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Lo enzo Conso i (Ins i u e o Ma ine Sciences, I alian Na ional Resea ch Council, T ies e)
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Wal e Joyce (Uni e si ä F eibu g, Swi ze land)
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489
COMPTES RENDUS PALEVOL • 2025 • 24 (24) © Publica ions scien i iques du Muséum e /and Académie des sciences, Pa is. www.c -pale ol.
An oine LOGGHE
Cen e de Reche che en Paléon ologie – Pa is (CR2P),
Muséum na ional d’His oi e na u elle, CNRS-MNHN-SU,
CP38, 8 ue Bu on, F-75005 Pa is (F ance)
[email p o ec ed] (co esponding au ho )
A naud REBILLARD
Leibniz-Ins i u ü E olu ions- und Biodi e si ä s o schung,
Museum ü Na u kunde, In alidens aße, 43, 10115 Be lin (Ge many)
and Ins i u ü Biologie Humbold -Uni e si ä zu Be lin,
In alidens aße, 42, 10115 Be lin (Ge many)
and Cen e de Reche che en Paléon ologie – Pa is (CR2P),
Muséum na ional d’His oi e na u elle, CNRS-MNHN-SU,
CP38, 8 ue Bu on, F-75005 Pa is (F ance)
Guillaume HOUÉE
Alicia SÁNCHEZ GIMENO
Cen e de Reche che en Paléon ologie – Pa is (CR2P),
Muséum na ional d’His oi e na u elle, CNRS-MNHN-SU,
CP38, 8 ue Bu on, F-75005 Pa is (F ance)
E wan COURVILLE
Biogéosciences, Uni e si é de Bou gogne, CNRS,
6 Boule a d Gab iel, F-21000 Dijon (F ance)
Laza e ELBAZ
Depa men o He pe ology, STBio pos -g adua ion p og am,
Museu de Zoologia da Uni e sidade de São Paulo, A enida Naza é 481,
Ipi anga, São Paulo 04263-000, SP (B azil)
Paloma-Ma ia TARABA
Mai ie de Sain -Y ieix La Pe che, 45 boule a d de l’hô el de ille,
F-87500 Sain -Y ieix La Pe che (F ance)
Rodolphe PASO
1 a enue du ec eu Pineau, F-86000 Poi ie s (F ance)
Max HERDE
Leibniz-Ins i u ü E olu ions- und Biodi e si ä s o schung,
Museum ü Na u kunde, In alidens aße, 43, 10115 Be lin (Ge many)
and Ins i u ü Biologie, Humbold -Uni e si ä zu Be lin,
In alidens aße, 42, 10115 Be lin (Ge many)
S anisla ŠTAMBERG
Museum o Eas e n Bohemia, H adec K álo é
Eliščino nábřeží 465, 500 01 H adec K álo é (Czech Republic)
Palaeoecology and palaeoen i onmen
o Discosau iscus Kuhn, 1933 wi h excep ional p ese a ion
o skin and in es ine molds om he ea ly Pe mian
o F anchesse (Allie , F ance)
490 COMPTES RENDUS PALEVOL • 2025 • 24 (24)
Logghe A. e al.
u n:lsid:zoobank.o g:pub:DBE4A506-5D02-4AFF-846E-3E367D472281
Logghe A., Rebilla d A., Houée G., Sánchez Gimeno A., Cou ille E., Elbaz L., Ta aba P.-M., Paso R., He de M.,
Š ambe g S., Pouillon J.-M. & S eye J.-S. 2025. — Palaeoecology and palaeoen i onmen o Discosau iscus Kuhn,
1933 wi h excep ional p ese a ion o skin and in es ine molds om he ea ly Pe mian o F anchesse (Allie , F ance).
Comp es Rendus Pale ol 24 (24): 489-504. h ps://doi.o g/10.5852/c -pale ol2025 24a24
ABSTRACT
The Ca boni e ous-Pe mian is a key pe iod conce ning he eme gence and di e si ica ion o majo
e apod clades such as amphibians and amnio es. This is pa icula ly he case o he Seymou iamo pha
Wa son, 1917. Some seymou iamo phs a e well-known, such as Seymou ia B oili, 1904 om No h
Ame ica and Ge many, bu he ecology o mos axa emains deba ed. This is he case o Discosau iscus
Kuhn, 1933 om he ea ly Pe mian o Eu ope. He e we p o ide a p elimina y s udy o new specimens
om he ea ly Pe mian Lage s ä e o F anchesse (Allie , F ance) whose excep ionally well-p ese ed
so issues including skin and in es inal cas s p o ide aluable insigh s in o he palaeoecology o
Discosau iscus. We desc ibe o he i s ime he comple e skin ex e nal and his ological ea u es o
a seymou iamo ph indica ing ha ju enile discosau iscids al eady displayed epide mal scala ion.
This sugges s a skin adap ed owa ds e es ial li es yle in ju enile Discosau iscus. The eco e y o an
in si u cololi e acili a es he iden i ica ion o seymou iamo ph cop oli es ound ex si u, and hus,
u u e in e p e a ions o he speci ic die o his axon. In o de o be e unde s and how hese so
issues ha e been p ese ed, we also conduc ed a aphonomic s udy including a s a ig aphic and
palaeoen i onmen al analysis o he ossil laye deposi s. Ou esul s sugges ha p onounced olcanic
ac i i y a wo k du ing he ea ly Cisu alian o he Bou bon-l’A chambaul Basin may be he cause
o he mul iple Pe mian mass mo ali y assemblages ound a F anchesse, as well as he excep ional
p ese a ion o he ossil emains om his si e.
RÉSUMÉ
Paléoécologie e paléoen i onnemen de Discosau iscus Kuhn, 1933 a ec p ése a ion excep ionnelle des
moules de peau e d’in es in du Pe mien p écoce de F anchesse (Allie , F ance).
Le Ca boni è e-Pe mien es une pé iode clé en ce qui conce ne l’éme gence e la di e si ica ion des
p incipaux clades de é apodes els que les amphibiens e les amnio es. C’es no ammen le cas des
Seymou iamo pha Wa son, 1917. Ce ains seymou iamo phes son bien connus, comme Seymou ia
B oili, 1904 d’Amé ique du No d, mais l’écologie de la plupa des axons es e déba ue. C’es le cas de
Discosau iscus Kuhn, 1933 du Pe mien in é ieu d’Eu ope. Ici, nous é udions de maniè e p éliminai e
de nou eaux spécimens p o enan du Lage s ä e du Pe mien in é ieu de F anchesse (Allie , F ance)
don les issus mous excep ionnellemen bien conse és, y comp is la peau e le moulage des in es ins,
pe me en d’en app end e da an age su la paléoécologie de Discosau iscus. Nous déc i ons pou la
p emiè e ois la mo phologie ex e ne e in e ne de la peau d’un seymou iamo phe, indiquan que
les discosau iscidés ju éniles p ésen aien des écailles épide miques. Cela suggè e une peau adap ée à
un mode de ie e es e chez les Discosau iscus ju éniles. La décou e e d’un cololi he insi u acili e
l’iden i ica ion des cop oli hes de seymou iamo phes ou és exsi u e ainsi de u u es in e p é a ions
su le égime alimen ai e de ce axon. A in de mieux comp end e commen ces issus mous on é é
p ése és, nous a ons égalemen mené une é ude aphonomique comp enan une analyse s a ig a-
phique e paléoen i onnemen ale des dépô s des couches ossili è es. Nos ésul a s suggè en qu’une
ac i i é olcanique ma quée au cou s du Cisu alien in é ieu dans le bassin de Bou bon-l’A chambaul
pou ai ê e à l’o igine des mul iples assemblages de mo ali é massi e ou és à F anchesse, ainsi
que de la p ése a ion excep ionnelle des es es ossiles de ce si e.
Jean-Ma c POUILLON
Rhinopolis, 179 ue des Pla iè es, F-38300 Ni olas Ve melle (F ance)
J.-Sébas ien STEYER
Cen e de Reche che en Paléon ologie – Pa is (CR2P),
Muséum na ional d’His oi e na u elle, CNRS-MNHN-SU,
CP38, 8 ue Bu on, F-75005 Pa is (F ance)
Submi ed on 4 Feb ua y 2025 | Accep ed on 20 June 2025 | Published on 6 No embe 2025
KEY WORDS
Pe mian,
Seymou iamo pha,
skin,
cololi e,
palaeoecology.
MOTS CLÉS
Pe mien,
Seymou iamo pha,
peau,
cololi he,
paléoécologie.
491
Palaeoecology and palaeoen i onmen o Discosau iscus
COMPTES RENDUS PALEVOL • 2025 • 24 (24)
INTRODUCTION
Complemen a y o he s udy o mine alised issues in ossil
e eb a es, new palaeoecological ai s can be in e ed om
ossilised so issues, such as skin (e.g. Vincen e al. 2017;
Hend ickx e al. 2022; Jagielska e al. 2024), and b omali es
(Ho mann e al. 2020; Halaçla e al. 2023; Q a ns öm e al.
2024). Indeed, he skin, as i s ba ie be ween he o ganism
and he ex e nal en i onmen , shows se e al adap a ions such
as p o ec ion agains mechanical s ess, UV- adia ion o p ed-
a o y a acks (Vicka yous & Si e 2009). I can also p e en
desicca ion (Aliba di 2003), a ou he mo egula ion (Lindg en
e al. 2014), and is a main componen o beha iou al displays
such as camou lage and/o sexual display, o example h ough
colou a ion (e.g. McNama a e al. 2021). Fossilised diges i e
emains, p ese ed ei he wi hin (e.g. consumuli es) o ou side
he body (e.g. cop oli es, egu gi ali es), also p o ide aluable
in o ma ion ega ding palaeoecology (e.g. en i onmen al
ac o s; Hun 1992; No hwood 2005) o palaeobiodi e si y
(some lo al and aunal elemen s a e only p ese ed in b o-
mali es; By ne e al. 2022). Especially, consumuli es such as
gas oli es o cololi es a e he mos compelling e idence in he
ossil eco d o he p eda o -p ey in e ac ions (e.g. Q a ns öm
e al. 2024). Fu he mo e, ossilised gu con en is p ecious
e idence allowing o di ec ly link a p eda o wi h i s p ey
and en i onmen , a a ela i ely p ecise empo al and spa ial
esolu ion (K iwe e al. 2008; Mondini 2023). Howe e ,
as bo h so issues and b omali es ossilise mo e a ely han
mine alised issues, hese key aspec s o he palaeoecology
o Palaeozoic e apods a e poo ly known (e.g. Eliason e al.
2017; Hun & Lucas 2021).
The Discosau iscidae Rome , 1947 is a clade o Seymou-
iamo pha Wa son, 1917. Thei palaeoecology is deba ed
be ween mainly aqua ic o mo e amphibious/ e es ial li e-
s yle (Klemba a 1995; Malakho 2000; K ilo e al. 2008;
Sanchez e al. 2008; Calábko á e al. 2022). Mainly, ex e nal
gills a e equen ly p ese ed in discosau iscids (I akhnenko
1987; Klemba a 1995; Bulano 2003). Cololi es ha e been
desc ibed in associa ion wi h discosau iscid emains, sugges ing
pu a i e cannibalism (Klemba a & Meszá oš 1992). De mal
co e age has been desc ibed bu only assigned o ossi ied
de mal scales, wi hou palaeoecological implica ions (Špina
1952; Klemba a & Ba ík 1999). Thus, almos no skin so
issue is known o his clade, and e en o all Seymou ia-
mo pha (Olson 1965).
The Pe mian Lage s ä e o F anchesse (Allie , F ance), dis-
co e ed by P o . Roge -Paul Dechamb e (MNHN) in 1964,
yielded hund eds o sub-comple e o comple e specimens o
Discosau iscus Kuhn, 1933, p ese ing so - issue emains (S eye
e al. 2012: ig. 2). F om 2007, one o us (J.-S.S.) o ganised
egula sys ema ic exca a ions a he Les Cha bonniè es si e in
F anchesse, unde a wo king con en ion wi h Epona Socie y,
owne o he si e. These ieldwo ks led o he disco e y and
desc ip ion o a e y ich lo a and auna including nume ous
aeduellids, one pa ial Bohemiacan hus Schneide , 1996 sha k,
nume ous exquisi e Discosau iscus Kuhn, 1933 specimens, and
a la al e yopoid emnospondyl (S eye e al. 2012). Then A.L.
co-o ganised u he ieldwo ks (2021, 2022, 2024) in o de
o collec mo e palaeon ological da a, o be e unde s and
he so issue p ese a ion and o p ecise he s a ig aphic log
o he locali y (Houée e al. 2022a, b; Logghe e al. 2023a).
He e we desc ibe new Discosau iscus specimens om he
la es ieldwo ks, wi h excep ionally p ese ed comple e skin
co e age iden i ied as epide mal skin, and a cololi e eco -
e ed in si u wi hin a comple e skele on. We also p ecise he
s a ig aphic log o he locali y. These new disco e ies allow:
1) he iden i ica ion o seymou iamo ph skin so issues and
i s compa ison wi h he cu aneous co e age o o he Palaeo-
zoic e apods; 2) he s udy o a seymou iamo ph cololi e;
3) he discussion o hese new da a in e ms o palaeoecology
(e.g. skin adap a ion o an aqua ic o amphibious/ e es ial
li e), and palaeoen i onmen ; and 4) o be e unde s and
he mass mo ali y o Discosau iscus om F anchesse and he
excep ional quali y o p ese a ion o he desc ibed so - issue
and cololi e emains.
GEOLOGICAL SETTING
The Les Cha bonniè es si e in Le Pon e , 4 km nea he il-
lage o F anchesse, is loca ed in he no h o he Bou bon-
l’A chambaul Basin (Allie , Massi Cen al, F ance), app ox.
10 km away om he ea ly Pe mian his o ic si e o La Queue-
de-l’É ang (de Sain -Seine 1949; Heyle 1969). Mo e p ecisely,
his si e belongs o he Sou igny sub-basin (S eye e al. 2012;
Fig. 1A). As wi h he o he Ca boni e ous-Pe mian basins
o Au un, Blanzy-Le C euso and Decize-La Machine, he
Bou bon-l’A chambaul Basin is loca ed in he no he n pa
o he Massi Cen al (Deb ie e 1992), belonging o he
Va iscan Bel o he Cen al Pangea (Châ eauneu & Fa janel
1989). The ossil-bea ing ho izons o he Les Cha bon-
niè es si e come om he uppe -mos shale sequence o he
Reniè e B Membe o he Reniè e Fo ma ion (S eye e al.
2012). The Reniè e Fo ma ion has been iden i ied om he
si e o Buxiè es-les-Mines, sou h o he Aumance sub-basin o
he Bou bon-l’A chambaul Basin (S eye e al. 2000, 2012;
Kaul uss 2003; Fig. 1B). The unde lying Buxiè es Fo ma ion
may be o ea ly A inskian age (288 ± 4 Ma; Kaul uss 2003
acco ding o he In e na ional Ch onos a ig aphic Cha
2024-12) bu ecen U-Pb da ing in he Buxiè es Fo ma-
ion a he p oposed a lowe mos Asselian age (Cisu alian,
lowe Pe mian) o 298.592 ± 0.075/0.15/0.35 Ma (Me cuzo
e al. 2023). An absolu e age o he Les Cha bonniè es si e
has no been calcula ed so a , bu a Sakma ian age has been
p oposed based on he p e iously assigned Sakma ian age
o he Buxiè es Fo ma ion (S eye e al. 2012, based on he
ICC 2010-09). An olde age o he Les Cha bonniè es si e
is in his way possible ega ding he new adiome ic ages
o he Buxiè es Fo ma ion.
Valli (2007) conduc ed he i s s a ig aphic su ey o
he Les Cha bonniè es si e (S eye e al. 2022: ig. 2A) and
obse ed a s ong ho izon al a iabili y o he laye s in he
span a o ew me e s, wi h changes in bo h sedimen ology
and ossil emain con en s. Ou ecen ieldwo ks (exca a ions
492 COMPTES RENDUS PALEVOL • 2025 • 24 (24)
Logghe A. e al.
ig. 1. — Geological con ex o he Les Cha bonniè es si e nea F anchesse, Bou bon-l’A chambaul Basin (Allie , F ance) wi h loca ions o he main ossili e ous
deposi s o La Queue-de-l’É ang, Buxiè es-les-Mines and he ea ly Pe mian Lage s â e o F anchesse: A, simpli ied map o he Ca boni e ous and Pe mian
Bou bon-l’A chambaul Basin (modi ied a e Paque e & Feys 1989); B, log o he main locali ies, om bo om o op: Buxiè es-les-Mines, F anchesse, La Queue-
de-l’É ang; ed s a , s a ig aphic posi ion o he Les Cha bonniè es si e (modi ied a e Deb ie e 1992); C, de ailed s a ig aphic log o he SM sec o o he
Les Cha bonniè es si e nea F anchesse.
pos -Pe mian ocks
lowe Pe mian
uppe Ca boni e ous
C ys alline basemen
Gipcy-Boubon idge
?
A
BC
100 m
5 km
N
F anchesse
F anchesse
Bou bon-l'A chambaul
Bou bon-
l'A chambaul
Aumance
sub-basin
Sou igny
sub-basin
Buxie es
Buxie es-
les-Mines
Allie i e
Mu a aul
Reniè e
Fo ma ion
Reniè e B
A
A
A
A
A
A
Reniè e A
Pe mian
Cisu alian
Buxiè es
Fo ma ion
Sup a
Buxiè es
In a
Buxiè es
Basal
conglo.
In é ieu e
Fo ma ion
Ca boni e ous
Pennsyl anian
Sec o SM
C ys alline ock
A gili es
Sands one
Oil-shale beds
Ash- u (A )
Soil
Clay
Clays one
Sil s one
Cine i e
Gypsum
Discosau iscus
Kuhn, 1933
Plan ossil
Ac inop e ygians
493
Palaeoecology and palaeoen i onmen o Discosau iscus
COMPTES RENDUS PALEVOL • 2025 • 24 (24)
and de ailed s a ig aphy; Fig. 1C) ha e been ealised in he
sec o SM (see S eye e al. 2022: ig. 2A), which showed a
high po en ial o eco e ing Discosau iscus specimens h ough
d illings (Valli 2007; S eye e al. 2022). The s a ig aphy o
his sec o is composed o , om bo om o op (Fig. 1C): an
al e nance (15 cm) o c ys allised gypsum, a possible cine i e
and shales; an al e nance (4 cm) o clays ones, clays and a da k
indu a ed shale le el (3 cm) wi h nume ous well-p ese ed
Discosau iscus emains o a ious sizes. F om his laye , a e
plan agmen s and ou slabs con aining (sub)comple e
Discosau iscus – hei aphonomy will be he subjec o ano he
a icle – we e sampled. Abo e, a hin (0.5 cm) o ange cin-
e i e, sampled o absolu e age in es iga ions, is associa ed
wi h c ys allised gypsum and whi e o g ey clay. I is opped
by a da k indu a e clays one (10 cm) ich in plan agmen s,
eco e ed in associa ion wi h ano he possible cine i e laye
(1 cm, also sampled). This plan laye is opped by an al e -
nance o clay (1 cm) wi h c ys allised gypsum, pape shale
laye s (5 cm), hen a da ke hin clays one laye (4 cm) wi h
nume ous ac inop e ygian emains. Abo e his “ ish”-laye ,
nume ous specimens o Discosau iscus ha e been ound in a
g ey shale le el (10 cm). They a e excep ionally well-p ese ed
and abundan , wi h se e al comple e specimens exposed side
by side (e.g. S eye e al. 2012, 2022; Houée e al. 2022a;
Logghe e al. 2023a). The ma e ial desc ibed he e comes
om his laye . I has been eco e ed in associa ion wi h an
ig. 2. — New specimens o Discosau iscus sp. om he Pe mian o F anchesse: A, RH-FR-2022-01, comple e and a icula ed ju enile indi idual (only he
igh o elimb is missing) p ese ing de mal so - issue alongside he whole body; B, RH-FR-2022-07, sub-comple e indi idual o poo c anial and so - issue
p ese a ion bu yielding an in si u mold o he in es ine; C, RH-FR-2022-03, sub-comple e indi idual p ese ing de mal so - issue alongside he whole body.
Scale ba s: A, B, 5 cm; C, 3 cm. C edi s: Lilian Cazes (CR2P).
A
B
C
494 COMPTES RENDUS PALEVOL • 2025 • 24 (24)
Logghe A. e al.
o ange dus y laye , possibly ano he cine i e (2 cm), sampled
as well o p ecise i s na u e and un absolu e da ing analyses.
Abo e was obse ed a g ey-yellowish clays one (17 cm) wi h
mos ly well-p ese ed ac inop e ygians, a e chond ich hyan
spines and some la ge and sub-comple e Discosau iscus a he
base. Finally, his laye is opped by da k and wea he ed clay-
s ones (20 cm) wi h isola ed small Discosau iscus agmen s
and nume ous mo e o less comple e ac inop e ygians, and
hen he g ass soil.
MATERIAL AND METHODS
We sampled ou specimens o a icula ed Discosau iscus
(RH-FR-2022-01, RH-FR-2022-02, RH-FR-2022-03 and
RH-FR-2022-07; Figs 2; 3) whose so issues a e p ese ed,
including a specimen con aining a cololi e ound in si u in he
abdomen (RH-FR-2022-07; Fig. 2B). All hese specimens we e
collec ed om he Pe mian o F anchesse (see he Geological
Se ing sec ion o mo e de ails) and a e pa o he collec ions
o he Rhinopolis Associa ion, which aims o p omo e and
p ese e he palaeon ological he i age o he Dépa emen
de l’Allie (Allie coun y, F ance). All specimen images we e
aken using a Nikon D800 came a (apn ull- ame 36 Mpx)
wi h a lens Mak o-Plana Zeiss 50 mm. Fo mac o, we used a
Canon 5DSR came a (apn ull- ame 50Mpx) wi h a Canon
MP-E65 mm lens moun ed on mic ome e ail and illumina ed
using cold ligh on ibe . The images we e p ocessed in Focus
S acking using Helicon Focus ( . 8.3.0). The images we e
aken and p ocessed by Lilian Cazes (CR2P). The specimens
a e s o ed in he collec ions o he Rhinopolis Associa ion in
Va ennes-su -Allie (Allie , F ance).
IDENTIFICATION
SyS ema icS
The s udied specimens show an exquisi e p ese a ion o
he so issues bu a e y poo p ese a ion o he skele al
elemen s, especially hose om he skull: his ende s hei
sys ema ic iden i ica ion e y di icul a he species le el.
The aim o his pape is no o gi e a de ailed sys ema ic
palaeon ological desc ip ion o he whole skele al elemen s.
Howe e , some skele al elemen s allow an iden i ica ion a
he amily and genus le el:
1) all he specimens ha e ela i ely sho unks and s ou
limbs. RH-FR-2022-02 p ese es a leas 24 p esac al e e-
b ae (Fig. 2C), and RH-FR-2022-01 small ibs a he le el
o he ce ical e eb ae and a le oo wi h a phalangeal
o mula o 2-3-4-5-3 (Fig. 2A). Also, RH-FR-2022-03
shows swollen neu al a ches, la ge cylind ical pleu ocen-
a, and haemal a ches a he le el o he ail (Fig. 3): hese
cha ac e s a e o en seen in Discosau iscidae (e.g. Klemba a
& Ba ík 1999; Lau in 2010).
2) hese specimens ha e ela i ely sho snou s, ounded
o bi s lying an e io ly o he middle o he skull leng h,
and a ela i ely high skull in he egion o he o ic no ches
when p ese ed: hese cha ac e s a e ypical o he genus
Discosau iscus, e.g. Klemba a 1997. Un o una ely, as men-
ioned abo e, he peculia p ese a ion o hese specimens
does no allow o iden i y he species: pending a mo e
ho ough sys ema ic s udy o he whole ma e ial o F anchesse
(composed o hund eds o specimens), we he e o e p e e
o s ay p uden and o e e he s udied specimens as o
Discosau iscus sp. This iden i ica ion is consis en wi h ha
o S eye e al. (2012, 2022).
ig. 3. — New specimen (RH-FR-2022-02) o Discosau iscus sp. om he Pe mian o F anchesse, p ese ing de mal so - issues: pos e io body egion (le ),
and an e io body egion ( igh ) o a p obable unique indi idual. Scale ba : 5 cm. C edi s: Lilian Cazes (CR2P).
495
Palaeoecology and palaeoen i onmen o Discosau iscus
COMPTES RENDUS PALEVOL • 2025 • 24 (24)
G ow h S aGe
The high ossi ica ion deg ee o he p ese ed long bones, ibs
and e eb ae; he well-de eloped o namen a ion pa e n o he
de mal bones; he ac ha he o bi s a e ela i ely an e io ly
loca ed wi hin he skull; he ela i ely long pos o bi al egion;
and he obus mandibles p ese ing well-de eloped ee h a e
ypical o a la e ju enile on ogene ic s age (Klemba a 1997;
S eye 2000; Sanchez e al. 2008). Howe e , acco ding o Klem-
ba a (2009) who dis inguished on ogene ic s ages based on skull
leng h only, hese specimens migh be younge in soma ic age.
Conside ing all he cha ac e s oge he , we p e e o s ay p uden
and o gi e an “a e age” ju enile age o he s udied specimens.
DESCRIPTION OF THE SOFT TISSUES
Pu a i e ePide mal ScaleS (Figs 4, 5)
Se e al ows o scales a e well-p ese ed on RH-FR-2022-01
(Fig. 4A, B). These scales seem o be posi ioned on he lanks
and/o do sal side o he specimen as hey do no appea
o be below he e eb al column bu a he cons ained
a ound i . These scaly skin emains a e mos ly p ese ed
on he body egion o he o he Discosau iscus skele ons,
and less de ailed skin emains a e isible a ound he ails.
Skin emains om he shoulde gi dles and pel ic egions
a e seldomly isible (see RH-FR-2022-03; Fig. 4C), and
o e all appea smoo he han in he ho acic egion, and
dep i ed o scales. These skin emains p esen a blackish
colo a ion, consis en wi h ha o he es o he skele on
and wi h o he e apod skin emains eco e ed om he
F anchesse locali y, and con as ing wi h he g ey colou
o he subs a e. They a e p ese ed in hee-dimensions
(Fig. 4B), and a e he e o e no he esul o he imp ession
o he skin on he subs a e. The scales a e mos ly iangula
(Fig. 4B), al hough some seem a he quad angula (Fig. 4C).
They measu e be ween 1 and 1.5-1.7 mm. No concen ic
ings seem o be p ese ed wi hin he scales. Howe e , hey
seem o p ese e idges, ha ollow he o e all shape o he
scales (Fig. 5A). Some kind o ube cule-like o namen a-
ion is also seen on some scales, howe e wi hou de ini e
A
B
C
D
ig. 4. — Close-ups on de mal so - issues o Discosau iscus sp. specimens om he Pe mian o F anchesse: A, RH-FR-2022-01, showing a angemen in
longi udinal and ans e sal ows; B, mac opho og aphy o he scales o RH-FR-2022-01 showing he iangula ( o quad angula in some egions) shape o
he scales. The a angemen in longi udinal and ans e sal ows is well isible he e; C, RH-FR-2022-02, showing a simila a angemen o he scales as in he
o he specimens, p ese ing howe e a mo e quad angula shape o he scales; D, RH-FR-2022-03, showing he longi udinal and ans e sal a angemen
o he scales. The ows seem less s aigh han hose in RH-FR-2022-01 bu display a mo e ‘wa y’ a angemen alongside he body. Scale ba s: A, D, 2 cm;
B, 2 mm; C, 1 cm. C edi s: Lilian Cazes (CR2P).
502 COMPTES RENDUS PALEVOL • 2025 • 24 (24)
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Submi ed on 4 Feb ua y 2025;
accep ed on 20 June 2025;
published on 6 No embe 2025.
