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Pie e MOISSETTE
Na ional & Kapodis ian Uni e si y o A hens, Facul y o Geology and Geoen i onmen ,
Depa men o His o ical Geology and Paleon ology 15784, A hens (G eece)
and CR2P (CNRS, MNHN, UPMC, So bonne Uni e si é),
Dépa emen O igines e É olu ion, UMR7207, Muséum na ional d’His oi e na u elle,
case pos ale 38, 57 ue Cu ie , F-75231 Pa is cedex 05 (F ance)
[email p o ec ed]
Assimina ANTONARAKOU
Geo ge KONTAKIOTIS
Na ional & Kapodis ian Uni e si y o A hens, Facul y o Geology and Geoen i onmen
Depa men o His o ical Geology & Paleon ology, 15784, A hens (G eece)
[email p o ec ed]
[email p o ec ed]
Jean-Jacques CORNÉE
Géosciences Mon pellie , Uni e si é de Mon pellie
Uni e si é des An illes, CNRS, Poin e à Pi e, Guadeloupe, FWI (F ance)
[email p o ec ed]
Vasileios KARAKITSIOS
Na ional & Kapodis ian Uni e si y o A hens, Facul y o Geology and Geoen i onmen
Depa men o His o ical Geology & Paleon ology, 15784, A hens (G eece)
[email p o ec ed]
Submi ed on 5 Feb ua y 2020 | accep ed on 6 June 2020 | published on 16 Decembe 2021
B yozoan aunas a he To onian-Messinian ansi ion.
A palaeoen i onmen al case s udy om C e e Island,
eas e n Medi e anean
KEY WORDS
B yozoa,
palaeoba hyme y,
palaeoen i onmen ,
palaeogeog aphy,
La e Miocene,
G eece,
C e e,
Aegean Sea.
u n:lsid:zoobank.o g:pub:14A6956D-54AD-48D2-9C5E-BA380EDACAA4
Moisse e P., An ona akou A., Kon akio is G., Co née J.-J. & Ka aki sios V. 2021. — B yozoan aunas a he To onian-
Messinian ansi ion. A palaeoen i onmen al case s udy om C e e Island, eas e n Medi e anean. Geodi e si as 43
(26): 1365-1400. h ps://doi.o g/10.5252/geodi e si as2021 43a26. h p://geodi e si as.com/43/26
ABSTRACT
Fou sedimen a y sec ions we e logged and sampled om uppe To onian-lowe Messinian ou c ops
on he island o C e e (G eece). The collec ed ma e ial yielded abou 60 b yozoan species belonging
o nine di e en colonial mo pho ypes. A ew species a e s enoba hic, indica ing ei he shallow- o
deep-wa e en i onmen s, bu mos o hem a e eu yba hic (wi h ba hyme ic anges ex ending in
some ins ances om he shel down o se e al hund eds o me es). B yozoan communi ies poin o
sea-le el a ia ions modula ed by local ec onics. Deep ci cali o al en i onmen s we e ecognized
a he base o h ee sec ions (whe eas he ou h sec ion s a s wi h in ali o al/shallow ci cali o al
en i onmen s passing o deep ci cali o al. Shallow ba hyal habi a s ollow upwa ds, succeeded in
u n by assemblages indica i e o deep ci cali o al and subsequen ly shallow ci cali o al/in ali o al
dep hs. In e ed ba hyme ic luc ua ions a e diach onous h oughou he island. Shallow-wa e species
ound associa ed, some imes abundan ly, wi h a e deep-wa e aunas in a ew le els, a e in e p e ed
as anspo ed by cu en s. Se e al dysoxic episodes ha e been also de ec ed: likely he esul o wa e
1366
GEODIVERSITAS • 2021 • 43 (26)
Moisse e P. e al.
INTRODUCTION
A he end o he Miocene, du ing he To onian-Messinian
T ansi ion (TMT), sea wa e exchanges be ween he Medi e -
anean and he A lan ic Ocean began o be es ic ed (e.g.,
Kon akio is e al. 2019; Co bí e al. 2020). This es ic ion
culmina ed by he end o he Messinian when he Be ic and
Ri ian co ido s we e closed (Ma ín e al. 2001; Capella e al.
2017; K ijgsman e al. 2018). This led o he well-known
Messinian Salini y C isis (MSC) cha ac e ised by he wide-
sp ead deposi ion o hick e apo i es in deep and/o ma ginal
sub-basins o he en i e Medi e anean Sea (Hsü e al. 1973;
Ryan e al. 1973; Aksu e al. 2018; Haq e al. 2020; Manzi
e al. 2020). An abundan li e a u e has been de o ed o he
iming and he causes o his e en , and a consensus p og es-
si ely eme ged (CIESM 2008; Manzi e al. 2013; Ro e i e al.
2014, 2018, 2020). To explain he la ge olume o e apo i es,
connec ions be ween he A lan ic Ocean and he Medi e -
anean Sea a e conside ed necessa y. A lan ic wa e s en e ed
he Medi e anean ei he h ough he Be ic and Ri ian co -
ido s o e en he Messinian Gib al a Co ido (Achalhi
e al. 2016; K ijgsman e al. 2018). The i s s ep o he MSC
should ha e occu ed in he 5.97-5.60 Ma in e al (Gau ie
e al. 1994; K ijgsman e al. 1999; Manzi e al. 2013). Du -
ing his in e al, he P ima y Lowe Gypsum uni (PLG), he
Te minal Ca bona e Complex (TCC; Es eban 1979), and
coe al basinal sedimen s we e deposi ed. The PLG is opped
by a majo e osional su ace which has been iden i ied bo h
onsho e and o sho e (e.g., Lo i e al. 2011). The second s ep
happened in he 5.60-5.54 Ma in e al; i is cha ac e ised
by he Resedimen ed Lowe Gypsum uni (RLG) and hali e
deposi ion. The hi d s ep o he MSC occu ed in he 5.54-
5.33 Ma in e al. I is ypi ied by he deposi ion o he Uppe
Gypsum uni and sedimen s displaying eshwa e in luences
(Lago Ma e acies), p io o a ma ine e looding a ca 5.33
Ma (base o he Pliocene).
The island o C e e (Aegean Sea, G eece) o e s excellen ly
exposed Miocene o Pliocene ma ine deposi s ha yielded
abundan , di e se, and gene ally well-p ese ed ossil o gan-
isms. Apa om o amini e a, p e ious s udies dealing wi h
ossil communi ies a e ela i ely ew (Agiadi e al. 2017).
Among in e eb a es, b yozoans a e a he common, bu
ha e no been s udied in de ail, e en in he la e Miocene
(Moisse e e al. 1993, 2018; D inia e al. 2009). B yozoans
a e ben hic sessile in e eb a es ha can con iden ly be used
o palaeoen i onmen al econs uc ions (e.g., Schop 1969;
Ha melin 1988; Moisse e 2000; Amini e al. 2004; Taylo
2005). S udies on p esen -day Medi e anean deep-wa e
b yozoans a e sca ce, bu ossil communi ies ha e been he
subjec o much less a en ion (Moisse e& Spjeldnaes 1995;
Rosso 2005; Di Ge onimo e al. 2005; Moisse e e al. 2017).
The pu poses o his pape a e hus: 1) o con ibu e o a be -
e knowledge o la e Miocene b yozoans and 2) use hem o
palaeoen i onmen al econs uc ions.
Fou sec ions (Po amida, Ke amou si, Kapa iana, and Fan-
e omeni) ha e been chosen (Moisse e e al. 2018) in h ee
di e en sedimen a y basins (Chania, He aklion, and Si ia)
in a wes -eas ansec along he Island o C e e, in o de o
column s a i ica ion and enhanced p oduc i i y. These p ocesses we e p obably boos ed by a com-
bina ion o changes in oceanic ci cula ion, clima e, global sea-le el, and he local/ egional ec onics
(in C e e and/o he ma ine ga eways be ween he Medi e anean and he A lan ic).
RÉSUMÉ
Faunes de b yozoai es à la limi e To onien-Messinien. É ude de cas des paléoen i onnemen s de l’île de
C è e, Médi e anée o ien ale.
Qua e coupes li hos a ig aphiques on é é le ées e échan illonnées dans les a leu emen s du To -
onien supé ieu -Messinien in é ieu de l’île de C è e (G èce). Le ma é iel p éle é a li é en i on 60
espèces de b yozoai es appa enan à neu mo pho ypes coloniaux di é en s. Quelques espèces son
s énoba hes, indiquan des en i onnemen s li o aux ou p o onds, mais la plupa son eu yba hes,
a ec des in e alles ba hymé iques s’é endan pa ois depuis le pla eau con inen al jusqu’à plusieu s
cen aines de mè es de p o ondeu . Les communau és de b yozoai es é èlen des a ia ions du ni eau
ma in modulées pa la ec onique locale. Des en i onnemen s du ci cali o al p o ond son econnus
à la base de ois des coupes (mais la qua ième coupe commence pa l’in ali o al/ci cali o al cô ie ,
passan ensui e au ci cali o al p o ond). Des habi a s du ba hyal supé ieu appa aissen ensui e, sui is
d’assemblages indiquan le ci cali o al p o ond, puis le ci cali o al cô ie /in ali o al. Ces luc ua-
ions ba hymé iques son o emen diach ones à a e s l’île. Des espèces li o ales son pa ailleu s
associées, pa ois en g and nomb e, a ec de plus a es aunes p o ondes dans quelques ni eaux où
elles on é é anspo ées pa des cou an s. Plusieu s épisodes dysoxiques on é é égalemen iden i iés,
ésul an p obablemen d’une s a i ica ion c oissan e de la colonne d’eau e d’une augmen a ion de
la p oduc i i é o ganique. Ces p ocessus on é é aisemblablemen a o isés pa une combinaison
de changemen s dans la ci cula ion océanique, le clima , le ni eau ma in global e la ec onique
locale/ égionale (en C è e e /ou au ni eau des co ido s ma ins en e la Médi e anée e l’A lan ique).
MOTS CLÉS
B yozoai es,
paléoba hymé ie,
paléogéog aphie,
paléoen i onnemen ,
Miocène supé ieu ,
G èce,
C è e,
Me Égée.
1367
B yozoan aunas a he To onian-Messinian ansi ion
GEODIVERSITAS • 2021 • 43 (26)
analyse he changes ha occu ed in b yozoan aunas be o e
he Messinian Salini y C isis (MSC). Ou sampling s a s
app oxima ely a 7.58 Ma, c.220 ka a e he To onian
Salini y C isis (TSC a 7.8 Ma; K ijgsman e al. 2000), and
e mina es a 6.72 Ma, c.750 ka be o e he onse o he MSC
(5.97 Ma; Manzi e al. 2013, 2018). The e o e, he s udied
sedimen s documen he TMT, a c ucial ime in e al o he
palaeoceanog aphic his o y o he Medi e anean Sea.
GEOLOGICAL BACKGROUND
A ound he middle/la e Miocene bounda y, he a ea now
occupied by he island o C e e (Fig. 1) became di ided in o a
se ies o basins bounded by aul s and illed mos ly by ma ine
sedimen s ( an Hinsbe gen& Meulenkamp 2006; Zacha iasse
e al. 2008, 2011). Du ing he la e Miocene, ma ls, clayey
limes ones and sap opels deposi ed, la e eplaced by ca bon-
a es, dia omi es and e apo i es (D inia e al. 2004, 2007b;
Ka aki sios e al. 2017a, b; Moisse e e al. 2018; An ona akou
e al. 2019; Kon akio is e al. 2019, 2020).
MATERIAL AND METHODS
The ou sec ions s udied in his pape ha e been logged and
sampled du ing se e al ield wo k campaigns (Figs 2-5) and
some da a ha e been al eady published in Moisse e e al. (2018).
Depending mos ly on ou c opping condi ions, an a e age
sampling in e al o app oxima ely 1m has been used ( ange
TUNISIA
ITALY
SICILY
TURKEY
LIBYA
GREECE
20 km
P e-NeogeneNeogene
He aklion
B
Si ia
Chania
Po amida
Fane omeni
Kapa iana
Ga dos
A
Re hymnon
Ke amou si
200 km
CRETE
35°30'
35°
35°30’
35°
26°
26°25°30’
25°30’25°
25°24°30’
24°30’ 24°
24°
23°30’
23°30’
ig. 1. — A, Si ua ion map o C e e wi hin he eas e n Medi e anean; B, geological ske ch map o he island o C e e (a e K ijgsman e al. 1994), wi h loca ion
o he s udied sec ions.
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GEODIVERSITAS • 2021 • 43 (26)
Moisse e P. e al.
be ween abou 1.6m in Ke amou si and 0.6m in Po amida
sec ion). An o e all numbe o 232 samples we e collec ed,
among which 171 yielded a leas one b yozoan specimen.
Fo each sample, 500 g o d y sedimen we e soaked o
se e al hou s in dilu ed hyd ogen pe oxide and la e washed
unde unning wa e h ough i e mesh sie es (2mm,
1mm, 0.5mm, 0.25mm, and 0.125mm). The esidues
we e o en-d ied and all iden i iable ossils we e picked and
coun ed unde a s e eomic oscope.
The ime amewo k and he s a ig aphic co ela ion
o he s udied sec ions was based on he quali a i e and
semi-quan i a i e analysis o he plank onic o amini e a
iden i ied in he samples (Moisse e e al. 2018; Kon aki-
o is e al. 2019; his s udy). The ch onology o he s udied
sec ions is based on he assemblage-based concep o he
ma ke species, which was ini ially de eloped by Zacha iasse
(1975) and elabo a ed by se e al au ho s (e.g., Hilgen e al.
1995; An ona akou e al. 2007; Ka aki sios e al. 2017a
Loza e al. 2018; Kon akio is e al. 2019; Vasilie e al.
2019; Zacha iasse e al. 2021) in Medi e anean Neogene
sec ions. A o al o c.300 specimens om he >0.125mm
size ac ion o he plank onic o amini e al assemblage
we e collec ed om he washed esidue o ob ain in o ma-
ion abou he abundance o ma ke species as a pe cen age
o he o al plank onic o amini e al auna. The sec ions
we e da ed by using p esence/absence pa e ns and coil-
ing o plank onic o amini e al ma ke species based on
he plank onic o amini e al bios a ig aphy o K ijgsman
e al. (1994, 1995). Ages o he plank onic o amini e al
bioe en s ecognized (Table 1) a e de i ed om he as o-
ch onology based on magne os a ig aphy and as onomical
uning o sedimen a y cycle pa e ns (Lou ens e al. 2004).
Each b yozoan specimen, agmen o whole colony, was
coun ed as one uni in he >0.25mm size ac ion and a
semi-quan i a i e ep esen a ion ( e y a e, a e, equen ,
common, abundan ) was hen p oduced (Figs 2-5). The pal-
aeoen i onmen al analysis was based on he known ecologi-
cal equi emen s o li ing ep esen a i es: colonial g ow h
o ms (zoa ial o ms) and species (Hageman e al. 1998;
Moisse e 2000; Amini e al. 2004; Taylo 2005). Selec ed
well-p ese ed specimens we e ul asonically cleaned and
hen obse ed and pho og aphed wi h a scanning elec on
mic oscope (JEOL JSM-6360) a he Na ional and Kapo-
dis ian Uni e si y o A hens (Depa men o His o ical
Geology-Paleon ology).
The ba hyme ic (bionomical) zona ion used in his pape
ollows ha o Pé ès& Pica d (1964). The dep h limi s a e
es ima es ela ed o ligh and empe a u e le els along la i-
udinal and longi udinal g adien s, also depending on bo -
om subs a e cha ac e is ics: in ali o al (0-40m), shallow
ci cali o al (40-80m), deep ci cali o al (80-200m), and
uppe ba hyal (200-500m). The app oxima e equi alen
e ms used by an Mo kho en e al. (1986) a e espec i ely:
inne shel (inne ne i ic), mid shel (mid ne i ic), ou e
shel (ou e ne i ic), and uppe slope (uppe ba hyal).
RESULTS AND INTERPRETATIONS
Bios a ig aphy
Mos esul s ela ed o he Po amida, Kapa iana and Fan-
e omeni sec ions ha e al eady been published by Moisse e
e al. (2018). New da a o he 84-m hick Ke amou si sec-
ion (He aklion basin; 35°16’34.77”N, 25°01’38.88”E) a e
gi en below and illus a ed in Figu e 3.
The plank onic o amini e al analysis o he s udy sec ions
e ealed he succession o 10 plank onic bioe en s (Table 1)
and he To onian-Messinian bounda y (7.24 Ma). O e all,
all s udy sec ions co e he TMT spanning he ime in e -
al be ween 7.58 and 6.72 Ma. The Fane omeni sec ion is
sligh ly olde han he o he h ee sec ions, wi h he dex al
o sinis al (d/s) coiling change o he Globo o alia sci ula
g oup eco ded a he base o his sec ion (Moisse e e al.
2018). The bios a ig aphic e idence om he op o he
sec ions u he poin s o an age younge han, a leas ,
he Las Occu ence (LO) o G.nicolae a 6.72 Ma, o
Fane omeni and Kapa iana compa ed o Ke amou si and
Po amida sec ions espec i ely (Figs 2-5).
able 1. — Plank onic o amini e al bioe en s iden i ied in he ou s udied sec ions (Moisse e e al. 2018; Kon akio is e al. 2019). Abb e ia ions: LO, Las Oc-
cu ence; LCO, Las Common Occu ence; FO, Fi s Occu ence; FCO, Fi s Common Occu ence.
S age Plank onic Fo amini e al Bioe en s
S a ig aphic le el (m) Ch onos a ig aphy
Po amida
Ke amou si
Kapa iana
Fane omeni
Age (Ma)
Messinian (10) LO G. nicolae – – 37.5 47.7 6.72
(9) FO G. nicolae 34.4 65.5 33.0 41.0 6.83
(8) LCO G. sci ula g oup (sin) – – 31.0 32.4 7.08
(7) FCO G. mio umida g oup 28.8 44.5 – 24.6 7.24
To onian (6) In lux G. mena dii 4 wi hin he ange o G. mena dii 5 26.8 – 15.1 22.2 7.28
(5) Pa acme end G. sci ula g oup (dex) 26.8 – – 22.2 7.28
(4) FO G. mena dii 5 (dex) 7.5 – 13.2 19.4 7.36
(3) LO C. pa ulus 4.3 28.0 7.0 12.8 7.45
(2) LCO G. mena dii 4 (sin) 0.5 21.0 – 8.2 7.51
(1) G. sci ula g oup coiling change D/S – – – 3.5 7.58
1369
B yozoan aunas a he To onian-Messinian ansi ion
GEODIVERSITAS • 2021 • 43 (26)
Desc ip ion o he sec ions
anD o hei B yozoan communi ies
The s udied sedimen s a e mos ly composed o ma ls
and clayey limes ones. Addi ionally, sands ones, sandy
ma ls, bioclas ic limes ones, e uginous conc e ions, and
bi al e shell beds a e common (Moisse e e al. 2018).
The associa ed skele al o ganisms comp ise abundan
o amini e a (ben hic and plank onic), common o a e
bi al es, a e o e y a e os acods, echinoids, p e o-
pods, and e y a e gas opods, scaphopods, decapods,
b achiopods and ishes.
B yozoan emains occu in almos 3/4 o he collec ed
samples, bu hey a e gene ally mino componen s wi h
only a ew agmen s and species in each sample. The
main esul s (including dis ibu ion o he species and
semi-quan i a i e abundances) a e p esen ed in
Figu es 2-5
.
Ma l
Clayey limes one
Sands one
Halimeda limes on
e
Fe uginous bed
Bi al e shells
C isia den icula a
C isia aculea a
En alopho oecia sp.
Exidmonea a lan ica
Sc upocella ia sp.
C isia sp.
Ybselosoecia ypica
Nellia enella
Re epo ella sp.
Smi ina ce ico nis
Tu bicellepo a sp.
Sc upocella ia sc upea
Chape ia annulus
Cheilopo ina campanula a
Dispo ella hispida
Ma ga e a ce eoides
Smi ina cana a ii
S eginopo ella mon ena i
Annec ocyma majo
Canda ec angula a
Cellepo ina cana iensis
Cella ia salico nioides
Bu onella ia mu iella
Calpensia nobilis
Bi lus a sa a ii
Cellepo ina c . siphuncula
Escha ina c . du e ei
Escha oides coccinea
Lagenipo a lep alioides
Mecynoecia delica ula
My iapo a unca a
Onychocella angulosa
Puellina sp.
Schizo heca issa
Tubulipo a sp.
MESSINIAN
–2
10
10
20
–1
20
0
0
39
30
m
10
30 20
40 m
1
30
TORTONIAN
SECT. POTAMIDA A
SECT.
POTAMIDA B
Dysoxia
T anspo ed
elemen s
DysoxiaDysoxia
equen (11-20)
a e (6-10)
e y a e (1-5)
common (21-50)
abundan (>51)
7.28
Ma
7.24
Ma
6.83
Ma
7.24
Ma
deep ci cali o al uppe ba hyal deep ci cali o al
shallow ci c.-
in ali o al
samples
Ba opo a osula
Anguisia e ucosa
O bi ulipo a excen ica
B yoc yp ella o qua a
7.51
Ma
7.45
Ma
7.36
Ma
ig. 2
. — Schema ic sedimen a y log o Po amida composi e sec ion wi h sample loca ion and semi-quan i a i e abundances o b yozoan species.
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GEODIVERSITAS • 2021 • 43 (26)
Moisse e P. e al.
Dysoxia
10
20
1
30
9'
40
50
T anspo ed elemen s
Annec ocyma majo
C isia is ulosa
C isia den icula a
En alopho oecia sp.
Exidmonea a lan ica
C isia sp.
Ybselosoecia ypica
Canda ec angula a
Cellepo ina cana iensis
Chlidonia py i o mis
Nellia enella
Re epo ella sp.
Smi ina ce ico nis
Tubulipo a sp.
Cella ia salico nioides
Bu onella ia mu iella
Calpensia nobilis
Cellepo ina c . siphuncula
Cheilopo ina campanula a
Ma ga e a ce eoides
equen (11-20)
a e (6-10)
e y a e (1-5)
common (21-50)
abundan (>51)
Ma l
Clayey limes one
Lamina ed ma l
Sands one
Bi al e shells
Fe uginous bed
Sandy ma l
Adeonella polys omella
Bi lus a sa a ii
Cella ia is ulosa
Cupulad ia c . cana iensis
Discopo ella eussiana
Escha oides coccinea
Hincksina sp.
Hippopo ina sp.
Mecynoecia delica ula
Sc upocella ia sp.
Onychocella angulosa
Tu bicellepo a sp.
S eginopo ella mon ena i
Schizo heca issa
Pa inella adia a
Sc upocella ia sc upea
80 m
70
60
50
40
30
20
10
0
MESSINIANTORTONIAN
C isia aculea a
7.24
Ma
uppe ba hyal deep ci c.
in ali o al
shallow ci cali o al-in ali o al deep ci cali o al
lagoonal
shallow ci c.-
samples
Dysoxia
7.51
Ma
7.45
Ma
7.24
Ma
6.83
Ma
Gemellipo a ebu nea
Ba opo a osula
Te ia i egula is
Anguisia e ucosa
O bi ulipo a excen ica
ig. 3
. — Schema ic sedimen a y log o Ke amou si sec ion wi h sample loca ion and semi-quan i a i e abundances o b yozoan species.
1371
B yozoan aunas a he To onian-Messinian ansi ion
GEODIVERSITAS • 2021 • 43 (26)
Gemellipo a ebu nea
Ba opo a osula
Te ia i egula is
Anguisia e ucosa
Dysoxia
26
PLIOCENE
0
10
20
30
40
50 m
01
010
020
030
036
1
10
12'
14'
20
MESSINIANTORTONIAN
T anspo ed elemen
s
Annec ocyma majo
C isia is ulosa
C isia den icula a
En alopho oecia sp.
Exidmonea a lan ica
Sc upocella ia sp.
C isia sp.
Ybselosoecia ypica
Canda ec angula a
Cellepo ina c . siphuncula
Chlidonia py i o mis
Re epo ella sp.
Smi ina ce ico nis
Tu bicellepo a sp.
Cella ia salico nioides
Sc upocella ia sc upea
Bu onella ia mu iella
Calpensia nobilis
Cheilopo ina campanula a
Dispo ella hispida
Halysisis c . diaphana
Ma ga e a ce eoides
Smi ina cana a ii
S eginopo ella mon ena i
Ma l
Clayey limes one
Lamina ed ma l
Sands one
Bioclas ic limes one
Bi al e shells
Ra inemen su ace
Dysoxia
equen (11-20)
a e (6-10)
e y a e (1-5)
common (21-50)
abundan (>51)
C isia aculea a
Ma
deep ci c.
uppe ba hyal deep ci cali o al
shallow ci cali o al-in ali o al
SECT. KAPARIANA E (KAP)
SECT. KAPARIANA W (CAP)
samples
7.24
7.24
Ma
7.08
Ma
6.83
Ma
6.72
Ma
7.28
Ma
7.36
Ma
7.45
Ma
ig. 4
. — Schema ic sedimen a y log o Kapa iana composi e sec ion wi h sample loca ion and semi-quan i a i e abundances o b yozoan species.
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GEODIVERSITAS • 2021 • 43 (26)
Moisse e P. e al.
ema ks
The gonozooid, cha ac e is ic o he genus, consis ing o a
simple elonga ed chambe wi h a e minal ubula ooecios ome
(J.-G. Ha melin, pe sonal communica ion), was no obse ed.
The enc us ing basis is isible in some o he s udied speci-
mens (Fig. 6B). The species c ea ed by Ne iani (1895) om
he Pliocene/Pleis ocene o no he n I aly, A.jullieni mos
p obably co esponds o he enc us ing basis o A. e ucosa.
Howe e , he homonymous A.jullieni desc ibed by Os o sky
(1998) om he p esen -day An a c ic is a di e en species.
Pe is ome diame e and leng h a e smalle in A. e ucosa and
he p ominen e ucae associa ed wi h he pseudopo es a e
absen in A.jullieni.
Family uBulipo iDae Johns on, 1838
ype genus.— Exidmonea Da id, Monge eau& Pouye , 1972, by
subsequen designa ion o Monge eau (1969: 212).
Genus Exidmonea Da id, Monge eau& Pouye , 1972
Exidmonea a lan ica (Fo bes in Johns on, 1847)
(Fig. 7C, D)
Idmonea a lan ica Fo bes in Johns on, 1847: 278, pl. 48, ig. 3.—
Buge 1966: 5, pl. B, igs 1-2.
Idmid onea a lan ica – Ha melin 1976: 182, pl. 32, igs 1-11.—
Vá a 1983: 73, pl. 1, igs 1-4.— Zabala 1986: 658, ex - igs
235a-d; pl. 28, ig. E.— Zabala& Maluque 1988: 174, ex - igs
569-572; pl. 33, ig. A.— El Hajjaji 1992: 44, pl. 2, ig. 11.—
Haddadi-Hamdane 1996: p. 45, pl. 1, ig. 8, pl. 2, ig. 4.
Exidmonea a lan ica – Monge eau 1970: 30, pl. 1, ig. 1; pl. 2, igs
1, 5.— Da id e al. 1972: 84.— Moisse e 1988: 48, pl. 6, igs
6-7.— Pouye & Moisse e 1992: 24, pl. 1, igs 7-8.—Moisse e
e al. 1993: 84, igs 3i-j.— Moisse e& Spjeldnaes 1995: 780, pl.
1, ig. 11.
Exidmonea i o is – Haywa d& McKinney 2002: 104, ig. 49A-H, 50.
o
ccu ence
.— Eocene: I aly, Aus ia, Hunga y, Romania (Zágo šek
2003). Oligocene: Ge many, I aly (B aga 2008). Ea ly Miocene:
F ance, I aly, Egyp (El Sa o i& El-So ogy 1999). Middle Miocene:
Aus ia, Hunga y, Poland, F ance, Libya (Vá a 1983). La e Mio-
cene: Alge ia (Moisse e 1988), Mo occo (El Hajjaji 1992), Tunisia
(Moisse e 1997), C e e. Pliocene: F ance, Sicily (Pouye & Moisse e
1992), Alge ia (Haddadi-Hamdane 1996), Tunisia. Pleis ocene: Sic-
ily (Di Ge onimo e al. 1997; Rosso 2005), Rhodes (Moisse e&
Spjeldnaes 1995).
This lis o ossil eco ds conce ns Idmonea-Idmid onea-Exidmonea-
like species, i.e. a g oup o species wi h he same g ow h o m
wi hou conside a ion o he species-speci ic mo phological c i e ia
which canno be, in mos cases, p ese ed in ossils (J.-G. Ha melin,
pe sonal communica ion). Recen : Eas e n A lan ic (No h Sea o
Angola) and Medi e anean, a dep hs anging om 10 o 850m.
In he Medi e anean, his species is pa icula ly abundan be ween
40 and 100m (Ha melin 1976).
Desc ip ion
Vincula ii o m colony. F on al su ace showing zoecial ubes
al e na ing in wo se ies o 4-5 ubes. Do sal la ened o
sligh ly conca e, o namen ed by hin in e ed U-shaped
g ow h lines. Gonozooid (b ood-chambe ) elonga e, si ua ed
along he axis o a b anch, be ween pe is omes.
ema ks
The oecios ome was no obse ed.
Family e iiDae Canu& Bassle , 1920
Genus Te ia Jullien, 1882
Te ia i egula is (Meneghini, 1844)
Idmonea i egula is Meneghini, 1845: 128.
Te ia i egula is – Malecki 1963: 71, ex - ig. 30; pl. 3, ig. 11.—
Vá a 1975: p. 523, pl. 2, igs 3-4.— Ha melin 1976: 163, pl. 26,
igs 1-11.— Ha melin& d’Hond 1982: 9, pl. 3, ig. 4.— Zabala
1986: 656, ex - ig. 234.— Moisse e 1988: 49, pl. 6, igs 10-12.—
Zabala& Maluque 1988: 174, ex - igs 563-568.— Pouye &
Moisse e 1992: 25, pl. 1, igs 11-12.— El Hajjaji 1992: p. 48, pl.
2, ig. 10.— Moisse e e al. 1993: 85, ig. 3h, l.— Moisse e&
Spjeldnaes 1995: 782, pl. 1, igs 9, 12.— Haddadi-Hamdane 1996:
46, pl. 1, ig. 10, pl. 2, igs 3, 8.
occu ence.— Eocene: Poland, Hunga y, Spain (Malecki 1963).
Oligocene: Ge many (Malecki 1963). Ea ly Miocene: F ance (Buge
1957). Middle Miocene: Aus ia, Hunga y (Moisse e e al. 2006),
F ance. La e Miocene: Alge ia (Moisse e 1988), Mo occo (El
Hajjaji 1992), C e e (Moisse e e al. 1993). Pliocene: Ne he lands
(Lagaaij 1952), Belgium, F ance, Alge ia, Sicily (Pouye & Mois-
se e 1992). Pleis ocene: Rhodes (Moisse e& Spjeldnaes 1995),
Ka pa hos (Moisse e e al. 2017). Recen . Eas e n A lan ic (down
o a dep h o 2650m in he Bay o Biscay), Medi e anean (60-
300m), Paci ic, Indian Ocean.
ema ks
Only a ew agmen s ha e been ound in wo samples. This
easily ecognizable species is no illus a ed he e because o
he ela i ely poo p ese a ion s a e o he a e eco e ed
specimens.
Family Diape oeciiDae Canu, 1918
Genus Ybselosoecia Canu& Lecoin e, 1933
Ybselosoecia ypica (Manzoni, 1878)
(Fig. 7E-G)
Filispa sa ypica Manzoni, 1878: 10, pl. 8, ig. 30.— Canu 1909:
115, pl. 14, igs 25-26.
Ybselosoecia ypica – Buge 1956: 13, pl. 1, igs 3-4; pl. 2, igs 3-4.—
Malecki 1963: 76, ig. 33; pl. 5, ig. 1.— Monge eau 1965: 317,
ig. 1.— Vá a 1974: 362, pl. 2, igs 9-10.— Vá a 1984: 226, pl.
1, igs 2-3.— Moisse e 1988: 56, pl. 7, igs 9, 13; 1997: 188, pl. 1,
ig. 6.— El Hajjaji 1992: 56, pl. 2, igs 2-3.— Pouye & Moisse e
1992: 28, pl. 2, ig. 4.— Moisse e e al. 1993: 87.— Moisse e&
Spjeldnaes 1995: 784, pl. 2, ig.4.— Haddadi-Hamdane 1996:
51, pl. 1, igs 11-12; pl. 2, ig. 11.— Pouye 1997: 26, pl. 1, igs
1-4.— Zágo šek 2003: 119, pl. 4, igs 5-6.— Zágo šek 2010a: 31,
pl. 15, igs 1-5.— Zágo šek e al. 2017: 115, ig. 2J.
1379
B yozoan aunas a he To onian-Messinian ansi ion
GEODIVERSITAS • 2021 • 43 (26)
occu ence.— Eocene: F ance, I aly, Ge many, Aus ia, Romania,
Poland (Malecki 1963; Zágo šek 2003). Oligocene: F ance (Buge
1964). Oligo-Miocene: I an (Zágo šek e al. 2017). Ea ly Miocene:
F ance (Monge eau 1965). Middle Miocene: F ance, Aus ia, Hunga y,
Romania, Czech Republic, Poland (Vá a 1984; Moisse e e al. 2006).
La e Miocene: Mo occo, Alge ia (Moisse e 1988), Tunisia (Moisse e
1997), C e e (Moisse e e al. 1993). Pliocene: Ne he lands, F ance,
Sicily (Pouye & Moisse e 1992), Alge ia (Haddadi-Hamdane 1996),
Tunisia (Buge 1956). Pleis ocene: Rhodes (Moisse e& Spjeldnaes 1995).
This ossil species has mos ly been eco ded om he Medi e anean
and Pa a e hys basins. I has been eco e ed in a ew ins ances oge he
wi h deep-wa e b yozoan aunas: Messinian o wes e n Alge ia (Mois-
se e 2000) and Pleis ocene o Rhodes (Moisse e& Spjeldnaes 1995).
Desc ip ion
Vincula ii o m colony. Dicho omous b anches mos ly la -
ened. F on al smoo h, zoecial ubes indis inc , quincuncially
a anged and isola ed, wi h well-de eloped sub-ci cula pe i-
s omes. Do sal sligh ly con ex, wi h hin ans e sal g ow h
A
DEF
BC
ig. 8. — A-C, C isia aculea a Hassall, 1841; A, in e node, on al iew, POTB13: AMPG(IV) 2828a; B, in e node, do sal iew, POTB13 2828b: AMPG(IV); C, in e -
node wi h gonozooid, KAP22: AMPG(IV) 3339. D-F, C isia den icula a (Lama ck, 1816); D, in e node, on al iew, POTB13: AMPG(IV) 2829a; E, in e node do sal
iew, POTB13: AMPG(IV) 2829b; F, gonozooid, POTB13: AMPG(IV) 2829c. Scale ba s: A, 500 µm; B-F, 200 µm.
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GEODIVERSITAS • 2021 • 43 (26)
Moisse e P. e al.
lines and pe o a ed by small pseudopo es. Gonozooid enla ged,
also pe o a ed by small pseudopo es, occupying he whole
wid h o a b anch and encompassing up o en zoecial ubes.
Ooecios ome wi h la ed, ans e se and con olu ed im.
Subo de ARTICULINA Busk, 1859
Family c isiiDae Johns on, 1838
Genus C isia Lamou oux, 1812
C isia aculea a Hassall, 1841
(Fig. 8A-C)
C isia aculea a Hassall, 1841: 170, pl. 7, igs 3-4.— Ha me 1891:
132, pl. 12, ig. 4.— Haywa d& Ryland 1985: 50, ig. 14.—
Ha melin1990: 1602, igs 2, 3-6.
occu ence.— Recen : Eas e n A lan ic (50-60m in he No h
Sea, much deepe in No way, he Bay o Biscay and Mo occo: 135-
1000m) and Medi e anean (115-480m).
Desc ip ion
Cella ii o m colony. In e nodes sho (8-12 zooids). The i s
ami ica ion a ises om he hi d zooid. Pseudopo es elon-
ga e and ela i ely a e. Gonozooid py i o m wi h a disc e e
ooecios ome si ua ed nea he base o he ollowing zooid.
ema ks
This species is poo ly known and has a ely been illus a ed.
I has o en been conside ed as a a ie y o C.ebu nea (Lin-
naeus) o which i esembles. Ha melin (1990) dis inguished
a no he n (A lan ic) and a sou he n o m (A lan ic coas o
Mo occo, Medi e anean). The main di e ences a e a highe
numbe o zooids (9-12) pe in e node and a mi e-shaped
gonozooid in he sou he n o m.
C isia den icula a (Lama ck, 1816)
(Fig. 8D-F)
Cella ia den icula a Lama ck, 1816: 137.
C isia den icula a – Vá a 1975: 523, pl. 1, ig. 1.— Haywa d&
Ryland 1985: 54, ig. 16.— Poluzzi e al. 1988: 64, igs 15b-d.—
Poluzzi& Rosso 1988: 99, pl. 1, ig. 1.— Zabala& Maluque 1988:
165, ex - igs 480-482, pl. 29B.— Bobies 1958: 153, pl. 13, igs
5-7.— Ma copoulou-Diacan oni& Wues 1999: 552, pl. 2, ig. 6.
occu ence.— Middle Miocene: Aus ia, Hunga y, Czech Republic
(Bobies 1958; Vá a 1975). La e Miocene: Sa dinia, Sicily (Moisse e
e al. 2002). Pliocene: UK, Ne he lands (Lagaaij 1952), Belgium,
C e e (Ma copoulou-Diacan oni& Wues 1999). Pleis ocene: Sicily
(Rosso 1987). Recen : wes e n and eas e n A lan ic (Canada o Gul
o Mexico, Madei a, Azo es, No way o Ghana), Medi e anean. This
species li es a dep hs be ween 0 and 100m, bu i is associa ed wi h
deep-wa e -co als in No way (Sula Ree : 275-295m; Mo ensen&
Fosså 2006) and in he Bay o Biscay (Cal e 1896).
ema ks
C.den icula a closely esembles he ossil species C.hoe nesi
Reuss, which howe e has a g ea e numbe o zooids pe
in e node (14-16).
Class GYMNOLAEMATA Allman, 1856
O de CHEILOSTOMATIDA Busk, 1852
Subo de FLUSTRINA Smi , 1868
Supe amily callopo oiDea No man, 1903
Family QuaD icella iiDae Go don, 1984
Genus Nellia Busk, 1852
Nellia enella (Lama ck, 1816)
(Fig. 9E)
Cella ia enella Lama ck, 1816: 135.
Nellia ocula a – Busk 1852: 18, pl. 64, ig. 6; pl. 65 (bis), ig. 4.—
Moisse e e al. 1993: 98, ig. 5i.— Moisse e 1997: 192, pl. 2, igs 5-6.
Nellia enella – Ziko e al. 2016: 18, pl. 2, ig. 6.
occu ence.— Eocene: UK, F ance, I aly, Ge many, Romania
(B aga 2008). Oligocene: USA, Mal a, Libya (El Sa o i& Mu ah
2007), UAE (B aga& Bah 2003). Ea ly Miocene: USA (Di Ma -
ino e al. 2017), Dominican Republic, F ance, Po ugal, Egyp
(El Sa o i& El-So ogy 1999). Middle Miocene: Jamaica, Hunga y
(Moisse e e al. 2006), Aus ia, F ance, Po ugal, Egyp (Ziko e al.
2016). La e Miocene: Mo occo (El Hajjaji 1992), Alge ia (Moisse e
1988), C e e, Tu key, Tunisia (Moisse e 1997). Pliocene: Cos a
Rica, Panama, Dominican Republic, Jamaica (Taylo & Fos e
1998; Chee ham e al. 1999). Pleis ocene: Rhodes (PM, pe s. obs.).
Recen : eas e n and wes e n A lan ic, Indian Ocean, Paci ic. This
species is widely dis ibu ed in opical o sub opical wa e s a ound
he wo ld. I li es a dep hs be ween 0 and 250m (F ansen 1986).
Desc ip ion
Colony cella ii o m. In e nodes s aigh wi h quad angula
sec ion and elonga e zooids sepa a ed by shallow g oo es.
Opesia o al, occupying he g ea es pa o he zooidal on al
wall. Gymnocys and c yp ocys well-de eloped p oximally.
Two small a icula ia occu on he p oximal pa o he gym-
nocys . O icells no obse ed.
ema ks
Nellia ocula a Busk is now ega ded as a junio synonym o
N. enella (Lama ck). Howe e , hei e y b oad ecological,
geog aphical and s a ig aphical eco ds may indica e ei he a
’’li ing ossil’’ species o a “c yp ic species complex” (Wins on
e al. 2014; Di Ma ino e al. 2017).
Family cupulaD iiDae Lagaaij, 1952
Genus Cupulad ia Canu& Bassle , 1919
Cupulad ia c . cana iensis (Busk, 1859)
(Fig. 9A, B)
Cupula ia cana iensis Busk, 1859: 66, pl. 23, igs 6-9.
Cupulad ia cana iensis – Manzoni 1869, p. 26, pl. 2, ig. 17; 1877: 72,
pl. 17, igs 56a-c.— Cipolla 1921: 31, pl. 2, igs 22-24.— Lagaaij 1952:
33, pl. 2, igs 1a-b.— Buge 1957: 139, pl. 9, ig. 5.— Annoscia 1963:
225, pl. 9, ig. 1; pl. 10, ig. 1; pl. 11, igs 1a-b; pl. 12, igs 1a-b.—
Cook 1965: 197, ex - igs 1a- , pl. 1, ig. 1; pl. 3, ig. 4.— P enan &
Bobin 1966: 307, igs 101-102.— Baluk& Radwansky 1984: 21, pl.
1, igs 1-4; pl. 8, igs 1-4.— Zabala& Maluque 1988: 89, ig. 112-
1381
B yozoan aunas a he To onian-Messinian ansi ion
GEODIVERSITAS • 2021 • 43 (26)
A
C
EFGH
D
B
ig. 9. — A, B, Cupulad ia c . cana iensis (Busk, 1859); A, on al iew o a whole colony, KER39: AMPG(IV) 3078a; B, do sal iew o a whole colony, KER39:
AMPG(IV) 3078b; C, D, Discopo ella eussiana (Manzoni, 1869); C, on al iew o a whole colony, KER30: AMPG(IV) 3102; D, do sal iew o a whole colony, KER30:
AMPG(IV) 3101; E, Nellia enella (Lama ck, 1816), de ail o an in e node showing wo zooids in on al iew, KER17: AMPG(IV) 3150a; F-H, Canda ec angula a
Udin, 1964; F, do sal iew o an in e node agmen , FAN35: AMPG(IV) 3506a; G, on al iew o an in e node agmen , FAN35: AMPG(IV) 3506b; H, De ail o he
same agmen , FAN35: AMPG(IV) 3506b. Scale ba s: A-D, 1 mm; E, H, 100 µm; F-G, 200 µm.
1382
GEODIVERSITAS • 2021 • 43 (26)
Moisse e P. e al.
114.— Pouye & Moisse e 1992: 33, pl. 3, igs 4-5.— Moisse e
e al. 1993: 92, igs 4g-i.— Haddadi-Hamdane 1996: 65, pl. 5, igs
1, 4.— Ma copoulou-Diacan oni& Wues 1999: 555, pl. 3, ig. 1.
occu ence.— Middle Miocene: F ance, Spain, Aus ia, Poland
(Baluk& Radwansky 1984). La e Miocene: Ge many, I aly, C e e
(Moisse e e al. 1993). Pliocene: UK, Ne he lands, Sicily (Pouye &
Moisse e 1992), Alge ia (Haddadi-Hamdane 1996), C e e (Ma co-
poulou-Diacan oni& Wues 1999). Pleis ocene: Sicily (Rosso 1987),
Rhodes (Moisse e 2012). Recen : eas e n Paci ic (Ecuado o no he n
Mexico), wes e n and eas e n A lan ic (B azil, Ca ibbean, Gabon,
Azo es, Madei a and Cana y islands o sou he n Po ugal), sou he n
Medi e anean (Rosso& Di Ma ino 2016). This wa m-wa e species
li es on sandy, mo e o less muddy bo oms a dep hs be ween 50
and 300m (P enan & Bobin 1966). Bu i has also been ound in
much shallowe (5-50m) and much deepe wa e s (down o 860m
in he Sa gasso Sea; Lagaaij 1963). Following Cadée (1979, 1981),
C.cana iensis seems howe e mos ly a wes e n A lan ic species. Reco ds
wi h his name mus consequen ly be ca e ully checked, elimina ing
possible misiden i ica ions (A. Rosso, pe sonal communica ion).
ema ks
Due o possible con usion wi h wo o he species o he same
genus (C.bipo osa (Canu& Bassle , 1923) and C. indobonensis
Baluk& Radwanski, 1984) a sys ema ic e ision is needed.
Cadée (1979) also c ea ed a new subspecies, Cupulad ia
cana iensis ca e nosa, o Mio-Pliocene Eu opean specimens
wi h in e media e cha ac e is ics be ween C.bipo osa and
C.cana iensis (Cadée 1979, 1981).
Genus Discopo ella d’O bigny, 1852
Discopo ella eussiana (Manzoni, 1869)
(Fig. 9C, D)
Cupula ia eussiana Manzoni, 1869: 27, pl. 2, ig. 19.
Cupulad ia eussiana – Annoscia 1963: 226, pl. 9, ig. 2; pl. 10, ig.
2; pl. 13, ig. 1; pl. 14, ig. 1a-b.— P enan & Bobin 1966: 316,
ig. 104.— Reguan 1969: 38, igs 1-3.
Reussi ella eussiana – Baluk& Radwansky 1984: 27, pl. 11, igs 1-2.
Discopo ella eussiana – Cook 1965: 219, ex - ig. 2 ; pl. 3, ig. 1.—
Poluzzi 1975: 52, pl. 18, igs 1a-m, 2, 3.— Pouye & Moisse e
1992: 44, pl. 5, ig. 4-6.— Moisse e e al. 1993: 96, igs 5c-d.—
Haddadi-Hamdane 1996: 66, pl. 5, igs 2, 3, 8.
occu ence.— Middle Miocene: Aus ia (Baluk& Radwansky
1984). La e Miocene: I aly, C e e (Moisse e e al. 1993). Pliocene:
Alge ia (Haddadi-Hamdane 1996), Sicily (Pouye & Moisse e
1992). Pleis ocene: Spain (Reguan 1969), Sicily, Umb ia, cen al
I aly (Bizza i e al. 2015). Recen : he p esen -day occu ence o
his species is doub ul (Cook 1965; P enan & Bobin 1966). The
nea es li ing species, R.doma, li es in opical o sub opical wa e s
o he A lan ic and Medi e anean, on sandy o muddy bo oms
a dep hs o 10 o 370m (Cook 1965; P enan & Bobin 1966).
Desc ip ion
Lunuli i o m colony. F on al su ace wi h al e na ing adial
se ies o homboidal zooids, each bea ing a dis al ib aculum.
The six c yp ocys al den icles do no use in he middle o he
opesia. The apical zone o he colony shows zooids co e ed
by a calci ied lamina. Basal su ace dep essed in he cen e,
displaying ma ked adial g oo es and s ong ube cles.
ema ks
D. eussiana esembles e y much Reussi ella doma (d’O bigny,
1851) o which i has been con used (P enan & Bobin 1966).
Acco ding o Cook (1965), he un used c yp ocys al den icles
a e especially cha ac e is ic o he wes e n A ican Reussi ella
owenii (G ay, 1828).
Supe amily BuguloiDea G ay, 1848
Family canDiDae d’O bigny, 1851
Genus Canda Lamou oux, 1816
Canda ec angula a Udin, 1964
(Fig. 9F-H)
Canda ec angula a Udin, 1964: 393, pl. 1, ig. 3.— Vá a 1979:
599, pl. 1, igs a, d, g; 1980: 58, pl. 2, igs 5-6.— Schmid 1989:
25, pl. 6, igs 1-5, 8.— Moisse e 1997: 193, pl. 2, igs 11-12.
o
ccu ence
.— Middle Miocene: Aus ia (Schmid 1989), Hunga y
(Moisse e e al. 2007). La e Miocene: Tunisia (Moisse e 1997).
Desc ip ion
Colony incula ii o m. F on al showing ec angula zooids
o ganized in wo al e na ing se ies wi h p ominen dis al and
la e al edges. C yp ocys conca e, p oximally well-de eloped and
hus esul ing in an elonga e V-shaped opesia. Do sal display-
ing enla ged iangula ib acula ia wi h long se al g oo es and
la ge adicula po e. F on al a icula ia and o icells no obse ed.
ema ks
The closeness wi h he Recen Canda e i o mis Pou alès
has been emphasized since he c ea ion o C. ec angula a.
A e ision o bo h ossil and Recen ma e ial o he genus
Canda is desi able.
Genus Sc upocella ia an Beneden, 1845
Sc upocella ia c . ellip ica (Reuss, 1847)
(Fig. 10A-C)
Bac idium ellip icum Reuss, 1847: 56, pl. 9, ig. 7-8.
Sc upocella ia ellip ica – Reuss 1874: 148, pl. 11, igs 1-9.— Mois-
se e 1988: 106, pl. 16, igs 5, 8.— Schmid 1989: 23, pl. 5, igs
1-7.— Zágo šek 2010a: 46, pl. 52, igs 1-6.
Sc upocella ia sp. – Be ning 2006: 32, igs 27-29.
occu ence.— Middle Miocene: Aus ia (Schmid 1989), Czech
Republic (Zágo šek 2010a). La e Miocene: Alge ia (Moisse e 1988),
Spain (Be ning 2006).
Desc ip ion
Cella ii o m colony. Each segmen consis s o wo al e na ing
se ies o abou i e zooids. Gymnocys smoo h. O al opesia
1383
B yozoan aunas a he To onian-Messinian ansi ion
GEODIVERSITAS • 2021 • 43 (26)
occupying app oxima ely one hal o he zooecial leng h. Two
la ge sep ula a e disce nible in he dis al pa o he opesial
ma gin. Fou spine bases occu on he ou e dis al pa o he
opesia and h ee on he inne angle, accompanied by one sligh ly
la ge scu al spine (scu um a ely p ese ed, co e ing almos
hal he opesia). No on al a icula ium. La e al a icula ia
iangula and well de eloped. Do sal su ace showing small
iangula ib acula ia and adicula po es. No obse ed o icells.
ema ks
As no ed by Schmid (1989) and Be ning (2006), many ossil
Sc upocella ia specimens ha e been mis akenly assigned o
S.ellip ica. Pa ly due o a equen poo s a e o p ese a ion
his has been accompanied by insu icien desc ip ion and
illus a ion, no ably conce ning he p esence and numbe o
spine bases. Spines a e no alluded o in Reuss (1847), bu
men ioned and pa ly illus a ed in Reuss (1874). Al hough
no epo ed, spines a e isible on some specimens o he la e
Miocene o Alge ia (Moisse e 1988: pl. 16, ig. 8) on. Scu-
um and spines (a leas 5) a e clea ly obse able on he mid-
dle Miocene ma e ial (Schmid 1989: pl. 5, ig. 2 and ig. 4,
espec i ely). Six spines (+scu um) a e epo ed and illus a ed
by Be ning (2006). Zágo šek (2010b) desc ibes “spines o en
a anged in 4-5 pai s”. A e ision o Neogene Medi e anean/
Pa a e hys Sc upocella ia is hus s ongly necessa y, no ably o
con i m he a ibu ion o specimens wi h se en o al spines o
S.ellip ica. A new app oach o he axonomy o he polyphyle ic
genus Sc upocella ia has been p esen ed by Viei a e al. (2014).
Sc upocella ia sc upea Busk, 1852
(Fig. 10D-E)
Sc upocella ia sc upea Busk, 1852: 83, pl. 9, igs 11-12.— Hincks
1880: 50, pl. 7, igs 11-14.— P enan & Bobin 1966: 432, ex - ig.
143.— Ryland& Haywa d 1977: 138, ex - ig. 66.— Zabala&
Maluque 1988: 99, ex - igs 167-168.— Moisse e& Spjeldnaes
1995: 788, pl. 3, igs 2-4.— Haywa d& McKinney 2002: 27, ig.
11A-E.
occu ence.— Pliocene: Calab ia (Ne iani 1900). Pleis ocene:
Calab ia (Di Ge onimo e al. 1997), Rhodes (Moisse e& Spjeldnaes
1995). Recen : eas e n A lan ic and Medi e anean om he su ace
o 150m (and down o 500m in he Gul o Gascony: Jullien&
Cal e 1903).
Desc ip ion
Cella ii o m colony. In e nodes comp ising wo al e na ing se ies
o zooids. Gymnocys smoo h. O al opesia occupying sligh ly
mo e han hal o he zooecial leng h wi h wo la ge sep ula
in he dis al pa o he opesial ma gin Fi e spine bases on he
dis al im ( h ee on he ou e edge and wo on he inne pa ),
accompanied by one sligh ly la ge scu al basis on he in e nal
edge. The scu um i sel , a ely p ese ed, is a he la ge and i s
p oximal lobe is mo e de eloped. P ominen iangula la e al
a icula ia. When p esen , he on al a icula ia a e small and
always loca ed nea he in e nal dis al pa o he o icells, which
a e ounded, smoo h and exhibi a small p oximal enes a.
Do sal su ace wi h small iangula ib acula ia.
ema ks
The numbe o spine bases is ela i ely cons an , bu one o hem,
on he dis al im, is occasionally lacking. Only wo spines a e
obse able on o icella e zooids.
Supe amily cella ioiDea Lamou oux, 1821
Family cella iiDae Hincks, 1880
Genus Cella ia Ellis& Solande , 1786
Cella ia salico nioides Lamou oux, 1816
(Fig. 10F-H)
Cella ia salico nioides Lamou oux, 1816: 127.— Audouin 1826: 236;
Sa igny 1817: pl. 6, ig. 7.— P enan & Bobin 1966: 382, ex - ig.
124.— Ryland& Haywa d 1977: 124, ex - igs 56A, 59.— Poluzzi&
Pado ani 1984: 109, ig. 5d.— Zabala 1986: 307, ex - ig. 84:3a-e; pl. 3,
igs E-F.— Moisse e 1988: 104, pl. 17, igs 1-2.— Zabala& Maluque
1988: 94, ex - igs 137-140; pl. 2, ig. H.— Schmid 1989: 18, pl. 4,
igs 3-6.— El Hajjaji 1992: 129, pl. 6, ig. 7.— Moisse e& Spjeldnaes
1995: 788, pl. 3, ig. 1.— Haddadi-Hamdane 1996: 71, pl. 6, igs 12-
13.— López de la Cuad a& Ga cía-Gómez 1996: 158, igs 1C, 3A-F, 4.
occu ence.— Ea ly Miocene: F ance (Pouye 1991). Middle Mio-
cene: Aus ia (Schmid 1989), Hunga y (Moisse e e al. 2006), Czech
Republic, Poland (Pouye 1997). La e Miocene: Mo occo (El Hajjaji
1992), Alge ia (Moisse e 1988). Pliocene: Alge ia (Haddadi-Hamdane
1996), C e e. Pleis ocene: Sicily (Poluzzi& Pado ani 1984), Rhodes
(Moisse e& Spjeldnaes 1995). Recen : eas e n A lan ic (Mo occo,
Madei a, up o he She land Islands), Medi e anean, Red Sea. The
species has been eco ded om he A lan ic a dep hs o 0-360m (and
e en down o 636m in Mo occo). In he Medi e anean, i occu s om
he shallow in ali o al down o 280m.
ema ks
A numbe o specimens, especially hose om he Fane omeni
sec ion, esemble C.salico nioides a . no mani, a a ie y c e-
a ed by Has ings (1946) and conside ed as a dis inc species,
C.no mani, by P enan & Bobin (1966). Howe e , López
de la Cuad a& Ga cía-Gómez (1996) conside ed ha he
sligh mo phological di e ences (mo e slende in e nodes and
la ge a icula ia in C.no mani) do no jus i y he e ec ion
o a dis inc axon.
Subo de ASCOPHORA Le insen, 1909
In ao de HIPPOTHOOMORPHA Go don, 1989
Supe amily hippo hooiDea Busk, 1859
Family pasy heiDae Da is, 1934
Genus Gemellipo a Smi , 1873
Gemellipo a ebu nea Smi , 1873
(Fig. 10I)
Gemellipo a ebu nea Smi , 1873: 35, pl. 7, igs 152-156.— Ha me
1957: 994, pl. 69, igs 28-29.— Moisse e 1988: 190, pl. 30, igs
6, 9, 12.
Gemella ia punc a a – Seguenza 1880: 127, pl. 12, igs 14-14a.—
Ne iani 1900: 148, pl. 16, igs 89.
1384
GEODIVERSITAS • 2021 • 43 (26)
Moisse e P. e al.
Pasy hea ebu nea – Busk 1884: 5, pl. 34, ig. 1a- .— Canu& Bassle
1928: 151, pl. 8, igs 11-12.
occu ence.— Ea ly Miocene: Indonesia (Di Ma ino& Taylo
2014). Middle Miocene: Hunga y (Moisse e e al. 2006). La e Mio-
cene: Dominican Republic (Chee ham e al. 1999), Alge ia (Mois-
se e 1988), Calab ia. Pliocene: Sicily (Rosso 2002). Pleis ocene:
Calab ia, Sicily (Rosso& Di Ge onimo 1998; Rosso 2005), Rhodes
(Moisse e& Spjeldnaes 1995), Ka pa hos (Moisse e e al. 2017).
Holocene: sou he n I aly (Di Ge onimo e al. 2001). Recen : wes -
e n and eas e n A lan ic (B azil, Ca ibbean, Azo es, Madei a, Bay o
Biscay, Gul o Cadiz), Indian Ocean (Indonesia), Paci ic (Hawaii,
New Zealand). This is a deep-wa e species ound in he A lan ic a
dep hs be ween 60m and 3300m (Ha me 1957; Ha melin 1977).
Desc ip ion
Cella ii o m colony. In e nodes made o one o ou pai s
o zooids. In each pai , he zooids a e sepa a ed by a hin
g oo e, placed back o back, sligh ly wis ed om one
ano he , and o ien ed a abou 90 deg ees o he p e ious/
A
E
FG
HI
B
CD
ig. 10. — A-C, Sc upocella ia c . ellip ica (Reuss, 1847); A, iew o some zooids showing opesia wi h 4 spine bases on he ou e dis al pa and 3 spine bases + 1
scu al spine on he inne dis al im, FAN36: AMPG(IV) 3587; B, zooids wi h 7 spine bases and he scu um, FAN35: AMPG(IV) 3586a; C, do sal su ace o a colony
agmen wi h small ib acula ia and adicula po es, FAN35: AMPG(IV) 3586b; D, E, Sc upocella ia sc upea Busk, 1852; D, o icella e colony agmen , FAN35:
AMPG(IV) 3590a; E, de ail on al iew showing opesia wi h i e spine bases, FAN35: AMPG(IV) 3590b; F-H, Cella ia salico nioides Lamou oux, 1816; F, pa o
a slende in e node, FAN18: AMPG(IV) 3513a; G, de ail iew showing den icles and endo oichal o icells, POTB13: AMPG(IV) 2821a; H, a icula ia in on al and
la e al iew (a ow), POTB13: AMPG(IV) 2821b; I, Gemellipo a ebu nea Smi , 1873, b oken in e node showing wo zooids wi h wo o al sca s sepa a ed by a hin
in e zooidal g oo e, CAP032: AMPG(IV) 3426. Scale ba s: A-E, G, I, 100 µm; F, H, 200 µm.
1385
B yozoan aunas a he To onian-Messinian ansi ion
GEODIVERSITAS • 2021 • 43 (26)
ollowing pai . Zooids elonga e, displaying a smoo h on-
al su ace wi h sca ce inconspicuous po es. A ew zooids
bea in hei cen al pa a small o al sca (sepa a ed by
he hin g oo e di iding wo zooids) co esponding o he
s a o a la e al b anch. Ape u es almos ci cula wi h wo
e y small inden a ions on he p oximal co ne s. Nei he
a icula ia no o icells.
ema ks
Smi (1873) desc ibes an enc us ing base (Smi 1873: pl. 7,
ig. 152) om which he e ec , ypical o m a ises. Ano he
c eeping colony (Smi 1873: pl. 9, ig. 178) is e oneously
a ibu ed by he same au ho o G.ebu nea. The con usion
was e idenced by Canu& Bassle (1928).
In ao de UMBONULOMORPHA Go don, 1989
Supe amily lep alielloiDea Vigneaux, 1949
Family B yoc yp elliDae Vigneaux, 1949
Genus B yoc yp ella Cossman, 1906
B yoc yp ella o qua a (Jullien, 1903)
(Fig. 11A-C)
C yp ella o qua a Jullien in Jullien& Cal e , 1903: 77, pl. 7, ig. 5a-c.
Po ella o qua a – No man 1909: 300, pl. 39, igs 5-8.— Ba oso
1912: 33, igs 6, 6a-e.
B yoc yp ella o qua a – Ál a ez 1991: 93, ig. 5; pl. 10, igs A-E.
occu ence.— Recen : eas e n A lan ic (Gul o Gascony, Ma-
dei a). Deep-wa e species eco ded a dep hs om 80m o 300m
(Ál a ez 1991).
Desc ip ion
Colony incula ii o m. Na ow subcylind ical b anches o med
by 3
-
4 al e na ing longi udinal ows o zooids opening on he
same on al side. Zooids hexagonal sepa a ed by deep u -
ows. F on al en icose wi h abou 20 la ge po es. Ape u e
subci cula ; a small a icula ium is some imes p esen on i s
p oximal edge. Pe is ome mo e o less de eloped, de oid o
po es. Do sal sligh ly con ex showing ma ked g oo es co -
esponding o zooecial limi s and a ew po es. O icells no
obse ed.
ema ks
This species was e oneously iden i ied as Cha acodoma sp.
in Moisse e e al. (2018).
Family essa aDomiDae Jullien& Cal e , 1903
Genus Tessa adoma No man, 1869
Tessa adoma bo eale (Busk, 1860)
(Fig. 11D)
Onchopo a bo ealis Busk, 1860: 213, pl. 28, igs 6-7.
Po ina bo ealis – Hincks 1880: 229, pl. 31, igs 4-6.— Ne iani
1891: 120, pl. 4, igs 4-5.
Tessa adoma bo eale – Gau ie 1962: 222.— Haywa d& Ryland
1979: 242, ex - ig. 104.— Moisse e 1988: 123, pl. 20, ig.
5.— Zabala& Maluque 1988: 142, ex - ig. 343.— El Haj-
jaji 1992: 225, pl. 13, ig. 2.— Pouye & Moisse e 1992: 53,
pl. 7, ig. 3.— Moisse e& Spjeldnaes 1995: 790, pl. 3, ig. 5.
o
ccu ence
.— Middle Miocene: F ance. La e Miocene: Mo occo
(El Hajjaji 1992), Alge ia (Moisse e 1988), Calab ia. Pliocene:
no he n I aly, Sicily (Pouye & Moisse e 1992). Pleis ocene:
Sicily, Calab ia (Rosso 2005), Rhodes (Moisse e& Spjeldnaes
1995), Ka pa hos (Moisse e e al. 2017). Recen : A lan ic
(wes e n, bu mos ly eas e n), A c ic, Medi e anean. T.bo eale
is p edominan ly a deep-wa e (Chee ham 1972), c yophilic
(max. 13° C) species (60-3500m in he A lan ic, 50-1300m in
he Medi e anean).
Desc ip ion
Vincula ii o m colony. Slende cylind ical b anches. O al,
elonga e zooids a anged in al e na ing se ies. F on al smoo h
and inely s ia ed, wi h a ow o ma ginal po es. A spi a-
men is si ua ed p oximally o he semici cula ape u e,
which bea s a sho ubula pe is ome. A icula ia (1
-
3)
and o icell no obse ed.
ema ks
T.g acile (Sa s) is gene ally conside ed as a synonym o
T.bo eale.
Family ascosiiDae Jullien, 1883
Genus Kionidella Koschinsky, 1885
Kionidella excelsa Koschinsky, 1885
(Fig. 11E-F)
Kionidella excelsa Koschinsky, 1885: 68, pl. 7, igs 5-12.— Mal-
ecki 1963: 133, ig. 58, pl. 15, ig. 1.— B aga 1975, p. 147, pl.
3, ig. 67.— Moisse e 1988: 192, pl. 31, igs 4-6.— Zágo šek
2001: 554, pl. 19, ig. 4.— Zágo šek 2003: 176, pl. 28, ig. 6.
Fedo a excelsa – Wa e s 1891: 29, pl. 4, ig. 6.
occu ence.— Eocene: Ge many, Aus ia, I aly, Slo akia,
Hunga y, Romania, Poland (Zágo šek 2003). La e Miocene:
Alge ia (Moisse e 1988).
Desc ip ion
Small conical, hollow and a he elonga ed conescha el-
lini o m
colonies. Hexagonal zooids a anged in al e na ing
adial se ies. Smoo h con ex on al.
Pea -shaped ape u e wi h a la ge an e sepa a ed by
wo s ong ca delles om a smalle pos e . One o wo
well-de eloped o al a icula ia a e gene ally p esen on he
la e al sides o he zooid. O icell no obse ed.
ema ks
As no ed by se e al au ho s (e.g., Zágo šek 2003), he
a icula ia a e mo e o less de eloped and may ei he occu
in pai s, o alone, o lack comple ely. The o icells ha e
a ely been obse ed.
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GEODIVERSITAS • 2021 • 43 (26)
Moisse e P. e al.
A
D
E
F
BC
ig. 11. — A-C, B yoc yp ella o qua a (Jullien, 1903); A, colony agmen , on al iew, POTB13: AMPG(IV) 2815a; B, colony agmen , do sal iew, POTB13:
AMPG(IV) 2815b; C, de ail iew o zooids, FAN31: AMPG(IV) 3500a; D, Tessa adoma bo eale (Busk, 1860), colony agmen , FAN28: AMPG(IV) 3613. E-F, Kio-
nidella excelsa Koschinsky, 1885; E, colony agmen , basal iew, FAN7: AMPG(IV)3570a; F, on al iew o a ew zooids, FAN16: AMPG(IV) 3572. Scale ba s:
A, 500 µm; B, E, 200 µm; C, F, 100 µm; D, 1 mm.
1387
B yozoan aunas a he To onian-Messinian ansi ion
GEODIVERSITAS • 2021 • 43 (26)
Supe amily conescha ellinoiDea Le insen, 1909
Family Ba opo iDae Ne iani, 1900
Genus Ba opo a Reuss, 1867
Ba opo a osula (Reuss, 1847)
(Fig. 12A-G)
Cellepo a osula Reuss, 1847: 78, pl. 9, ig. 17; 1867: pl. 1, ig. 7a-
c; pl. 2, ig. 1a-c.
Ba opo a osula – Manzoni 1877: 54, pl. 2, ig. 6.— Malecki 1963:
135, pl. 15, ig. 4.— B aga 1975: 147, pl. 3, igs 10-11.— Cook&
Lagaaij 1976: 351, pl. 3, igs 2-3; pl. 4, igs 1-2.— Moisse e e al.
1993: 113, igs 7g-i.— Haddadi-Hamdane 1996: 121, pl. 2, ig.
4.— Moisse e 1996: 193, igs 1A-C.— Pizza e i& B aga 2000:
61, ig. 2.— Zágo šek 2010b: 168, pl. 83, ig. 4.
Lac imula sp. – Moisse e 1988: 193, pl. 31, igs 7-9.
occu ence.— Eocene: I aly, Romania (Malecki 1963; B aga 1975).
Ea ly Miocene: Spain, Co sica, I aly, Mal a (Moisse e 1996). Middle
Miocene: Aus ia, Hunga y (Moisse e e al. 2006), Czech Republic
(Zágo šek 2010b), no he n I aly, Calab ia, Mal a (Moisse e 1996).
La e Miocene: Alge ia (Moisse e 1988), Sa dinia, Sicily, Calab ia
(Pizza e i& B aga 2000), Mal a (PM, pe s. obs.), C e e (Moisse e
e al. 1993). Pliocene: Spain, Alge ia (Haddadi-Hamdane 1996),
no he n I aly, Sicily, C e e (PM, pe s. obs.), Ka pa hos (PM, pe s.
obs.). Pleis ocene: Rhodes (PM, pe s. obs.). This ossil species has
exclusi ely been eco ded om he Medi e anean-Pa a e hys ealm
(Moisse e 1996). Th ee p esen -day Ba opo a species a e known
om he Indo-Paci ic a dep hs be ween 285 and 880m (Cook&
Lagaaij 1976; Haywa d& Cook 1979).
Desc ip ion
Small conical conescha ellini o m colonies wi h a la ened base
an apical ube comp ised o kenozooids and e mina ed by a
small pi . Hexagonal zooids a anged in concen ic al e na ing
se ies. F on al con ex wi h ai ly la ge po es. La ge ci cula
ape u e loca ed in he dis al pa o each zooid. No a icula ia.
Ra e small b oken hype s omial o icells a e isible (Fig. 12A).
ema ks
A ew ju enile colonies a e p esen in a ai numbe o samples.
This was also obse ed by se e al au ho s (Cook& Lagaaij
1976; Pizza e i& B aga 2000).
Family o Bi ulipo iDae Canu& Bassle , 1923
Genus O bi ulipo a S olickza, 1862
O bi ulipo a excen ica Seguenza, 1880
(Fig. 12H-I)
O bi ulipo a excen ica Seguenza, 1880: 130, pl. 12, igs 22-22a.—
Ne iani 1900: 188, pl. 17, igs 15-16.— Wa e s 1919: 90, ex - ig.
2a-c.— Rosso& San ilippo 1991: 202, pl. 1, igs 1-5; pl. 2, igs
1-8.— Moisse e e al. 1993: 113, igs a-c.
occu ence.— La e Eocene: I aly (Wa e s 1919). Ea ly Miocene:
Sa dinia (Rosso& San ilippo 1991). La e Miocene: Calab ia (Rosso&
San ilippo 1991), C e e (Moisse e e al. 1993). O bi ulipo a and he
o bi ulipo i o m mo pho ype ( ossil and Recen ) a e conside ed as
deep-wa e ma ke s and li ing ep esen a i es as ypical o muddy
bo oms (Cook 1981; Rosso& San ilippo 1991).
Desc ip ion
Discoidal bilamina mo phology (o bi ulipo i o m). A sho
kenozooidal ube occu s a he apical pa o each colony.
Subci cula o subhexagonal zooids a anged in i egula
concen ic se ies and p og essi ely inc easing in size om he
apex o he base. F on al con ex wi h ela i ely la ge po es.
Ve y la ge ci cula ape u e loca ed in he cen e o each
zooid. No a icula ia. A ew o icells (o hei sca s) a e isible
a he g owing edge o some colonies. They a e hype s omial,
sphe ical, as la ge as a zooid and pe o a ed by po es simila
o hose o he zooecial on al.
DISCUSSION
aphonomy
Mos b yozoans (and o he skele al o ganisms) a e well p e-
se ed, excep a numbe o yellowish, ab aded agmen s o
li o al species (e.g., Adeonella polys omella, Ma ga e a ce e-
oides, Re epo ella sp., Smi ina ce ico nis, and S eginopo ella
mon ena i) indica ing anspo om shallow o deep wa e s.
Examples o such anspo ed ma e ial a e ound in h ee sec-
ions bu only in a ew beds (among which Po amida B13;
Kapa iana CAP18; Ke amou si KER22, KER28), whe e
nume ous shallow-wa e b yozoan emains a e ound oge he
wi h ewe deepe wa e elemen s.
palaeoen i onmen al signi icance
o he la e miocene B yozoans o c e e
Dep h
A gene al end is obse ed in all s udied sec ions (Figs 2-5).
S a ing in he la e To onian, a ela i ely deep basal
assemblage (deep ci cali o al) is la e eplaced by an
uppe ba hyal one. A shallowing-upwa d sequence ollows,
wi h deepe ci cali o al condi ions succeeded by shallow
ci cali o al o in ali o al ones. A sligh dissimila i y is
obse ed in he Ke amou si sec ion wi h e y shallow se -
ings (in ali o al-lagoonal) a he base, o e lain by shallow
ci cali o al en i onmen s and la e he same sequence as
in he o he h ee sec ions. This pa e n (Fig. 6), wi hin
di e en sub-basins, is in e p e ed as esul ing mos ly
om he s epped closu e o he Be ic-Ri ian co ido s. I
is alida ed by he analysis o he associa ed ben hic aunas
o o amini e a, bi al es, and os acods (Moisse e e al.
2018). Howe e , sligh di e ences a e no iced be ween
he wo esul s wi h he ecogni ion in he p esen pape
o shallowe en i onmen s (deep ci cali o al) a he base
o all sec ions.
Co ela ions be ween he p o ided plank onic o amini -
e al bios a ig aphic zona ion and he palaeoen i onmen al
scheme based on b yozoan (and o he ) aunas a e di icul .
The posi ion o he ba hyme ic e en s is ma kedly diach onic,
p obably because o di e en ial e ical mo ions in sepa a e
sub-basins (Fig. 13). The uppe ba hyal (deepes ) assemblage
is howe e loca ed in he la es To onian in h ee sec ions
(Po amida, Kapa iana, and Fane omeni) and in he ea lies
Messinian in he Ke amou si sec ion.
1394
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