu , he adial elemen s un o e he sha ed walls o he
adjacen se ies’ co alli es c ea ing a ladde -like a angemen simila o ha obse ed
o e he idges in congene s like Pa ona a ians and Pa ona chi iquiensis. Despi e he
lack o idges, P. giannii sp. no . has mo phological a ini ies wi h hese wo species.
Howe e , based on p e iously published mo phome ics and me is ics, P. giannii sp.
no . co alli es a e la ge and mo e c owded, and he p ima y sep a a e longe and mo e
nume ous. In i o, he new species is dis inguished by ully ex ended whi e o beige
en acles du ing he day ime, gi ing i a whi e-bea ded appea ance. Despi e he ecolog-
ical ele ance o Pa ona co als, a axonomic e ision o he genus is o e due, and he
exis ing molecula s udies indica e ha i is polyphyle ic. He e, he p oposed placemen
o P. giannii sp. no . in he genus is based on mo phological e idence alone and phylog-
enomic analyses a e cu en ly in p og ess.
Key wo ds: Mo phome ics, museum collec ions, ee -building co al, skele al mo phol-
ogy, axonomy
In oduc ion
The colonial scle ac inian genus Pa ona Lama ck, 1801 is a common com-
ponen o he shallow-wa e co al ee communi ies ac oss he Indo-Paci -
ic om he Red Sea and he Sou hwes e n Indian Ocean o he Eas Paci ic,
spanning opical and sub- opical la i udes (Du ham and Ba na d 1952; Ve on
and Pichon 1980; Wijsman-Bes e al. 1980; Pichon and Benzoni 2007; Glynn e
al. 2017; Be umen e al. 2019). Pa ona co als a e zooxan hella e ee -builde s
Academic edi o :
Ka eh Samimi-Namin
Recei ed:
30 July 2025
Accep ed:
1 Oc obe 2025
Published:
19 No embe 2025
ZooBank: h ps://zoobank.
o g/0072E066-E864-4B37-8D67-
5EDE6D5F81A2
Ci a ion: Benzoni F (2025) A new
ee -dwelling co al, Pa ona giannii
sp. no . (Scle ac inia, Aga iciidae),
wi h an o e iew o he skele al
mo phology o he ype specimens
o he genus Pa ona. ZooKeys 1260:
123–147. h ps://doi.o g/10.3897/
zookeys.1260.167263
ZooKeys 1260: 123–147 (2025)
DOI: 10.3897/zookeys.1260.167263
124
ZooKeys 1260: 123–147 (2025), DOI: 10.3897/zookeys.1260.167263
F ancesca Benzoni: A new Pa ona co al om Indian Ocean ee s
ac i ely con ibu ing o he s uc u e and unc ioning o co al ee ecosys ems
(To ole o-Langa ica e al. 2022). Some ondose o columna species can
o m gian colonies (Mezaki e al. 2014; Siena e al. 2025) and/o monospe-
ci ic s ands locally eaching me ic size (Ve on 1986; Mon agne e al. 2013).
Pa ona species suppo a di e se and o en specialized associa ed auna and
mic obiomes (Hoeksema and an de Meij 2013; Mon ano and Maggioni 2018;
Hu e al. 2020; Meh o a e al. 2020; Yu e al. 2020; Cheng e al. 2023; Delgadillo
O donez e al. 2022; an de Schoo and Hoeksema 2024; Bähe e al. 2025).
Mo eo e , some species a e dep h gene alis s and h i e below 30 m dep h as
componen s o mesopho ic co al ecosys ems (Luck e al. 2013; K ame e al.
2019; Longenecke e al. 2019). Pa ona co als a e conside ed s ess- ole an
(Gleason 1993; Hun ing on e al. 2022) and ha e been s udied o hei acclima-
o y abili y in ma ginal en i onmen s and esilience o changes in en i onmen-
al condi ions due o ocean wa ming (McClanahan, 2000; Soland e al. 2003;
Guzman and Co és 2007; To ole o-Langa ica e al. 2022; Zhang e al. 2022,
2025; Khen e al. 2024). Despi e i s ecological ele ance and being he a ge
o undamen al and expe imen al esea ch e o s, species bounda ies wi hin
he genus Pa ona and phylogene ic ela ionships a e s ill only pa ially explo ed
h ough a gene ic app oach. A ailable molecula da a on a subse o species
sugges ha he genus is polyphyle ic and equi es o mal axonomic e ision
(Ma é 2003; Moo hien Pillay e al. 2006; Ki aha a e al. 2012; Luck e al. 2013;
Waheed e al. 2015; Te aneo e al. 2017; Quek e al. 2023).
Belonging o he amily Aga iciidae G ay, 1847, Pa ona cu en ly includes 20
alid ex an species al hough 57 nominal ex an species ha e been his o ically
asc ibed o i (Hoeksema and Cai ns 2024). Mo phologically, species cu en ly
in he genus display ema kable di e ences in co allum and co alli e shape, co -
alli e a angemen , and columella s uc u e (Ve on and Pichon 1980; La ypo
2014). Typically o an aga iciid, co alli e a angemen in Pa ona is hamnas-
e oid: he adial elemen s un o e he co allum su ace om a co alli e cen-
e o he adjacen ones, passing o e hei walls, which a e gene ally poo ly
de eloped and sca cely isible (Che alie and Beau ais 1987). The o ma ion
o new polyps in a colony, and new co alli es in i s co allum, occu s h ough in-
a en acula budding and can con inue wi hou he ac ual loss o o ganic con-
nec ion among he buds (Ma hai 1948c). This p ocess in some species can
lead o he o ma ion o se ies o polyps aligned in alleys. Adjacen se ies a e
sepa a ed by a iably con inuous sha ed co alli e walls ha can de elop in o
aised longe idges o sho e mon icules (Ma hai 1948a; Ve on and Pichon
1980: igs 47–53). These indeed a y in heigh , leng h and o ien a ion depend-
ing on he species, and on hei posi ion in a single colony (Ma hai 1948b).
The adial elemen s unning abo e any shape o elonga ed idge in Pa ona a e
pa allel among hem and pe pendicula o he idge’s main axis gi ing a ypical
ladde -like a angemen (Ve on and Pichon 1980: igs 21, 24).
An undesc ibed species o ee -dwelling Pa ona was collec ed a di e en lo-
cali ies in he Gul o Tadjou a, Gul o Aden, A abian Sea, S ai o Oman, he SW
Indian Ocean, and he NE Indian Ocean. I is ecognized based on i s enc us ing
g ow h o m, a smoo h co allum su ace de oid o idges o mon icules, a c owd-
ed co alli e a angemen , s ongly al e na ing adial elemen s wi h he alle ones
always lush wi h he co allum su ace gi ing i an e en appea ance. In i o, he
species has ully ex ended polyp en acles a day ime. O al disks and/o en acles
125
ZooKeys 1260: 123–147 (2025), DOI: 10.3897/zookeys.1260.167263
F ancesca Benzoni: A new Pa ona co al om Indian Ocean ee s
a e whi e and gi e he colonies a dis inc i e whi e bea ded appea ance. The new
species skele al mac o- and mic omo phology a e desc ibed and illus a ed in de-
ail. Measu emen s and coun s o skele al cha ac e s p e iously analyzed o wo
congene s wi h an enc us ing g ow h o m and co alli e mo phological a ini ies
wi h i , Pa ona chi iquiensis Glynn, Ma é & S emann, 2001 and Pa ona a ians (Ve -
ill, 1864) (Ma é 2003), we e ob ained om he undesc ibed Pa ona hus allowing
a di ec quan i a i e compa ison. In he discussion sec ion, ype ma e ial o nine
congene s and e e ence ma e ial om wo mo e is illus a ed o show he mo -
phological a ia ion o Pa ona species and he di e ences wi h he new species.
Ma e ials and me hods
Collec ion and p ocessing
Ma e ial was collec ed du ing SCUBA a 13 locali ies in he sou hwes e n Indi-
an Ocean (Fig. 1A) and he Gul s o Tadjou a and Aden, he A abian Sea, and
he Gul o Oman (Fig. 1B) du ing mul iple expedi ions be ween 2007 and 2022.
Be o e collec ion, li ing co al colonies we e imaged in si u wi h digi al came -
as o documen colony g ow h o m, appea ance and colo a ion, and polypa
ea u es. Specimens, mos ly colony agmen s, we e sampled by hamme and
chisel. Once on land, each specimen was agged and subsampled o p ese e
co al issue in molecula g ade e hanol o DNA ex ac ion. Co alla we e hen
le o e nigh in a sodium hypochlo i e solu ion (household bleach) o emo e
o ganic ma e , insed in eshwa e , and d ied.
Imaging and mo phological analysis
Fo mac o-mo phological obse a ions o co allum su ace and co alli e a -
angemen , specimens we e pho og aphed wi h a Nikon Coolpix digi al came a
wi h a e e ence scale and h ough a Leica M80 mic oscope equipped wi h
Figu e 1. Map showing he ype (s a ) and o he collec ion (ci cle) locali ies o Pa ona giannii sp. no . A. Collec ion locali-
ies in he SW Indian Ocean, and B. The seas a ound he A abian Peninsula. 1 = Maskali Island, Djibou i; 2 = Aden, Yemen;
3 = Balha , Yemen; 4 = Hyllanyia Island, Bi Ali, Yemen; 5 = Bu um, Yemen; 6 = Al Mukallah, Yemen; 7 = Hawla , Soco a
Island, Yemen; 8 = Dhalku , Oman; 9 = Mi ba , Oman; 10 = Qinqa i Bay, Oman; 11 = Musca , Oman; 12 = Mayo e Island; 13
= Mahé Island, Seychelles; 14 = Galle, S i Lanka; 15 = Pulau Songsong, Malaysia. Scale ba : 200 km.
126
ZooKeys 1260: 123–147 (2025), DOI: 10.3897/zookeys.1260.167263
F ancesca Benzoni: A new Pa ona co al om Indian Ocean ee s
a Leica IC80HD came a a known magni ica ion. To in es iga e and desc ibe
he skele al ea u es mic o-mo phology and hei a iabili y, agmen s we e
ob ained om wo dis inc specimens displaying di e en deg ees o calci i-
ca ion o he co allum, namely UNIMIB BAL253 ( hinne ) and UNIMIB AD016
( hicke ). F agmen s we e g inded, moun ed on s ubs using sil e glue, spu -
e -coa ed wi h conduc i e gold ilm and examined using a Vega Tescan Scan-
ning Elec on Mic oscope a Uni e si y o Milano-Bicocca, I aly. Fo specimen
size measu emen s, a calipe was used. Co alli e and adial elemen measu e-
men s and coun s we e aken on digi al images wi h isible e e ence scale
using Image J . 1.54p (Schneide e al. 2012) o 13 specimens. Fo each
specimen, i e co alli es we e selec ed o measu emen s and coun s. Only ull-
g own co alli es, no adjacen and no unde going budding we e conside ed.
Following Ma é (2003: ig. 3), he ollowing skele al cha ac e s ha p o ided
s a is ically signi ican di e ences among Pa ona species we e measu ed and
coun ed: maximum calicula diame e , minimum calicula diame e , maximum
columella diame e , minimum columella diame e , main sep a (S1, 2) leng h,
numbe o sep a, numbe o sep a eaching he columella. S a is ical analysis
o e alua e i any o he cha ac e s we e s a is ically signi ican ly di e en in
he new species compa ed o he enc us ing species examined by Ma é (2003)
was no possible because he o iginal da ase was no included in which only
a e ages (± SE) we e epo ed o each cha ac e . The e o e, compa ison o
cha ac e s is he ea e pe o med based on he a ailable in o ma ion.
Reposi o ies and ins i u ional ac onyms
The specimens collec ed o his s udy a e deposi ed in he ollowing eposi o-
ies: Muséum na ional d'His oi e na u elle (MNHN), Pa is, F ance; Flo ida Mu-
seum o Na u al His o y (UF), Gaines ille, Flo ida, USA; King Abdullah Uni e si y
o Science and Technology (KAUST), Thuwal, Saudi A abia; Uni e si y o Mila-
no-Bicocca (UNIMIB), Milan, I aly; Uni e si y o he Seychelles (UNISEY), Anse
Royale, Mahé, Seychelles.
In Sep embe 2004, a isi o he Na u al His o y Museum (NHMUK), London,
UK, allowed he disco e y o his o ical specimens showing he same diagnos-
ic mac o and mic o-mo phological cha ac e s as he collec ed ma e ial o
his s udy. In he accompanying labels, hey had all been iden i ied as Pa ona
explanula a (Lama ck, 1816). Collec ed in he las cen u y by di e en scien-
is s om he Seychelles, S i Lanka, and Malaysia, hese specimens ex end he
known geog aphic dis ibu ion o he new species o wo addi ional locali ies in
he NE Indian Ocean (Fig. 1A).
Examined ype and e e ence ma e ial
Fo compa a i e pu pose, ype o ep esen a i e specimens o Pa ona spe-
cies examined o collec ed by he au ho , espec i ely, we e included in his
s udy and a e lis ed he ea e . The holo ype o Pa ona diminu a Ve on, 1990
(QMT G32480) was examined and pho og aphed a he Queensland Museum
T opics (QMT), Towns ille, Queensland, Aus alia, in May 2009. The holo ype
o Pa ona minu a Wells, 1954 (USNM 44786) was s udied and pho og aphed
in June 2009 a he Uni ed S a es Na ional Museum (USNM), Washing on DC,
127
ZooKeys 1260: 123–147 (2025), DOI: 10.3897/zookeys.1260.167263
F ancesca Benzoni: A new Pa ona co al om Indian Ocean ee s
USA. Du ing a subsequen isi in Ma ch 2023, he ype specimens o Pa ona
decussa a Dana, 1846 (USNM 201) and Pa ona chi iquiensis Glynn, Ma é &
S emann, 2001 (USNM 100871) we e also examined and imaged. Images o
he holo ypes o Pa ona di luens (Lama ck, 1816) (MNHN IK-210-587) and
Pa ona maldi ensis (Ga dine , 1905) (NHMUK 1937.11.17.806) we e aken
a he MNHN in Feb ua y 2014 and a he NHMUK in Sep embe 2024, e-
spec i ely. In Oc obe 2024, he ype se ies o Pa ona a ians Ve ill, 1864,
a pa a ype o P. chi iquiensis (YPM 24153), and syn ypes o Pa ona cla us
(Dana, 1846) (YPM 6249) and Pa ona gigan ea (Ve ill, 1869) (YPM 1679A)
we e s udied and imaged a he Yale Peabody Museum (YPM). An image wi h
scale o specimen YPM IZ.001806.CN (syn ype) was kindly p o ided by E.
Lazo-Wasem. The ype specimens o Pa ona cac us (Fo skål, 1775), ype lo-
cali y Red Sea, and Pa ona explanula a (Lama ck, 1816), ype locali y Indian
Ocean, we e no examined. Ins ead, specimens KAUST SA 205, ma ching he
o iginal desc ip ion o P. cac us, and KAUST SA 231, P. explanula a, om he
Fa asan Banks, Saudi A abian Red Sea, we e imaged a he Red Sea Resea ch
Cen e (RSRC), KAUST a e collec ion in 2013.
Taxonomic accoun
Family Aga iciidae G ay, 1847
Genus Pa ona Lama ck, 1801
Pa ona giannii sp. no .
h ps://zoobank.o g/8396733D-1C70-4F02-BD5C-655C050C3C98
Figs 2–7
Type locali y. Yemen: Shabwa P o ince, Bi Ali, Hyllanyia Island, 13°59.183'N,
48°19.137'E; dep h 5 m; 11 No embe 2008, F. Benzoni leg.
Type ma e ial. Holo ype • 1 colony agmen (10.5 x 5.1 cm, Fig. 2); O iginal
label: “Bi Ali, Hyllanyia Island, Yemen; 13°59.183'N, 48°19.137'E; 11 No . 2008;
F. Benzoni leg.; UNIMIB-C eocean-To al Yemen Co al Biodi e si y exped.; col-
lec ion code BA034; MNHN-IK-2012-14233”.
O he ma e ial. Djibou i • 1 colony agmen (Fig. 7C, pa o he colony
in si u); Maskali Island; 11°42.38'N, 43°9.24'E; 29 Feb. 2020; F. Benzoni leg.;
Dolphin C uise exped.; KAUST DJ403. Yemen • 1 colony agmen (Figs 3A,
H, 5A, D, F, H–L); Aden, Ras An ouk; 12°45.085'N, 45°1.659'E; 8 Ma . 2009; F.
Benzoni and M. Pichon leg.; UNIMIB-C eocean-To al Yemen Co al Biodi e si y
exped.; UNIMIB AD016 • 1 colony agmen (Fig. 6A, whole colony in si u);
Balha ; 13°58.402'N, 48°11.549'E; 22 Ma . 2014; F. Benzoni leg.; C eocean-To-
al Balha LNG Plan moni o ing p og am exped.; UNIMIB BAL252 • 1 colony
agmen (Fig. 5B, C, E, G); Balha ; 13°58.413'N, 48°10.532'E; 24 Ma . 2014; F.
Benzoni leg.; C eocean-To al Balha LNG Plan moni o ing p og am exped.;
UNIMIB BAL253 • 1 colony agmen ; Bi Ali; 13°59.116'N, 48°15.372'E; 16
No . 2008; F. Benzoni leg.; UNIMIB-C eocean-To al Yemen Co al Biodi e si y
exped.; UNIMIB BA010 • 1 colony agmen ; Bi Ali; 13°59.094'N, 48°14.018'E;
16 No . 2008; F. Benzoni leg.; UNIMIB-C eocean-To al Yemen Co al Biodi e si-
y exped.; UNIMIB BA017 • 2 colony agmen s (Fig. 3C); Bi Ali; 13°59.180'N,
48°15.692'E; 19 No . 2008; F. Benzoni leg.; UNIMIB-C eocean-To al Yemen
128
ZooKeys 1260: 123–147 (2025), DOI: 10.3897/zookeys.1260.167263
F ancesca Benzoni: A new Pa ona co al om Indian Ocean ee s
Co al Biodi e si y exped.; UNIMIB BA066 • 2 colony agmen s (Fig. 6G, colony
in si u); Bu um; 14°19.266'N, 48°59.641'E; 18 Ma . 2009; F. Benzoni and M.
Pichon leg.; UNIMIB-C eocean-To al Yemen Co al Biodi e si y exped.; UNIMIB
BU049 • 2 colony agmen s; Al Mukallah; 14°30.923'N, 49°9.254'E; 17 Ma .
2007; F. Benzoni and M. Pichon leg.; UNIMIB-C eocean-To al Yemen Co al Bio-
di e si y exped.; UNIMIB MU085 • 1 colony (Fig. 3B); Al Mukallah; 14°30.696'N,
49°9.360'E; 18 Ma . 2007; F. Benzoni and M. Pichon leg.; UNIMIB-C eocean-To-
al Yemen Co al Biodi e si y exped.; UNIMIB MU128 • 1 colony agmen (Figs
3D, 6C, colony in si u); Soco a Island, Hawla ; 12°40.662'N, 54°4.497'E; 14
Ma . 2010; F. Benzoni and M. Pichon leg.; UNIMIB-C eocean-To al Yemen
Co al Biodi e si y exped.; UNIMIB SO078. Oman • 1 colony agmen (10 x 8
cm); Dhalku ; 16°41.235'N, 53°11.749'E; dep h 9.4 m; 4 Dec. 2022; F. Benzoni
leg.; Oman Biobli z exped.; collec ion code OM0895; UF 17903 • 2 colony ag-
men s (Fig. 6B, colony in si u); Mi ba , Eagles Bay; 16°56.377'N, 054°47.799'E;
dep h 5.4 m; 9 Jan. 2022; F. Benzoni leg.; Oman Biobli z exped.; collec ion
code OM0096; UF 17957 • 1 colony agmen (Fig. 3F); Mi ba , Qinqa i Bay;
17°0.561'N, 55°1.240'E; dep h 6 m; 12 Jan. 2022; F. Benzoni leg.; Oman Biobli z
exped.; collec ion code OM0226; UF 18088 • 1 colony agmen (Fig. 3G); Mus-
ca , Jazi a Al Fahl; 23°40.953'N, 58°30.011'E; dep h 5.9 m; 1 Feb. 2022; F.
Benzoni leg.; Oman Biobli z exped.; collec ion code OM0722; UF 18089 • 1 col-
Figu e 2. Holo ype o Pa ona giannii sp. no . MNHN-IK-2012-14233 A. he whole colony in si u a Hyllanyia Island, Bi
Ali, Yemen, p io o sampling and B. co allum o he sampled agmen . C. Top iew o he co allum showing co alli e
a angemen and he posi ion o some o he co alli e se ies bounda ies (dashed pink lines) ma ked by he p esence o
pa allel adial elemen s unning o e and ac oss hem, and D. side iew showing i s e en su ace gi en by equally high
and la op ma gins o S1 and S2 sep a, and he deep-sea ed ossae. Scale ba s: 1 cm (B); 5 mm (C, D).
129
ZooKeys 1260: 123–147 (2025), DOI: 10.3897/zookeys.1260.167263
F ancesca Benzoni: A new Pa ona co al om Indian Ocean ee s
ony agmen ; Mi ba , Ma io W eck; 16°56.933'N, 054°43.686'E; dep h 10.4
m; 7 Jan. 2022; F. Benzoni leg.; Oman Biobli z exped.; collec ion code OM0017;
UF 18090. Mayo e • 3 colony agmen s (Figs 3E, 5E, pa o he colony in
si u); Îlo M sambo o; 12°38.031'S, 45°1.140'E; 1 Jun. 2010; F. Benzoni leg.;
Ta a Oceans exped.; UNIMIB MY069. Seychelles • 1 colony agmen ; Mahé
Island, Ho se Shoe Ree ; 24 Feb. 2019; F. Benzoni and R. A igoni leg.; Uni e -
si y o Seychelles Ou e Islands co al collec ion acili y exped.; UNISEY SY079
• 1 colony agmen ; same da a as o p eceding; UNISEY SY080 • 1 colony;
Mahé Island, Le Cap; 12 Jan. 1966; B.R. Rosen leg.; NHMUK 1981.3.5.425 •
1 colony; Mahé Island, Baie Te nay; 28 Dec. 1965; B.R. Rosen leg.; NHMUK
1981.3.5.426 • 1 colony (Fig. 4A, B); Mahé Island, No h Eas Poin ; 22 Dec.
1965; B.R. Rosen leg.; NHMUK 1981.3.5.427. S i Lanka • 1 colony (Fig. 4C, D);
Galle; W.C. Ondaa je leg.; NHMUK 1883.3.24.4 • 1 colony; W.C. Ondaa je leg.;
NHMUK 1883.5.23.8. Malaysia • 1 colony (Fig. 4E, F); Pulau Songsong, Wes
Coas , Peninsula Malaysia; C. Be e on leg.; NHMUK 1979.9.24.66.
Desc ip ion. Colonial, co allum enc us ing wi h a ached ma gins, g owing
on he unde lying subs a e and b oadly ollowing i s su ace elie (Figs 2–4, 6,
7); maximum obse ed hickness 1 cm. Co allum su ace smoo h and de oid o
idges, c es s, mon icules o hydnopho ae (Figs 2B–D, 3, 4). Co alli es c owded,
less han a co alli e diame e apa , and hamnas e oid in a angemen (Figs
2B–D, 3, 4, 5A–F). Budding in a en acula , locally leading o he o ma ion o
a iably de eloped co alli es se ies by epea ed incomple e sepa a ion o walls
a e budding. Thei ou line is a iable e en wi hin he same colony anging
om ellip ical o kidney-shaped (Fig. 3H), polygonal (Fig. 3F) o i egula (Figs
2C, 3C). Co alli es 2.4 mm (± 0.1 SE) in a e age maximum diame e and 1.7
mm (± 0.1 SE) in a e age minimum diame e . On a e age, 23 sep a (± 1 SE)
occu pe co alli e (Table 1), he 6 S1 eaching he columella, hei inne ma gin
using wi h i ; hei a e age leng h is 0.9 mm (Table 1). Sep a a anged in h ee
o de s (Fig. 5G, K, L), hose o he i s and second (S1 and S2, espec i ely)
sub-equal in hickness and leng h, all a aining he same heigh (Fig. 5J–L).
S2 may each he columella o emain sligh ly sho e ei he wi h ee ma gin
o occasionally using wi h he columella mo e deeply in he ossa. Thi d o de
sep a (S3) a e always p esen , almos comple e. S3 sep a a e less han ½ o S1
and S2 in leng h (Fig. 5J, K), ne e eaching he columella, and a e hinne and
Table 1. Compa ison o mean (±SE) co alli e and adial elemen s dimensions (mm) and coun s among Pa ona giannii
sp. no ., Pa ona chi iquiensis, and Pa ona a ians. Cha ac e names, measu emen s and coun s o P. chi iquiensis and
P. a ians a e de i ed om Ma é (2003: able 6) e aining he o iginal numbe o decimal places used he ein.
P. giannii sp. no . P. chi iquiensis P. a ians
Mean (±SE) Mean (±SE) Mean (±SE)
Maximum calicula diame e 2.39 (±0.09) 1.85 (±0.05) 1.31 (±0.03)
Minimum calicula diame e 1.66 (±0.10) 1.11 (±0.04) 1.50 (±0.02)
Maximum columella diame e 0.46 (±0.05) 0.34 (±0.03) 0.26 (±0.01)
Minimum columella diame e 0.20 (±0.02) 0.20 (±0.02) 0.16 (±0.01)
Main sep a leng h 0.90 (±0.06) 0.62 (±0.02) 0.47 (±0.01)
Numbe o sep a 23.35 (±0.74) 20.02 (±0.53) 19.86 (±0.62)
Numbe o sep a eaching he columella 6.34 (±0.31) 7.17 (±0.16) 7.88 (±0.15)
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Figu e 3. Top iews o he co alla showing co alli e a angemen in he Pa ona giannii sp. no . specimens collec ed o
his s udy: A. UNIMIB AD016, om Aden, Yemen; B. UNIMIB MU128, om Al Mukallah, Yemen; C. UNIMIB BA066, om Bi
Ali, Yemen; D. UNIMIB SO078, om Soco a Island, Yemen; E. UNIMIB MY069, om Mayo e Island; F. UF 18088 (collec-
ion code OM226), om Mi ba , Oman; G. UF 18089 (collec ion code OM722), om Musca , Oman; H. Same specimen as
in A. Dashed pink lines indica e he posi ion o some o he co alli e se ies bounda ies ma ked by he p esence o pa allel
adial elemen s unning o e and ac oss hem. Scale ba s: 1 cm (A, B); 5 mm (C–H).
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F ancesca Benzoni: A new Pa ona co al om Indian Ocean ee s
sho e in heigh han S1–2 (Fig. 5L). Sep al sides o S1 and S2 o namen ed
wi h sca e ed blun g anules, S3 sides smoo h (Fig. 5J–L). Synap iculae con-
nec ing adial elemen la e al sides ho izon ally can be isible in less densely
calci ied specimens (Fig. 5B, C, E). S1 and S2 uppe ma gins lush, la ened
and unning pa allel o co allum su ace gi ing i an o e all e en appea ance
(Figs 2D, 5C, F). Whe e co alli e se ies occu , adial elemen s un beyond he se-
Figu e 4. Co allum and co alli e mac o-mo phology o he Na u al His o y Museum (UK) specimens o iginally iden i ied as
Pa ona explanula a and he e e-iden i ied as Pa ona giannii sp. no . om locali ies 13–15 in Figu e 1. A. View o he whole
co allum o specimen NHMUK 1981.3.5.427 om Mahé Island, Seychelles, enc us ing a la ge piece o co al ubble also col-
onized by o he ben hic o ganisms including c us ose co alline algae (CCA) and ben hic o amini e a; B. De ail o A showing
co alli e a angemen ; C. Top iew o specimen NHMUK 83.3.24.4 om Galle, S i Lanka; D. De ail o he co allum su ace o
he specimen in C; E. Top iew o specimen NHMUK 1979.9.24.66 om Pulau Songsong, Malaysia, enc us ing a block o co -
al ubble also o e g own by CCA (bo om igh ); F. Close iew o he same specimen in E. Scale ba s: 1 cm (A–E); 5 mm (F).
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F ancesca Benzoni: A new Pa ona co al om Indian Ocean ee s
ea ly-de elopmen s age colonies o hese species a e dis inc om P. giannii
sp. no ., ypically de oid o idges. Mo eo e , co alli es in P. cac us, P. danai,
P. ondi e a and P. mino a e smalle han 2 mm in la ges diame e (Table 2)
while P. giannii sp. no . o ms co alli es wide han 2 mm in diame e .
Among he Pa ona species o ming p edominan ly enc us ing co alla de oid
o idges, bo h Pa ona minu a Wells, 1954 (Fig. 8G) and Pa ona xa i ae Schee
& Pillai, 1974 o m co alli es less han 2 mm in diame e , hence smalle han in
he new species, while P. explanula a (Lama ck, 1816) (Fig. 8H) has co alli es
mo e han 4 mm in diame e , la ge han in he new species. Fu he mo e, in
hese h ee species, co alli es can be mo e han 1 co alli e diame e apa while
hey a e always less han one co alli e apa in P. giannii sp. no .
Table 2. Lis o he co allum and co alli e cha ac e s and hei s a es o Pa ona giannii sp. no . and he cu en ly alid
Pa ona species. Co allum g ow h o m (CGF): massi e (m ), submassi e (sm), columna (cl), ondose ( ), oliose ( l),
enc us ing (en); Co alli e a angemen in se ies (CS): mos ly p esen (yes), pa ially p esen , o only in some co alla bu
no in o he s (pa s), mos ly absen (no); Raised co allum idges (CR) p esen (yes), absen (no); sho (sh), long (lo);
Co alli e dis ance (CDIS): mo e han 1 co alli e diame e apa (>), less han 1 co alli e diame e apa (<); Co alli e wall
e ical de elopmen (CW): aised om he co allum su ace ( ), lush wi h he co allum su ace ( ); Co alli e diame e
(CD): CD < 2 mm (a); 2mm ≤ CD <4 mm (b), CD ≥4 mm (c); co alli e ou line (CO): ci cula (ci), ellip ical (el), polygonal
(po), i egula (i ), indis inc (in); Sep al hickness (ST): equal o sub-equal (eq), unequal (un); Sep al uppe ma gin shape
abo e he co alli e wall (SUM): ounded ( o), la ened ( l); Columella e ical de elopmen in he ossa espec i e o he
sep al uppe ma gin (CDE): high (hi), low (lo), indis inc (in). * = co allum ma gins ee, no a ached o he subs a e. na
= in o ma ion no a ailable om he o iginal desc ip ion and illus a ion, ype no examined. ^ = om he subsequen
desc ip ion by Milne-Edwa ds (1860: 67).
Pa ona species CGF CS CR CDIS CW CD CO ST SUM CDE
P. cac us yes yes > a el-in eq l lo
P. danai yes yes > a ci-el-in un o hi
P. ondi e a^ yes yes na a na un na hi
P. decussa a yes yes < bel-po-in un o- l lo
P. di a ica a yes yes </> bel-in eq o- l hi
P. mino yes yes na a po-in na o in
P. diminu a m pa s no < bel-in un l hi
P. dis inc a m * no no < na in eq l in
P. gigan ea m pa s no > c ci-el un o hi
P. due deni m pa s no < a ci-el eq o lo
P. di luens sm-c-en* no no < - c ci-el un o hi
P. cla us sm-c pa s no < bci-el-i un o hi
P. bipa i a sm-en* pa s no < bci-el-i un o hi
P. enosa m-sm-en pa s yes < bpo-i eq o lo
P. maldi ensis cl, l, en* no no </> bci-el eq o hi
P. explanula a l, en* pa s no > c ci-el un o hi
P. minu a l, en pa s no > - a ci-el-in eq o lo
P. xa i ae en* no no > a ci-el-in un o lo
P. a ians en* pa s yes < a i -in un o- l lo
P. chi iquiensis en pa s yes < a i -in un l lo
P. giannii sp. no . en pa s no < bpo-i -in un l lo
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Figu e 9. Type specimens o Pa ona chi iquiensis, Pa ona decussa a and Pa ona a ians showing he posi ion, a ange-
men , and leng h o he co allum idges, hei uppe ma gin ma ked by anspa en dashed pink lines. A. Top iew o spec-
imen USNM 100871, holo ype o P. chi iquiensis om U a Island, Panama; he anspa en dashed g ey polygon indica es
he posi ion o he su ace po ion shown in B; B. De ail o A showing i egula ly a anged and shaped idges on he co al-
lum, some mo e elonga ed and o he s educed o conical s uc u es de ined by di e en au ho s as “hydnopho ae” (Glynn
e al. 2001), o “conical mon icules” (Ve on and Pichon 1980); C. Close up iew o he co alli es and idges o P. chi iqui-
ensis pa a ype (YPM 24153); D. Side iew o he anas omosing onds in specimen USNM 201, syn ype o P. decussa a
om Fiji, he wo whi e a ows on he sides o i show he o iginal colony g ow h di ec ion om he base o he co allum
o he onds op ma gin; he anspa en dashed g ey polygon indica es he posi ion o he su ace po ion in B; E. De ail
o C showing long and well-de ined, idges g owing along he ond and sepa a ing adjacen co alli e se ies, he ladde -like
a angemen o he adial elemen s unning o e he idges is isible; F. View o specimen YPM IZ.001806.CN, syn ype o
P. a ians, wi h well-de eloped c es -like idges sepa a ing co alli e se ies. Scale ba s: 1 cm (A, B, D–F); 5 mm (C).
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The wo enc us ing Pa ona species ha bea mo e simila i y a he co alli e
le el o he new species desc ibed he e a e Pa ona chi iquiensis Glynn, Ma é &
S emann, 2001 and Pa ona a ians (Ve ill, 1864) (Table 2). Howe e , bo h spe-
cies ha e smalle co alli e diame e (Table 1) based on he mo phome ic anal-
yses pe o med by Ma é (2003). Mo eo e , Pa ona chi iquiensis has S1 = S2 >
S3 and S3 a e o namen ed wi h same shape and size g anules as S1–2 (Glynn
e al. 2001: ig. 2B), while Pa ona giannii sp. no . has smoo h S3 (Fig. 5G–L).
While he new species and P. a ians ha e a simila minimum co alli e diame e ,
co alli es in he la e a e on a e age 1 mm smalle in maximum diame e han
in P. giannii sp. no . co alli es. Consequen ly, P. a ians has sho e S1 adial
elemen s and has on a e age less sep a pe co alli e, bu mo e o hem each
he columella (Table 1). Finally, bo h P. a ians and P. chi iquiensis o m a iably
long idges ha ne e de elop in o onds (Fig. 9A–C, F). In P. a ians, idges
can be s aigh o con o ed, de elop adially o pe pendicula ly o he colony
ma gin (Lewis e al. 2025: ig. 2), and in la ge colonies hei dis ibu ion and de-
elopmen become mo e i egula (Ma hai 1948b). Because o such a iabili y,
hese s uc u es in he genus Pa ona ha e been e e ed o wi h di e en e ms,
namely “c es s” (Ve ill 1864), “ e ical keels” (Ga dine 1898), “ idges” (Ma hai
1948a, 1948b; Ve on and Pichon 1980; La ypo 2014; Lewis e al. 2025), “collin-
es” (Ma hai 1948a, b; Ve on and Pichon 1980; Sheppa d and Sheppa d 1991;
Glynn e al. 2001; Ma é 2003; La ypo 2014), o he mo e elonga ed a ian s,
and “mon icules” (Ma hai 1948a), “conical mon icules” (Ve on and Pichon
1980), and “hydnopho ae” (Glynn e al. 2001) o he sho e ones. Rega dless
o he leng h o o ien a ion, adial elemen s un o e he aised idges in Pa ona
species pe pendicula ly o hei main axis leading o a ypical ladde -like a -
angemen (dashed pink lines in Fig. 9). This becomes less ob ious when a
idge u he de elops in heigh in o a c es o a ond. Al hough P. giannii sp.
no . is de oid o idges on he co allum su ace, mo e o less elonga ed walls
enclosing se ies o co alli es can o m. Thei posi ion is de ec able because he
s ongly al e na ing sep a a e nea ly and egula ly a anged pe pendicula o
hei di ec ion and o m he same ladde -like a angemen (dashed pink lines in
Figs 2, 3, 5) obse ed o e idges in species whe e hese o m.
His o ically, polyp ea u es in scle ac inian co als a e seldom conside ed
in axonomic desc ip ions especially o axa desc ibed be o e sampling and
obse a ions could ake place du ing SCUBA di ing o in aqua ium acili ies.
Some no able excep ions a e ep esen ed by gene a like Euphyllia Dana, 1846
and Fimb iaphyllia Ve on & Pichon, 1980 in which polyp shape and colo a ion
bea mo e in o ma i e cha ac e s han he ac ual skele on (Ve on and Pichon
1980; Luzon e al. 2017; A igoni e al. 2023). In axa like Blas omussa Wells,
1968, Al eopo a Blain ille, 1830, Goniopo a de Blain ille, 1830, Be na dpo a Ki-
ano & Fukami, 2014 and Po i es Link, 1807 polyp cha ac e s ha e been shown
o be in o ma i e o diagnos ic (Benzoni e al. 2014; Ki ano e al. 2014; Te aneo
e al. 2016, 2019). In mos Aga iciidae animal ea u es in i o a e ba ely isible
due o he small size and anspa en na u e o he polyps (Ve on and Pichon
1980). Ne e heless, polyp expansion in he day ime and pale colo a ion o he
en acles a e ypical and diagnos ic ea u es o P. giannii sp. no . h oughou
i s known geog aphic dis ibu ion ange ega dless o he local en i onmen al
condi ions. Ma é (2003: 431) ema ked ha in he Paci ic coas o Panama
in his s udy o P. ondi e a, P. chi iquiensis and P. a ians “none o he h ee
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F ancesca Benzoni: A new Pa ona co al om Indian Ocean ee s
Pa ona species displays expanded polyps du ing he day excep du ing pe iods
o apid wa e low”. Glynn e al. (2001) and Ma é (2003) indica e ha issue
colo a ion in i o is a diagnos ic cha ac e o iden i y P. chi iquiensis in he ield
and o dis inguish i om congene s. Ma é (2003: 438) ema ked ha “da k
polyps and coenosa c, and con as ing b igh whi e o sil e y o al discs and
en acles” cha ac e ized he P. chi iquiensis colonies in Panama. This cha ac e
seems o be conse ed also a o he loca ions. In ac , he same colo a ion was
consis en ly obse ed du ing he co al biodi e si y su eys in he Ma quises
A chipelago, F ench Polynesia, whe e P. chi iquiensis was he mos equen ly
encoun e ed scle ac inian co al ound a 94% o he su eyed si es (Sal a e
al. 2016: ig. 4A). Amongs congene s, P. gigan ea and P. explanula a (Fig. 7C)
usually ha e polyp en acles a iably ex ended du ing he day ime, howe e
hey a e g ey o whi e in he o me and g een o b own in he la e , and bo h
species ha e di e en g ow h o ms and polyp/co alli e size (Table 1). Finally,
he in i o colo a ion o P. dis inc a is unknown (La ypo 2014).
Conclusions
Pa ona giannii sp. no ., a new species o colonial ha d co al in he genus Pa ona,
is desc ibed, highligh ing he diagnos ic alue o bo h skele al and polyp mo -
phology. This species, wi h i s dis inc i e enc us ing g ow h o m, smoo h co al-
lum su ace, and cha ac e is ic day ime polyp expansion and colo a ion, s ands
apa om all cu en ly ecognized Pa ona species. I inhabi s shallow wa e co -
al ee s and ha d g ounds in he Indian Ocean. A de ailed compa a i e analysis,
including mo phome ic da a and compa isons wi h ype and e e ence ma e ial,
unde sco ed he mo phological dis inc i eness o P. giannii sp. no . and p o ided
an illus a ed summa y o he mo phological a iabili y wi hin he genus. Fu u e
esea ch on Pa ona and o he Aga iciidae, should p io i ize expanded species
sampling om di e se geog aphic locali ies including ype locali ies, coupled
wi h phylogenomic analyses. Such e o s will be essen ial o esol e he com-
plex phylogene ic ela ionships wi hin Pa ona and ul ima ely es ablish a obus
and s able axonomic amewo k o his ecologically impo an co al genus.
Acknowledgemen s
The au ho is g a e ul o B.W. Hoeksema, S.D. Cai ns, K. Samimi-Namin, and
an anonymous e iewe o hei use ul sugges ions and e isions which ha e
helped imp o e he manusc ip and Na ahlie Yonow o he inal p oo ing. Field-
wo k o ganiza ion, logis ics, and sampling pe mi s om he ele an au ho i ies
in Yemen we e possible hanks o he suppo o E. Du ieux (C eocean), C.H.
Chaineau (To al SA), R. Hi s , and M. Abdul Aziz (YLNG). I am g a e ul o S. Ba-
sheen (P o essional Di e s Yemen) o echnical and logis ical suppo . I am es-
pecially g a e ul o M. Pichon (Museum o T opical Queensland) o in i ing me
o be pa o he 2005-2014 biodi e si y expedi ion and moni o ing su eys in
Yemen. Sampling in Mayo e was possible hanks o he Ta a Oceans scien i ic
expedi ion and he OCEANS Conso ium. I am g a e ul in pa icula o E. Ka sen-
i (EMBL) and E. Bougois (Ta a Expedi ions) o allowing ee esea ch du ing
he expedi ion, o S. Kandels-Lewis (EMBL), R. T ouble (Fonds Ta a), R. F ied-
e ich (Wo ld Cou ie ) o expedi ion o ganiza ion and logis ics, and o Cap ain
142
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F ancesca Benzoni: A new Pa ona co al om Indian Ocean ee s
H. Bou maud and he Ta a c ew, and o M. O io and J.J. Ke d aon in pa icula .
I am especially indeb ed o L. Bigo (ECOMAR) o his assis ance and suppo
o ieldwo k in Mayo e. Ta a Oceans. I hank he commi men o he ollow-
ing people and addi ional sponso s who made his singula expedi ion possible:
CNRS, EMBL, Genoscope/CEA, VIB, S azione Zoologica An on Doh n, UNIMIB,
ANR (p ojec s POSEIDON/ANR- 09-BLAN-0348, BIOMARKS/ANR-08-BDVA-003,
PROMETHEUS/ ANR-09-GENM-031, and TARA-GIRUS/ANR-09-PCSGENM-218),
EU FP7 (Mic oB3/No.287589), FWO, BIO5, Biosphe e 2, he Veolia En i onmen
Founda ion, Région B e agne, Wo ld Cou ie , Illumina, Cap L’O ien , he EDF Foun-
da ion EDF Di e si e e, FRB, he P ince Albe II de Monaco Founda ion, E. Bou -
gois, he Ta a schoone , and i s cap ain and c ew. Ta a Oceans would no exis
wi hou con inuous suppo om 23 ins i u es (h ps://oceans. a aexpedi ions.
o g). This is con ibu ion numbe 162 based on ma e ial collec ed du ing he
Ta a Oceans Expedi ion 2009–2012. I am g a e ul o D. Ba shish and M. Fine
o he in i a ion o join he 2020 Djibou i Dolphin c uise which hey ully o ga-
nized and suppo ed. The Minis y o he En i onmen and Sus ainable De elop-
men (MEDD) and Minis è e de l’U banisme, de l’En i onnemen e du Tou isme
(MUET) a e acknowledged o hei c i ical assis ance in suppo ing he esea ch
in Djibou i, pa icula ly Mme Bilan Hassan Ismail, h ough coo dina ion o he
esea ch, pe mi s, and sample anspo . Co als in Djibou i we e collec ed unde
he pe mi ing au ho i y o he MUET and MEDD. Field wo k and sampling in he
Seychelles was possible hanks o he Uni e si y o he Seychelles and James
Michel Blue Economy Resea ch Ins i u e. I am g a e ul o K. Hoa eau and R. Wal-
on o o ganizing and suppo ing he co al biodi e si y wo kshop and e e ence
collec ion es ablishmen unde he GoS/UNDP/GEF Ou e Islands p ojec , and M.
Leo a o he guidance in he ield. M.-M. Muzungaile (Minis y o En i onmen ,
Ene gy and Clima e Change) is acknowledged o he suppo wi h he collec ion
pe mi s. Collec ion ieldwo k in Oman was suppo ed by NSF DEB-1457817 o G.
Paulay (Flo ida Museum o Na u al His o y). C. McFadden, M. and E. Clae eboud ,
N. S au , K. Samimi-Namin, S. Wilson, and o he membe s o he Oman Biobli z
expedi ions a e acknowledged o acili a ing ieldwo k, and he En i onmen Au-
ho i y o Oman o pe mi s. I am deeply indeb ed o my colleagues R. A igoni
(SZN), Tullia I. Te aneo (KAUST) o hei suppo , collabo a ion and help wi h
sampling and e e ence collec ion cu a ion on mul iple sampling expedi ions
o e he yea s. Fo museum e e ence collec ion s udy isi s I am since ely g a e-
ul o A. Qua ini and T. Co e (USNM), B.W. Hoeksema and J. Goud (RMNH),
M. Lowe and N. San odomingo (NHMUK), A. Andouche (MNHN), E. Lazo-Wasem
and L. Rojas (YPM), A. Bemis and G. Paulay (UF), and F. Ba eca (KAUST).
Addi ional in o ma ion
Con lic o in e es
The au ho ha e decla ed ha no compe ing in e es s exis .
E hical s a emen
No e hical s a emen was epo ed.
Use o AI
No use o AI was epo ed.
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F ancesca Benzoni: A new Pa ona co al om Indian Ocean ee s
Funding
This s udy was unded by he au ho ’s baseline a KAUST and he Red Sea Resea ch
Cen e FCC/1/1973-49-01.
Au ho con ibu ions
The au ho solely con ibu ed o his wo k.
Au ho ORCIDs
F ancesca Benzoni h ps://o cid.o g/0000-0001-8906-1309
Da a a ailabili y
All o he da a ha suppo he indings o his s udy a e a ailable in he main ex .
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