New, large actinopterygian fishes from the upper Carboniferous of Nýřany, Czech Republic

Ac a Palaeon ol. Pol. 69 (3): 501–522, 2024 h ps://doi.o g/10.4202/app.01162.2024
New, la ge ac inop e ygian ishes om
he uppe Ca boni e ous o Nýřany, Czech Republic
PAVEL BARTÁK, MARTIN IVANOV, EVA TIHLAŘÍKOVÁ, MARTIN OLBERT,
and VILÉM NEDĚLA
Ba ák, P., I ano , M., Tihlaříko á, E., Olbe , M., and Neděla, V. 2024. New, la ge ac inop e ygian ishes om he uppe
Ca boni e ous o Nýřany, Czech Republic. Ac a Palaeon ologica Polonica 69 (3): 501–522.
The lacus ine coal deposi s a Nýřany, Czech Republic, yielded a di e si ied e eb a e assemblage o he Middle
Pennsyl anian (Mosco ian) age, ep esen ed by he emains o ea ly e apods, as well as nume ous eshwa e ich hy-
o auna, including xenacan hi o m sha ks, acan hodians, dipnoans, and ay- inned ishes. Howe e , unlike some o he
uppe Ca boni e ous locali ies, he ac inop e ygian di e si y is limi ed o he h ee small-bodied species, mos o hem
endemic o Nýřany locali y and he equi alen s a a elsewhe e, indica ing ha he ue axonomic di e si y o he g oup
a he locali y may be biased. He e we desc ibe i s skele al emains o la ge ac inop e ygian ishes om he si e, in-
cluding a new genus and species, S ambe gich hys mac odens gen. e sp. no ., which is ep esen ed by a well-p ese ed
mandible wi h ee h. The mic o-compu ed omog aphic echniques e ealed in he specimen a p esence o a complex
neu o ascula sys em inne ing he ee h and he jaw, and supplying bo h wi h blood essels. The den i ion consis s o a
single ow o massi e, homodon , conical ee h, which possess simplexodon pliciden ine on hei base, he cha ac e is-
ics suppo ing he p eda o y ecology o he new species. The isola ed skele al emains o la ge-bodied ac inop e ygians
expand he knowledge on he di e si y o he g oup in Nýřany, and hei occu ence in coal deposi s o ela i ely shallow
lake indica es hey ep esen alloch honous, poo ly known aqua ic e eb a e associa ion, likely o igina ing om he
b aided i e sys em. These indings unde line he impo ance o less comple e skele al ma e ials occu ing in well-
known e eb a e assemblages o he uppe Ca boni e ous coal-bea ing locali ies.
Key wo ds: Ac inop e ygii, den i ion, con inen al basins, mo phology, neu o ascula sys em, palaeoen i onmen ,
pliciden ine, Palaeozoic.
Pa el Ba ák [[email p o ec ed]; ORCID: h ps://o cid.o g/0009-0009-3738-4849 ] and Ma in I ano
[ [email p o ec ed]; ORCID: h ps://o cid.o g/0000-0001-9108-9239 ], Depa men o Geological Sciences, Facul y
o Science, Masa yk Uni e si y, Ko lářská 267/2 611 37 B no, Czech Republic.
E a Tihlaříko á [[email p o ec ed]; ORCID: h ps://o cid.o g/0000-0002-7983-2971 ], Ma in Olbe [[email p o ec ed];
ORCID: h ps://o cid.o g/0000-0003-2280-3341 ], and Vilém Neděla [[email p o ec ed]; ORCID: h ps://o cid.o g/
0000-0001-6029-5435 ], En i onmen al Elec on Mic oscopy G oup, Ins i u e o Scien i ic Ins umen s o he Czech
Academy o Sciences, K álo opolská 147, 612 00 B no, Czech Republic.
Recei ed 17 Ap il 2024, accep ed 25 July 2024, published online 30 Sep embe 2024.
Copy igh © 2024 P. Ba ák e al. This is an open-access a icle dis ibu ed unde he e ms o he C ea i e Commons
A ibu ion License ( o de ails please see h p://c ea i ecommons.o g/licenses/by/4.0/), which pe mi s un es ic ed use,
dis ibu ion, and ep oduc ion in any medium, p o ided he o iginal au ho and sou ce a e c edi ed.
In oduc ion
The Nýřany locali y is pa o a coal ield si ua ed sou hwes
o Pilsen in he Czech Republic, which con ains se e al
uppe Ca boni e ous (Middle Pennsyl anian) coal seams,
de eloped wi hin he g ey-colou ed sequence o lu iola-
cus ine sedimen s o he Pilsen Basin. Remnan s o e -
eb a es come om se e al ossili e ous sap opelic ho i-
zons inside he Main Nýřany Coal Seam (F i sch 1879;
Milne 1980; Ba ák 2020) o he Nýřany Membe (Kladno
Fo ma ion), da ed as he la e Mosco ian (As u ian) age
(Opluš il e al. 2016). Thei collec ion began in he second
hal o he 19 h cen u y (F i sch 1870), wi h mos impo an
indings being de i ed om he Humbold and K imich
mines in he icini y o he Nýřany illage. O e he pas
ca. 150 yea s, nume ous e eb a e body ossils ha e been
desc ibed om he si e, ep esen ing one o he bes known
la e Mosco ian aunal assemblages wo ldwide. Much o he
e eb a e skele al emains a e ai ly comple e, a icula ed
specimens, indica ing a apid bu ial unde he anoxic con-
di ions wi h minimal e ec s o anspo (Milne 1980).
The enowned s a us o he Nýřany locali y undoub edly
a ises mainly om he highly di e si ied and especially
well-p ese ed assemblage o ea ly e apods, including
502 ACTA PALAEONTOLOGICA POLONICA 69 (3), 2024
baphe ids (Beaumon 1977; Milne e al. 2009), embo-
lome es (Ca oll 1970), a ious g oups o emnospondyls
and “lepospondyls” ( o he ecen o e iew, see Ba ák
2020), and he ea lies amnio es (Ca oll and Bai d 1972;
Reisz 1975). Howe e , he ich hyo auna is also well- ep-
esen ed and ela i ely di e se, comp ising acan hodians
(Zajíc 1988), xenacan hi o m sha ks (Schneide and Zajíc
1994; Heid ke 1998), dipnoans (F i sch 1888) and ac inop e-
ygian ishes (Š ambe g 1978, 1991, 2013). In compa ison
o he o he la e Ca boni e ous locali ies such as Newsham
(No humbe land, UK), Lin on (Ohio, USA), Mazon C eek
(Illinois, USA), Mon ceau-les-Mines (F ance), and Kouno
(Czech Republic), he coal-bea ing deposi s o Nýřany ha e
so a p oduced su p isingly ew ac inop e ygian species.
Al hough abundan in e ms o specimen numbe , hese
a e cu en ly con ined o only h ee small-bodied o ms,
he haplolepid Py i ocephalus sculp us and wo species o
he endemic genus Scele opho us, i.e., S. bise ialis and S.
e ucosus, none o hem exceeding he o al leng h o 150
mm (Wes oll 1944; Ga dine 1967; Š ambe g 1978, 1991,
2013). These axa ha e been pe cei ed as inhabi an s o
small and shallow, poo ly oxygena ed eshwa e s (Wes oll
1944; Milne 1980), he en i onmen al cons ain s o which
would po en ially p e en ed he se lemen by a popula ion
o la ge-bodied ac inop e ygians known om elsewhe e.
He e we desc ibe he i s skele al emains o la ge ac i-
nop e ygian ishes om he uppe Ca boni e ous o Nýřany,
consis ing o isola ed and agmen a y elemen s, o which a
single lowe jaw wi h well-p ese ed ee h is desc ibed as a
new genus and species, S ambe gich hys mac odens gen. e
sp. no . The mic o-compu ed omog aphy o his specimen
e eals he p esence o a p imi i e in e nal o ganiza ion o
he oo h den ine, which is discussed in he con ex o ecen
indings o pliciden ine in he Ac inop e ygii. He e desc ibed
ma e ials no only expand he knowledge on he di e si y o
he g oup a he locali y, bu also con ibu e signi ican ly o
he p oblema ic o aunal associa ions and palaeoen i on-
men a he Nýřany locali y, and expand he s a ig aphic
eco d o he la ge-bodied ac inop e ygians in he Cen al
and Wes Bohemian basins o he Czech Republic.
Ins i u ional abb e ia ions.—CGS, Czech Geological
Su ey, P ague, Czech Republic; NHMW, Na u his o isches
Museum Wien, Aus ia; NMP, Na ional Museum, P ague,
Czech Republic; ÚGV PAL, Palaeon ological collec ion o
he Depa men o Geological Sciences, Facul y o Science,
Masa yk Uni e si y, B no, Czech Republic.
Nomencla u al ac s.—This published wo k and he nomen-
cla u al ac s i con ains ha e been egis e ed in ZooBank:
u n:lsid:zoobank.o g:pub:24B92487-A962-474F-B9E2-
D819D0BCB203
Geological se ing
The Cen al and Wes Bohemian con inen al basins, ex end-
ing in he wes e n and cen al pa o he Czech Republic, ep-
esen he la e Ca boni e ous lu iolacus ine sedimen a y
in ill deposi ed on he Neop o e ozoic and ea ly Palaeozoic
Fig. 1. A. The s a ig aphic posi ion o he ac inop e ygian ma e ial desc ibed in p esen s udy. The s a ig aphic scheme is modi ied om Ma ínek e al.
(2017). B. Pe mo-Ca boni e ous con inen al basins o he Czech Republic wi h ma ked loca ion o Nýřany locali y.
B
Age Li hos a ig aphy
Global
s age
Regional
s age Subs age Fo ma-
ion
Coal
g oup
Asselian Au unian
C
Líně
B
Gzhelian
S ephanian
Sabe ian
Slaný
Kasimo ian
Ba ue-
lian
Týnec
Can a-
b ian
Kladno
Mosco ian
Wes phalian
As u ian
Bolso-
ian
Kladno
Kamenný Mos
Kouno
Ledce
Hředle
Mšec
Jelenice
Nýřany
Radnice
Klobuky
Zdě ín
Kouno
Mělník
Ne řeň
Cho íko
Nýřany
Touško
Lubná
Radnice
Plzeň
S a ig aphic hia us
A
50 km
S
P ague
Membe
S a ig aphic hia us
S a ig aphic hia us
S a ig aphic hia us
Duck-
man ian
BARTÁK ET AL.—ACTINOPTERYGIAN FISHES FROM THE UPPER CARBONIFEROUS OF CZECH REPUBLIC 503
basemen , and subdi ided in o Pilsen, Maně ín, Radnice,
Žihle, Kladno-Rako ník and Mšeno-Roudnice basins, he
li hos a ig aphic subdi ision o which ollows he same
gene al e minology (Fig. 1). In he Pilsen Basin, he old-
es li hos a ig aphic uni ep esen s he Kladno Fo ma ion,
subdi ided in o he Radnice and Nýřany membe s, be ween
which he s a ig aphic hia us ela ed o he Leonian Phase
o he Va iscan O ogeny, las ing abou 3.6 Ma, has been
documen ed (Opluš il e al. 2016; Ma ínek e al. 2017). The
deposi s o he Radnice Membe a e uncon o mably o e -
laid by he lu ial sedimen a ion o he Nýřany Membe ,
consis ing o basal conglome a es and domina ed by he
g ey-colou ed a coses, muds ones and clays ones, in he
Pilsen Basin a aining a hickness o abou 290 m (Pešek e
al. 1998, 2001). In addi ion o hese deposi s, in e cala ed
olcaniclas ics and g oups o coal seams occu ac oss he
Nýřany Membe . The zi con samples ob ained om he ol-
canogenic in e laye s ha e been 206Pb/238U da ed, p o iding
he absolu e age o he Nýřany Membe o 308.3–305.9 Ma
(Opluš il e al. 2016), co esponding o he la e Mosco ian
(As u ian) o ea ly Kasimo ian (Can ab ian) s age, i.e., he
bounda y o he Middle/Uppe Pennsyl anian. The nume -
ous coal seams in he Nýřany Membe , ep esen ing lacus-
ine deposi s, a e usually o ela i ely low hickness (up
o 2.5 m), and subdi ided in o Touško , Nýřany, Cho íko
and Ne řeň coal g oups (Pešek 1994). The Nýřany g oup o
coal seams consis s o wo, in some cases h ee, seams a -
aining a hickness o abou 1 me e , o which he lowe , i.e.,
he Main Nýřany Coal Seam, yielded abundan e eb a e
body- ossils. These a e de i ed om se e al sap opelic
ho izons a he base o he coal seam, and we e his o ically
documen ed in some coal mines (e.g., Humbold , K imich)
in he Nýřany and Třemošná coal ields (F i sch 1879). The
sap opelic ho izons de eloped wi hin he Main Nýřany Coal
Seam, which p oduced all e eb a e emains hus a doc-
umen ed a he Nýřany locali y, including he ma e ial de-
sc ibed he ein, indica e anoxic condi ions in s agnan wa e s
o a ela i ely shallow eshwa e lake (Milne 1980).
Ma e ial and me hods
The holo ype o S ambe gich hys mac odens gen. e sp. no .
(ÚGV PAL00174) is housed in he Palaeon ological collec-
ions o he Depa men o Geological Sciences, Masa yk
Uni e si y, B no (Czech Republic), and consis s o an iso-
la ed igh mandible wi h ee h in medial iew. The isola ed
and agmen a y elemen s o wo o he specimens om
he palaeon ological collec ions o he Na ional Museum,
P ague, Czech Republic, and he Na u his o isches Museum
Wien, Aus ia, a e e e ed o inde e mina e la ge ac inop e-
ygian ishes. These a e ep esen ed by he ollowing ma-
e ial: NMP M546, igh maxilla wi h ee h om la e al
iew and NHMW-Geo-2023/0311/0001, le clei h um om
medial iew. All specimens a e p ese ed on a slab o da k
coal shale coming om he uppe Ca boni e ous deposi s o
Nýřany, Czech Republic. In mos cases, he ma e ial con-
sis s o he mine alized bone issue o a ligh b own colou ,
which ep esen s a common ype o p ese a ion o he e -
eb a e body- ossils a he locali y. The holo ype o S. mac-
odens gen. e sp. no . was scanned wi h he high- esolu ion,
mic o-compu ed omog aphy de ice, GE phoenix | ome|x
L240, a he Cen al Eu opean Ins i u e o Technology
(CEITEC) in B no (Czech Republic). The ollowing pa am-
e e s we e used in he cou se o he specimen scanning:
he accele a ing ol age o 140 kV, he cu en o 220 µA,
he oxel size o 27 µm, 0.2 mm coppe beam il e . The
esul ing µCT slices we e manually segmen ed in a so wa e
ITK-SNAP . 3.8.0 (Yushke ich e al. 2006) in o de o ob-
ain he 3D su ace model. The su ace mo phology o he
ee h was imaged using lase scanning con ocal mic oscope
Keyence VK-X 1100 wi h he magni ica ion o 5× and he
ield o iew o 2730 µm; ull- ing ligh mode and lase
con ocal mode we e used. Fo imaging o mic o-mo pho-
logical de ails o uncoa ed oo h samples wi h a la ge ield
o iew and dep h o ield, he Ad anced en i onmen al
scanning elec on mic oscopy (A-ESEM) was used (Neděla
e al. 2020; S ela e e al. 2021). All A-ESEM expe i men s
we e pe o med a he Ins i u e o Scien i ic Ins umen s o
he Czech Academy o Sciences using modi ied mic oscope
QUANTA 650FEG (The mo Fishe Scien i ic). The obse -
a ion condi ions we e as ollows: wa e apou p essu e o
300 Pa, beam accele a ing ol age o 20 kV, a p obe cu en
o 80 pA, and a wo king dis ance o 12 mm. Mic og aphs
we e eco ded using wide ield ape u e de ec o WFAD o
A-ESEM (Bačo ský e al. 2022). Mac og aphic/la ge ield
images we e made by composing mic og aphs using Maps
so wa e (The mo Fishe Scien i ic). The addi ional mo -
phological de ails o he specimen we e imaged wi h Leica
DMC5400 (20 mpx) digi al came a connec ed o he Leica
MZ-16 s e eomic oscope.
Sys ema ic palaeon ology
Gna hos oma a Gegenbau , 1874
Os eich hyes Huxley, 1880
Ac inop e ygii Cope, 1887 sensu Good ich, 1930
Genus S ambe gich hys no .
ZooBankLSID: u n:lsid:zoobank.o g:ac :07AE2773-07C0-45AA-BE3B-
44FFB4C2DE0A
E ymology: In honou o S anisla Š ambe g (Museum o Eas e n Bo-
hemia, H adec K álo é, Czech Republic) o his long- e m esea ch on
he Pe mo-Ca boni e ous ac inop e ygians om he Eu opean basins.
Gende is masculine.
Type species: S ambe gich hys mac odens gen. e sp. no ., by mono-
ypy; see below.
Diagnosis.—As o ype and only known species.
S a ig aphic and geog aphic ange.—Mosco ian (Middle
Pennsyl anian) o Nýřany, nea Pilsen, Czech Republic.
504 ACTA PALAEONTOLOGICA POLONICA 69 (3), 2024
S ambe gich hys mac odens sp. no .
Figs. 2–5, 6A.
ZooBankLSID: u n:lsid:zoobank.o g:ac :9068DCBE-3FC8-4507-A250-
85EFEC9E7591
E ymology: F om G eek μακρóς (mak ós), meaning “long” o “la ge”,
and La in dens, i.e., “ oo h”; in e e ence o he conspicuous ee h o
he holo ype.
Holo ype: ÚGV PAL00174, an isola ed igh mandible wi h ee h expo-
sed in medial iew (Fig. 2A1).
Type locali y: Nýřany, coal mine 13 km sou hwes o Pilsen, Czech
Republic.
Type ho izon: Main Nýřany Coal Seam, Nýřany Membe , Kladno Fo -
ma ion, Pilsen Basin. A olcanic ash bed loca ed ca. 85 m abo e he
base o he Nýřany Membe has been 206Pb/238U da ed o 307.05 Ma ±
0.16 Ma, co esponding o he la es Mosco ian s age (As u ian sub-
s age) o he Middle Pennsyl anian (Opluš il e al. 2016).
Diagnosis.—A la ge ac inop e ygian ish (es ima ed o al
leng h 600–700 mm) dis inguished om all o he ea ly ac i-
nop e ygians by he ollowing unique combina ion o cha -
ac e s: well-de eloped pos e odo sal p ocess o mandible;
ex e nal de mal sculp u e consis s o anas omosing idges
and g oo es; single ow o la ge, smoo h, homodon , cone-
like ee h wi h bulbous bases; monocuspid ma ginal oo h
apices la e omedially comp essed and dis inc ly cu ed
medially; sub le mesial cu ing edge wi hou se a ion e-
s ic ed o apical po ion o ee h.
S ambe gich hys mac odens gen. e sp. no . di e s om
Ac olepis sedgwicki Agassiz, 1833, in mo e densely a anged
ee h la ge in size, and om Ac opholis s ensioei Aldinge ,
1937, and Plegmolepis kochi Aldinge , 1937, by la ge and
mo e obus ee h ewe in numbe . S ambe gich hys mac o-
dens gen. e sp. no . di e s om Ac olepis gigas (F ič, 1877)
by mo e widely ounded pos e o en al pa o he mandible,
deepe middle po ion o he mandible, angula no ex ending
a an e io ly, and by he p esence o anas omosing idges
and g oo es on he den a y ex e nal su ace. S ambe gich hys
mac odens gen. e sp. no . di e s om B azilich hys mac o-
gna hus Cox & Hu chinson, 1991, P ogy olepis specio sus
(F ič, 1875), P ogy olepis heyle i Poplin, 1999, Usclasich hys
mac odens Heyle , 1977, Zabo ich hys agmen alis Š am-
be g, 1989, and NMMNH P-77557 ( he unnamed la ge ac-
inop e ygian om Tinajas Membe , New Mexico; Ha is
and Lucas 2017) by homodon den i ion a anged in a single
ow close oge he , wi h sligh ly bulbous bases and medi-
ally ecu ed, la ened oo h ips. I u he di e s om
P ogy olepis speciosus, P ogy olepis heyle i, and U. mac o-
dens by smoo h su ace o ma ginal ee h, and om Z. ag-
men alis by well-de eloped pos e odo sal p ocess o man-
dible. S ambe gich hys mac odens gen. e sp. no . di e s
om B achydegma caela um Dunkle, 1939, by mo e closely
spaced ee h wi h bulbous bases, which ex end a pos e io ly
o he base o he pos e odo sal p ocess.
Desc ip ion.—Mandible: The holo ype o S ambe gich hys
mac odens gen. e sp. no . consis s o he isola ed igh man-
dible wi h ee h, exposed in he ock ma ix in medial iew
(Fig. 2A1). The p ese ed po ion o he mandibula amus
measu es 52 mm and i is incomple e an e io ly. Mos o
i s pos e io pa is p esen in he o m o mine alized bone
issue, al hough he small po ion co esponding o he pos-
e odo sal p ocess is p ese ed as an imp in o he la e al
su ace (Fig. 2A1). The mandibula amus is dis up ed by he
nume ous c acks, some o which a e obliquely di ec ed and
esul ed in bo h he do so en al and mediola e al displace-
men wi hin he jaw agmen . Simila ly, he bone medial
su ace o he mandible is damaged in he an e io egion and
along he en al ma gin o mos o he ee h, esul ing in he
exposu e o he in e nal s uc u es o he den a y (Fig. 2A1,
A3). The gene al o m o he mandible is boome ang-like,
wi h he pos e o en al bo de o he jaw smoo hly cu ed
do sally o o m a dis inc pos e odo sal p ocess o quad an-
gula shape, which is ele a ed abo e he ma ginal den i ion
o he mandible. The p ese ed po ion o he mandible is
ma kedly deep in he cen al pa , and g adually ape s in
bo h he an e io and pos e io di ec ions.
The majo i y o he lowe jaw is o med by he den a y.
I o ms he en i e an e io egion o he jaw la e ally and
ex ends pos e io ly sligh ly behind he le el o he ma ginal
ee h, whe e i con ac s he angula and su angula , al hough
he su u es canno be clea ly obse ed (Figs. 2A1–A3, 6A).
The la e al su ace o he den a y, imaged h ough he µCT,
e eals he p esence o a sculp u e consis ing o a sys em
o elonga e, anas omosing idges and g oo es (Fig. 2A2).
The o namen a ion ex ends along he en i e leng h o he
den a y and is con ined o i s cen al pa , being absen in
he en al, as well as do sal egions. I is cu en ly unclea ,
o wha deg ee his ea u e is a ec ed by he aphonomy and
he µCT p ocessing, o whe he he sculp u e dis ibu ion
ep esen s a na u al condi ion. The medial su ace o he
den a y is deeply exca a ed and smoo h o mos o i s ex en
(Fig. 2A1, A3). I has a p ominen medial wall jus en al o
he oo h ow, o which su ace is damaged and exposes he
in e nal s uc u es o he den a y, including he ascula ised
bone issue (bone o a achmen ), as well as a se o hin
canals pa ly illed wi h a whi e aluminosilica e seconda y
mine aliza ion. These canals a e in e p e ed he e as a pa
o he mandibula neu o ascula sys em inne ing he ee h
and he mandible (see below). The na ow Meckelian g oo e
is pa ly p ese ed in he an e o en al po ion o he den-
a y, ex ending mo e pos e io ly o o m a la ge Meckelian
ossa bo de ed do sally by a p ominen medial wall o he
bone. The angula o ms he caudal bo de o he mandibula
pos e odo sal p ocess, and cons ic s he la e al ex en o he
den a y, al hough he exac bounda y be ween he wo bones
is unclea (Figs. 2A2, A3, 6A). The la e al ex en o he angu-
la on he pos e odo sal p ocess o he mandible is appa en
om he imp in o i s la e al de mal sculp u e, consis ing o
hin and ela i ely densely a anged idges (Figs. 2A1). The
angula is con ined o he pos e io bo de o he p ocess and
is ex ended an e io ly in he do sal egion, while en ally i
has a subquad angula shape (Figs. 2A2, 6A). The smoo h
la e al su ace an e io o he angula in pos e odo sal pa
o he bone p obably co esponds o he su angula , and
BARTÁK ET AL.—ACTINOPTERYGIAN FISHES FROM THE UPPER CARBONIFEROUS OF CZECH REPUBLIC 505
A1
2
A
4
A
3
A
10 mm
Fig. 2. The ac inop e ygian ish S ambe gich hys mac odens gen. e sp. no . om Mosco ian, Middle Pennsyl anian, Nýřany Membe , Kladno Fo ma ion,
Nýřany, Czech Republic. The igh mandible o he holo ype, ÚGV PAL00174, as p ese ed in medial iew (A1). µCT su ace model o he mandible
in la e al (A2), medial (A3), and do sal (A4) iews. The g een colou in A3 and A4 ep esen s he sedimen a y in ill o neu o ascula mandibula canal.
Dashed lines wi h a ows indica e ac u es whe e displacemen has occu ed.

506 ACTA PALAEONTOLOGICA POLONICA 69 (3), 2024
ep esen s he a ea o o e lap wi h he pos e io egion o he
maxilla y pos o bi al pla e (Figs. 2A1, A2, 6A).
Den i ion: The ma ginal den i ion is implan ed in shal-
low dep essions o he ac odon o m and consis s o a single
ow o la ge, conical ee h (Fig. 2A4). The e is no e idence
on he p esence o a se o smalle la e al ee h common in
many o he ea ly ac inop e ygians (Poplin and Heyle 1993).
In addi ion o he nine mo e o less comple e ma ginal ee h,
se en mo e oo h posi ions a e p esen in he p ese ed po -
ion o he mandibula amus, making he o al coun no less
han six een ee h in he lowe jaw o S. mac odens gen. e
sp. no . The ee h a e homodon and se close oge he ; he
dis ance be ween he wo adjacen oo h posi ions is only
0.8 mm. The la ges ee h a e 3.5 mm high and 2 mm long
on he base. The dimensions o he ma ginal den i ion a e
ela i ely cons an h oughou he leng h o he jaw, and only
mino size dec easing is pe cep ible in he pos e io egion
o he oo h ow. The p opo ions o he ee h a e ela i ely
la ge in compa ison o he do so en al dep h o he jaw
amus, which eaches abou 2.5 o 3.5 imes o hei api-
co-basal heigh . The gene al o m o he ee h is cone-like
and esembles he la ge lania y ee h o o he Palaeozoic
p eda o y ac inop e ygians (Fig. 3A1–A13; Poplin 1999;
Š ambe g 2018, 2020). The ee h a e se pe pendicula o he
mandibula amus and a e up igh along hei en i e heigh .
The base is subo al in he c oss sec ion and sligh ly bul-
bous bo h mediola e ally and mesiodis ally. A sub le mesial
cu ing edge wi h no se a ion is con ined o he medially
cu ed uppe -mos pa o he oo h c own as a esul o
he la e al oo h comp ession (Fig. 3A3–A5, A12, A13). The
su ace o he ee h is smoo h h oughou hei heigh and
shows no p esence o idges o mic osculp u e in a o m o
ube cles and p o ube ances seen in some o he ac inop e-
ygian o ms (Rich e 1983; Poplin 1999; Š ambe g 2018,
2020). The smoo h enameloid laye is p esen in he speci-
men and occupies abou 40% o he oo h heigh in he apical
pa (Fig. 3A1, A2, A6–A11). I s b igh su ace o ms a sha p
ansi ion be ween i and he colla enamel o he oo h sha ,
and can be seen as a e y hin, supe icial laye in he µCT
ans e sal and axial slices o he ee h (Figs. 3A15, 4A6;
Ge main e al. 2016). The in e nal s uc u e o he ma ginal
den i ion, unco e ed by he µCT, shows he p esence o a
la ge pulp ca i y su ounded by a mode a ely hick laye o
he den ine (Figs. 3A15, 4A1–A5). In he c oss sec ion, his
den ine laye displays he p esence o a simple o m o plici-
den ine, which is es ic ed o he e y base o he ee h and
canno be obse ed in any o m (e.g., g oo es and idges) on
he ex e nal su ace o he implan ed ee h. The indi idual
den ine olds a e no ami ied and do no ex end o he cen-
e o he la ge pulp ca i y o he ee h. This simple o m o
he den ine in olding co esponds well o he simplexodon
ype o pliciden ine epo ed ecen ly in some ac inop e y-
gian and sa cop e ygian ishes (Meunie e al. 2015a, b).
Neu o ascula sys em: The X- ay µCT imaging ech-
niques e ealed inside he mandibula amus o S. mac o-
dens gen. e sp. no . he p esence o a slende bony canal,
loca ed immedia ely below he ma ginal den i ion, which is
in e p e ed he e as a passage o he mandibula b anch o
he igeminal ne e (V, amus mandibula is igemini; Figs.
2A1, A3, 3A14–A16, 5). The con inui y o he mandibula ca-
nal has been dis up ed by he nume ous ac u es and sho
displacemen s ha occu in he den a y o he specimen. The
descending b anch o he mandibula ne e, coming om he
igeminal ganglion, p obably en e ed he mandibula amus
o S. mac odens gen. e sp. no . in i s pos e io egion, as
epo ed in some ex an lowe ac inop e ygians (Allis 1922;
Pio owski and No hcu 1996). The pos e io -mos po ion
o he p ese ed pa o he mandibu la canal spli s in o ou
o i e subpa allel b anches which mo e an e io ly, a he
le el o he pos e io -mos ee h, pass in o he la e al amus o
he mandibula canal. The la e al amus al e s in mo phology
om nodula o od-like along i s cou se, and is accompanied
by he subpa allel medial amus in he pos e io pa o he
mandible, which e ains he o m o a simple ba h oughou
i s leng h. Bo h ami me ge a he le el o he en h pos e i-
o -mos oo h posi ion o o m a single main amus ha passes
in o he mo e an e io po ion o he mandible. I s o m is well
exposed in he longi udinal sec ion a he medial su ace o
he den a y due o he b eakage o he bone (Figs. 2A1, A3,
5A4). Bo h ami o he mandibula canal pos e io ly, as well
as a single main amus an e io ly, spli in o se e al b anches
owa ds he den a y ee h o o m a den al plexus, h ough
which he ne es and blood essels passed, supplying he
pulp ca i ies. In he an e io po ion o he jaw agmen , a
obus b anch o he mandibula neu o ascula canal ex ends
pos e ola e ally om i s main unk and e mina es in se e al
smalle b anches leading o he po es on he la e al su ace o
he den a y which migh ha e inne ed he ex e nal su ace
o he mandible. No mandibula senso y canal o he la e al
line sys em (Leuzinge e al. 2020) could be clea ly obse ed
in he den a y o S. mac odens gen. e sp. no ., al hough a ew
na ow, elonga e, pa allel canaliculi p esen in he mo e en-
al posi ion o he jaw amus migh be ela ed o he cephalic
senso y sys em.
Rema ks.—The agmen a y na u e o he ype specimen
o S. mac odens gen. e sp. no . me i s de ailed accoun o
i s ac inop e ygian cha ac e is ics be o e he axonomic as-
signmen a he speci ic le el and compa ison o o he axa
can be add essed. The hin enameloid laye con ined o he
apical po ion o he ee h ep esen s he only ac inop e y-
gian synapomo phy p esen in he holo ype o S. mac odens
gen. e sp. no . (Ø ig 1978b; Pa e son 1982). Al hough a
numbe o o he cha ac e is ics o he jaw agmen ha e
a wide dis ibu ion among he ea ly os eich hyians, all o
hese ea u es occu o a ying deg ees in ea ly ac inop e y-
gians, u he suppo ing i s a ibu ion o his g oup. These
include (i) simplexodon pliciden ine a he oo h base, (ii)
mandibula amus do so en ally deep in pos e io pa , (iii)
smoo hly cu ed pos e o en al ma gin o mandible, (i )
p ominen pos e odo sal p ocess o quad angula shape, ( )
la e ally la ened mandibula amus, ( i) ex e nal de mal
sculp u e o med by elonga ed idges and g oo es.
BARTÁK ET AL.—ACTINOPTERYGIAN FISHES FROM THE UPPER CARBONIFEROUS OF CZECH REPUBLIC 507
A hin enameloid laye is clea ly p esen in he den i ion
o S. mac odens gen. e sp. no . (Figs. 3A1, A2, A15, 4A6) and,
unlike chond ich hyans whe e he enameloid co e s mos o
he oo h sha (e.g., Jambu a e al. 2019), i is con ined o he
apical po ion o he ee h, o ming he cha ac e is ic ac o-
din caps o ac inop e ygians. Fu he mo e, he simplexodon
o m o pliciden ine is p esen a he base o he ma ginal
ee h, which is now known o be widely dis ibu ed among
4
A
5
AA
1 2
A
3
A
250 mµ
1 mm
6
A
7
A8
A
3
A
9
A10
A11
A
12
A
13
A
14
A15
A16
A
17
A
14
A15
A
1
A2
A
16
A
1 mm
1 mm
1 mm 1 mm 1 mm
800 mµ
200 mµ100 mµ100 mµ
10 mm
1 mm 1 mm 1 mm
Fig. 3. The ac inop e ygian ish S ambe gich hys mac odens gen. e sp. no . om Mosco ian, Middle Pennsyl anian, Nýřany Membe , Kladno Fo ma ion,
Nýřany, Czech Republic. Ma ginal den i ion and ans e sal sec ions o he igh mandible o he holo ype, ÚGV PAL00174. Pho og aphs o he ma ginal
ee h in medial iew (A1, A2). An e io -mos p ese ed oo h showing apically es ic ed cu ing edge in mesial iew (A3, A4, A5). Close up images om
A-ESEM showing apical po ion o he ma ginal ee h in medial iew (A6, A7, A8). Mo phology o he well-p ese ed an e io ma ginal ee h om A-ESEM
in medial iew (A9, A10, A11). 3D model o he hi d an e io -mos p ese ed oo h displayed h ough he con ocal mic oscopy showing mesial cu ing edge
(A12, A13). T ans e sal sec ions h ough he mandibula amus (A14, A15, A16). In e p e a i e d awing o he mandible showing he cou se o slices and de ails
o he ee h depic ed in o he igu es (A17). Abb e ia ions: c , cu ing edge; en, enameloid; mc, mandibula neu o ascula canal; mcb, mandibula neu o ascu-
la canal b anch; Mg, Meckelian g oo e; pc, pulp ca i y; pmcb, pos e io mandibula neu o ascula canal b anches. Scale ba s o A12 and A13 no a ailable.
508 ACTA PALAEONTOLOGICA POLONICA 69 (3), 2024
he a ious ac inop e ygian o ms (see Vi iani e al. 2022 o
he ecen summa y), al hough no necessa ily con ined o
hem (Meunie e al. 2015b). The ma ginal ee h a anged in
wo se ies ha e been in e p e ed o ep esen a plesiomo phic
condi ion o he ac inop e ygians, wi h he la ge, cone-like
ee h placed medially in espec o he diminu i e la e al
den i ion (Poplin and Heyle 1993; Lund and Poplin 1997).
Al hough his cha ac e is ic indeed equen ly occu s in he
Palaeozoic o ms, he p esence o a single ma ginal oo h ow
p esen in S. mac odens gen. e sp. no . does no p eclude i s
a ibu ion o Ac inop e ygii, as i is cu en ly known om
many o he ea ly ac inop e ygian gene a (e.g., Dunkle 1939;
Lund 2000; Poplin and Lund 2000; Hamel 2005; Š ambe g
2016; S ack e al. 2021; A gy iou e al. 2022), and is consid-
e ed o ep esen an apomo phic s a e o his g oup (Lund
and Poplin 1997). The mandible o S. mac odens gen. e sp.
no . is deepes in i s pos e io pa in on o he pos e o-
do sal p ocess, and i g adually na ows bo h an e io ly and
pos e odo sally along he smoo hly cu ed en al ma gin
o he bone (Figs. 2, 6A). The same gene al o m o he
lowe jaw can be seen in many Palaeozoic ac inop e ygians,
and he esemblance is especially e iden in axa such as
Pa a a asius hibba di Lund & Mel on, 1982, “Elonich hys”
hypsilepis Hay, 1900, P ogy olepis heyle i, Eu yno us c e-
na us Agassiz, 1835, and B achydegma caela um, in which
he p onounced pos e odo sal p ocess is well exposed (Lund
and Mel on 1982; Schul ze and Ba dack 1987; Poplin 1999;
Š ambe g 2018; F iedman e al. 2019; A gy iou e al. 2022).
Mo eo e , he mandible is ma kedly la ened mediola e -
ally, as p esen in ea ly os eich hyians (Bo ella e al. 2007;
Zhu e al. 2009), as well as sa cop e ygian (e.g., Je e y 2002;
Po o e al. 2015a) and ac inop e ygian ishes (e.g., A a ia
and Clou ie 1996; Giles e al. 2015; Figue oa e al. 2019),
bu absen in ea ly e apods (Ahlbe g and Clack 1998; Po o
e al. 2015b). In ea ly ac inop e ygians, he o m o de mal
sculp u e on he ex e nal su ace o he jaws and skull oo
a ies mos commonly om he g anula o idged, bo h
capped wi h a ganoine laye (Ø ig 1978a, b). The idges may
be ela i ely coa se and p onounced (e.g., Lund 2000; Mickle
e al. 2009), o a he ine and e micula , as in P ogy olepis
(Š ambe g 1991, 2018; Poplin 1999). Al hough he p esence
o he ganoine canno be eadily con i med in he mandible
o S. mac odens gen. e sp. no ., he de mal sculp u e on he
den a y con o ms o he i s pa e n, whe eas he second
condi ion is p esen on he ex e nal su ace o he angula
(Fig. 2A1). All he cha ac e is ics discussed abo e he e o e
indica e ha he holo ype o S. mac odens gen. e sp. no .
ep esen s an ac inop e ygian ish.
The mandible o S. mac odens gen. e sp. no . is in gen-
e al o m simila o ha o o he la ge-bodied p eda o y gen-
e a like B azilich hys mac ogna hus (Cox and Hu chinson
1991; Figue oa e al. 2019; Figue oa and And ews 2023),
B
4
A5
A6
A
A1
2
A
3
A
7
A
A1
2
A3
A
6
A
5
A
4
A
3 mm
3 mm
3 mm
1 mm 1 mm
1 mm
Eus henop e on
Chei olepis
Polyp e us
Hoplias
A apaima
S ambe gich hys
Sa cop e ygii
Ac inop e ygii
Teleos ei
10 mm
Fig. 4. The ac inop e ygian ish S ambe gich hys mac odens gen. e sp. no . om Mosco ian, Middle Pennsyl anian, Nýřany Membe , Kladno Fo ma ion,
Nýřany, Czech Republic. C oss sec ion h ough ma ginal den i ion o he holo ype, ÚGV PAL00174, showing he mo phology and dis ibu ion o plici-
den ine. A. µCT axial slices a di e en le els o ma ginal ee h (A1, A2, A3). In e p e a i e d awings o c oss sec ions a he base (A4), in he middle (A5)
and close o apical ip (A6) o a single oo h. In e p e a i e d awing o he specimen showing he cou se o slices in o he igu es (A7). B. Phylogene ic
diag am showing dis ibu ion and mo phology o pliciden ine in ma ginal ee h o selec ed ex inc and ex an ac inop e ygian ishes. Abb e ia ions:
de, den ine; d , den ine old; en, enameloid; pc, pulp ca i y; b, ascula ised bone (bone o a achmen ).
BARTÁK ET AL.—ACTINOPTERYGIAN FISHES FROM THE UPPER CARBONIFEROUS OF CZECH REPUBLIC 509
B achydegma caela um (Dunkle 1939; A gy iou e al.
2022), Ras olepis iojaensis López-A ba ello e al., 2006,
and ac olepids (Aldinge 1937; Ga dine 1963; Š ambe g
1991, 2018; Poplin 1999; López-A ba ello e al. 2010), ha -
ing elonga ed mandibula amus ape ing an e io ly, and he
p onounced pos e io a icula ing egion ele a ed abo e he
ma ginal oo h ow. The well-p ese ed mandibles om he
lowe Pe mian o F ance and B azil e eal ha he p omi-
nen pos e odo sal egion o he lowe jaw in P ogy olepis
heyle i and B azilich hys mac ogna hus consis s o bo h
angula and su angula , whe eas he den a y is pos e io ly
ounded and is no in ol ed in i s cons uc ion (Š ambe g
2018; Figue oa and And ews 2023). Al hough he su u es
a e di icul o be aced in S. mac odens gen. e sp. no .,
he cong uen a angemen o hese elemen s can also be
assumed o his egion (Fig. 6A).
The p ese ed po ion o he jaw agmen o S. mac o-
dens gen. e sp. no . measu es 52 mm. Howe e , he mandi-
ble is clea ly incomple e an e io ly, and we assume ha abou
one-qua e o one- hi d o he lowe jaw migh be missing
(Fig. 6A). I his es ima ion is co ec , hen he comple e man-
dible o he specimen could ha e eached abou 70–80 mm
in leng h. Based on he simila ly sized lowe jaws, Š ambe g
(1991) and Poplin (1999) es ima ed he o al body leng h o
P ogy olepis a abou 600–700 mm. Al hough no in o ma-
ion on body p opo ions, such as he a io o skull leng h
o o al body leng h, a e a ailable in S. mac odens gen. e
sp. no ., we assume he es ima ed leng h o he specimen
could be oughly simila o ha o P ogy olepis speciosus
and P ogy olepis heyle i. In he Pe mo-Ca boni e ous con i-
nen al basins o he Czech Republic, se e al ac inop e ygian
species eaching a size compa able o S. mac odens gen. e
sp. no . a e known, including Ac olepis gigas, P ogy olepis
speciosus, and Zabo ich hys agmen alis, he ma e ial all o
which was i s hand e-examined o compa ison pu poses.
The lec o ype o Ac olepis gigas is an almos comple e spec-
imen p ese ed as pa and coun e pa in a la ge side -
i e conc e ion, which includes, among o he s, he skull and
mandible, and measu es 1250 mm in o al leng h (Š ambe g
1991). The mandible o A. gigas shows he gene al plesio-
mo phic mo phology sha ed wi h simila ac inop e ygians
o easonable size, bu is much la ge compa ed o ha o S.
mac odens gen. e sp. no ., eaching a o al leng h o a leas
120 mm (Fig. 6B; Š ambe g 2006). The mandibula amus
is e y slende and s aigh an e io o he pos e odo sal
A
12
A
4
A
3
A
10 mm
B
A
10 mm
20 mm
Fig. 5. The ac inop e ygian ish S ambe gich hys mac odens gen. e sp. no . om Mosco ian, Middle Pennsyl anian, Nýřany Membe , Kladno Fo ma ion,
Nýřany, Czech Republic. Neu o ascula sys em in he igh mandible o he holo ype, ÚGV PAL00174, displayed h ough he µCT. The mandible in medial
(A1), la e al (A2), and do sola e al (A3) iews. Recons uc ion o he mandibula neu o ascula sys em in la e al iew (A4). G een colou ep esen s he
canals o mandibula neu o ascula sys em, yellow colou ma ks he posi ion o la e al po es. Abb e ia ions: dp, den al plexus; mcb, mandibula neu o as-
cula canal b anch; m , mandibula canal amus; pca, pa allel canaliculi; pmcb, pos e io mandibula neu o ascula canal b anches.
Fig. 6. The compa ison o he mandible o ac inop e ygian ishes (A) S am-
be gich hys mac odens gen. e sp. no . (Mosco ian, Middle Pennsyl anian,
Nýřany Membe , Kladno Fo ma ion, Nýřany, Czech Republic) and (B) Ac o-
lepis gigas (F ič, 1877) (Gzhelian, Uppe Pennsyl anian, Mšec Membe ,
Slaný Fo ma ion, Žilo , Czech Republic). A. ÚGV PAL00174, holo ype,
he econs uc ion o he igh mandible ( e e sed) in la e al iew. B. NMP
M125, lec o ype, he pho og aphy o he le mandible in la e al iew.
516 ACTA PALAEONTOLOGICA POLONICA 69 (3), 2024
acco ding o Lopez e al. (2008), he ish emains a e de i ed
om he lowe mos pa o he Tuilliѐ es-Loi as Fo ma ion,
ep esen ed by he black shale deposi s, indica ing a deepwa-
e lacus ine en i onmen wi h anoxic condi ions nea he
bo om (Lopez e al. 2008; Pocha and Van Den D iessche
2011). Finally, he medium-sized (<400 mm o al leng h) elon-
ich hyid Rhabdolepis mac op e us is known om he lowe
Pe mian limnic deposi s o he Saa -Nahe Basin, Ge many
(Ga dine 1963; Schindle 2018b). I s equen occu ence
has been documen ed om he clay i ons one acies o he
Humbe g Black Shale, which a e gene ally conside ed o con-
ain he au och honous auna in he deepes pa o he ex en-
si e Humbe g-Lebach Lake (Boy 1987; Schoch 2009, 2014).
Thus, based on he o e iew p esen ed abo e, i could be
concluded ha he la ge eshwa e ac inop e ygians a e com-
monly ound in deepwa e en i onmen s ac oss he Pe mo-
Ca boni e ous con inen al basins in he cen al Pangea
(Fig. 8). Al hough he occu ence o such ishes a Nýřany
indeed could indica e a p esence o deep and ex ensi e wa e
o body, as p e iously sugges ed by Milne (1980), we no e
ha palaeobiological e idence alone is ambiguous and insu -
icien o in e p e deposi ional en i onmen , and o he lines
o e idence should be aken in accoun .
As no ed abo e, sedimen ological da a suppo nei he
he p esence o deepwa e en i onmen la e ally o he shal-
low swamp lake, no subsidence o he basin, allowing he
accumula ion o a la ge wa e column. Despi e his, an al-
loch honous aqua ic e eb a e associa ion appa en ly oc-
cu s a he locali y (Table 1). These axa can be ecognized
based on he c i e ia es ablished by Milne (1980), includ-
ing he ela i e abundance o specimens, deg ee o hei
comple eness and a icula ion, unc ional mo phology, as
well as compa ison o o he aunal associa ions om else-
whe e. The la ge xenacan hi o m sha k O hacan hus bo-
hemicus has been desc ibed a he locali y om some skull
and pos c anial skele on ma e ials, isola ed do sal spines and
nume ous isola ed ee h (F i sch 1889; Heid ke 1998; Sole -
Gijón 2004). P e ious au ho s no ed he wide geog aphic and
s a ig aphic dis ibu ion o his and o he xenacan hi o m
gene a ac oss he Pe mo-Ca boni e ous limnic basins o he
Cen al and Wes e n Eu ope, and p oposed a palaeogeo-
g aphical scena io o hei con inen al dispe sion h ough he
i e channels in e connec ing hese basins (Schneide and
Zajíc 1994; Schneide 1996; Luccisano e al. 2022). This is
suppo ed by he ecen iso opic analyses o xenacan hi o m
ee h and spines (Fishe e al. 2013; Luccisano e al. 2023),
indica ing ha la ge-bodied o ms like O hacan hus we e
es ic ed o la ge and deep eshwa e lakes and i e sys-
Table 1. The lis o aqua ic axa conside ed in his s udy o be alloch honous a he locali y o Nýřany (Czech Republic).
Taxon Ma e ial Re e ences
Xenacan hi o mes
O hacan hus bohemicus F ič, 1875
wo ai ly comple e ju enile indi iduals; a icula ed skull
wi h mandibles and pec o al gi dle; mandible wi h skull
elemen s; pel ic gi dle; nume ous isola ed spines and ee h
F i sch (1889, 1907); Heid ke
(1998); Sole - Gijón (2004)
Ac inop e ygii
Ac inop e ygii inde . isola ed maxilla wi h in ao bi al, clei h um his s udy
S ambe gich hys mac odens gen. e sp. no . isola ed mandible wi h ee h his s udy
Dipnoi
Sagenodus sp. isola ed scales F i sch (1888); Wa son and Gill
(1923)
Te apoda
Baphe es o ien alis Milne e al., 2009 wo a icula ed skulls, mandibles and pec o al gi dle S een (1938); Beaumon (1977);
Milne e al. (2009)
Diplo e eb on punc a um F ič, 1879 disa icula ed c anial and pos c anial elemen s F i sch (1885); Ca oll (1970);
Klemba a e al. (2014)
moun ains
land mass
shallow sea p esen
absen
la ge-bodied
acnop e ygians
7
1
2
4
6
5
3
Pangea
Paleo e hys
Fig. 8. The la e Ca boni e ous (Pennsyl anian) palaeogeog aphic map
showing he occu ence o la ge-bodied ac inop e ygians in selec ed esh-
wa e o ma ine-in luenced locali ies wi hin con inen al basins o cen al
Pangea. 1, Nýřany, Czech Republic; 2, Mšec, Czech Republic; 3, Newsham,
No humbe land, UK; 4, Mon ceau-les-Mines, F ance; 5, Lin on, Ohio,
USA; 6, Mazon C eek, Illinois, USA; 7, Tinajas, New Mexico, USA.

BARTÁK ET AL.—ACTINOPTERYGIAN FISHES FROM THE UPPER CARBONIFEROUS OF CZECH REPUBLIC 517
ems. Simila ly, he dipnoan ish Sagenodus sp., known in
Nýřany om se e al isola ed scales (F i sch 1888; Wa son
and Gill 1923), has been documen ed no only om a ious
Eu opean con inen al basins (F i sch 1888; Schul ze 1993;
Oli e e al. 2012; Beeby e al. 2020), bu also om nume ous
locali ies in he USA (Schul ze and Cho n 1997). The la e
au ho s p oposed ha he genus Sagenodus was eu yhaline
and sp ead om he No h Ame ica o Eu ope ia a ma ine
connec ion, ollowed by i s dispe sal ac oss he Eu opean
con inen al basins h ough he i e channels. I s emains
a e equen ly ound a Newsham (Beeby e al. 2020) and
occu , among o he s, in he Mšec and Kouno membe s
(F i sch 1888; Lojka e al. 2009), indica ing he genus migh
ha e p e e ed a deepwa e en i onmen , al hough no being
necessa ily es ic ed o i . The aqua ic specializa ions o he
baphe oids, in Nýřany ep esen ed by Baphe es o ien alis,
a e well suppo ed by he a ious mo phological adap a-
ions on he skull and mandible (Beaumon 1977; Beaumon
and Smi hson 1998). Wes oll (1944) no ed ha membe s o
he g oup o en co-occu wi h haplolepids in deposi s in-
e p e ed as s agnan shallow wa e s, while Milne (1987)
assumed he baphe oids inhabi ed coas al b ackish deepwa-
e en i onmen s, al hough he poin ed o hei possible wide
ange o ecological adap a ions. Al hough he Baphe oidea
ep esen s a a e aunal componen in bo h he Lin on (Hook
and Bai d 1986) and Nýřany (Milne 1980) locali y, he
comple e a icula ed skulls ha e been epo ed om bo h
si es (Rome 1930; S een 1938; Beaumon 1977; Milne e al.
2009), indica ing ha hei ma e ials did no unde go a long
anspo o he place o deposi ion. We hypo hesize ha he
sca ce occu ences o he baphe oids in Nýřany e lec hei
ecological p e e ences, and hese o ms migh ha e indeed
p ima ily inhabi ed he i e sys ems, wi h only occasional
excu sions o he ela i ely shallow lake o o age o b eed.
Finally, Diplo e eb on punc a um is cu en ly known om
a single incomple e specimen (F i sch 1885: pl. 52) which
ep esen s sca e ed skele al emains o a possible embo-
lome e (Ca oll 1970; Klemba a e al. 2014). The na u e o
i s p ese a ion, as well as spa se occu ence, indica e i is
alloch honous a he locali y (Milne 1980), al hough oo
incomple e o in e i s possible habi a . The alloch honous
aqua ic e eb a e associa ion documen ed in deposi s o he
ela i ely shallow lake o Nýřany hus does no equi e he
p esence o a nea by la ge and deep eshwa e lake, as p e-
iously hypo hesized (Milne 1980), bu i can equally be in-
e p e ed as o igina ing om he b aided i e sys em known
o occu ac oss he Nýřany Membe (Opluš il e al. 2005). In
con ex o his, he mandible o S ambe gich hys mac odens
gen. e sp. no . and emains o he same o o he la ge ac i-
nop e ygian ishes in he locali y a e in e p e ed he e also as
de i ed om a i e channel ha could ha e been connec ed
o he shallow lake o some ime (Pešek 1994; Pešek e al.
1998). I is also in e es ing o poin ou he e ha , al hough
la gely incomple e, he alloch honous aqua ic e eb a e as-
socia ion o Nýřany seems o esemble in gene al compo-
si ion o some o he Ca boni e ous locali ies in e p e ed as
deepwa e en i onmen s (Fig. 9). This aunal conco dance
can be pe cei ed as pa ly a i icial and e lec ing a wide dis-
ibu ion o i s cons i uen s in a ious deposi ional se ings.
In o he wo ds, ce ain aqua ic e eb a es (e.g., xenacan-
hi o m sha ks, ac inop e ygian and dipnoan ishes) migh
ha e used he i e channels p ima ily o each he mo e sui -
able des ina ions such as deep eshwa e basins, whe e hei
occu ence is indeed well-documen ed. This migh also be
he case o la ge-bodied ac inop e ygians like P ogy olepis,
i.e., P ogy olepis speciosus and P ogy olepis heyle i, which
ha e been eco ded in he Pe mo-Ca boni e ous basins o
he Czech Republic (Š ambe g 1991, 2013; Lojka e al. 2009),
1
4
5
3
2
1
2
3
4
5
6
7
8
4
3
2
1
5
3
4
6
7
5
S epsodus sau oides
O hacan hus gibbosus
Pholide pe on a heyi
Megalocephalus pachycephalus
Ac olepis hopkinsi
Sagenodus inaequalis
Acan hodopsis wa di
pla ysomids
Rhabdode ma elegans
Ac inop e ygii inde .
O hacan hus bohemicus
Baphe es o ien alis
Diplo e eb on punc a um
S ambe gich hys mac odens
Sagenodus sp.
O hacan hus kouno iensis
Ac olepis gigas
P ogy olepis speciosus
Zabo ich hys agmen alis
Sagenodus sp.
Plica odus plica us
Acan hodes i schi
small-bodied ac inop e y
gians
7
8
6
1
2
3
5
4
2
1
5
3
6
7
Mšec
Newsham Nýřany
4
1
2
Bashki ian Mosco ian Gzhelian
Fig. 9. The compa ison o Pennsyl anian aqua ic e eb a e associa ions o deepwa e en i onmen (Newsham and Mšec) wi h alloch honous aqua ic
e eb a e associa ion o Nýřany, in e p e ed he e as o igina ing om b aided i e sys em. Black silhoue es ep esen sha ed aunal componen s a he
genus le el. Faunal lis s based on Lojka e al. (2009) o Mšec, on a ious sou ces o Newsham (no e ha lis ed auna o Newsham is no exhaus i e).
518 ACTA PALAEONTOLOGICA POLONICA 69 (3), 2024
F ance (Poplin 1999; Š ambe g 2018; Š ambe g and S eye
2021) and Spain (Sole -Gijón and Díez Ruiz 2023). The ab-
sence o S. mac odens gen. e sp. no . om o he con inen al
basins o Eu ope may hus be ela ed ei he o i s endemism,
sampling bias, o he simple lack o Eu opean basins o he
co esponding age.
The den i ion o S. mac odens gen. e sp. no . consis s o
la ge, massi e, uni o m ee h wi h poin ed ips and smoo h
su ace, which a e a anged in a single ow, and clea ly dis-
play he ca ni o ous adap a ions. The well-de eloped sim-
plexodon o m o pliciden ine, ound in all ee h o he jaw,
indica es he den i ion was adap ed o wi hs and he high
p essu e loads associa ed wi h p ey cap u ing. These mo -
phological and s uc u al ea u es, combined wi h he ela-
i ely la ge dimensions o he species (es ima ed o al leng h
up o 700 mm), suppo he in e p e a ion i occupied high
ophic le els in he ecosys ems i inhabi ed. Howe e , he
emains o much la ge ac inop e ygians a e documen ed in
Nýřany (NHMW-Geo-2023/0311/0001; he es ima ed o al
body leng h p obably no less han 1200 mm). Due o he
lack o o e lapping ma e ial, i cu en ly canno be esol ed
whe he his ma e ial can be a ibu ed o S. mac odens gen.
e sp. no ., o i ep esen s ano he so a unknown la ge-
bodied species. In any case, his o m would ha e ep e-
sen ed a gigan ic ac inop e ygian ish likely occupying op-
mos posi ions o he ood web (Fig. 9), and p obably able o
compe e wi h la ge-bodied xenacan hi o m sha ks and ea ly
e apods co-occu ing a he locali y.
Conclusions
The isola ed skele al emains o la ge ac inop e ygian ishes
a e epo ed he e o he i s ime om he Mosco ian,
Middle Pennsyl anian o Nýřany locali y, Czech Republic.
These include a new genus and species, S ambe gich hys
mac odens gen. e sp. no ., based on a single lowe jaw
wi h well-p ese ed ee h, which has been scanned h ough
he mic o-compu ed omog aphy and shows a p esence o a
complex in e nal neu o ascula sys em inne ing he lowe
jaw and ee h, and supplying bo h wi h blood essels. The
ma ginal den i ion o he new species exhibi s a single ow
o massi e, homodon and conical ee h wi h he p esence
o simplexodon pliciden ine on hei base, he ea u es ha
suppo i s p eda o y ecology. The occu ence o isola ed and
agmen a y emains o la ge-bodied ac inop e ygians in he
coal deposi s o a ela i ely shallow lake indica es ha hei
emains we e p obably de i ed om he b aided i e sys em
lowing h ough he allu ial plain o he Nýřany Membe ,
and ep esen he oldes s a ig aphic eco d o la ge ac i-
nop e ygian ishes in he Pe mo-Ca boni e ous sedimen s
o Czech Republic. These esul s unde line a po en ial axo-
nomic and palaeoecological impo ance o some isola ed and
agmen a y specimens ound in coal-bea ing deposi s o he
Cen al and Wes e n Eu ope, being a pa o he classic and
well-known la e Ca boni e ous e eb a e assemblages.
Acknowledgemen s
We a e indeb ed o Nela Doláko á (Masa yk Uni e si y, B no, Czech
Republic), U sula Göhlich (NHMW), Bo is Ek (NMP), and Pe
Budil (CGS) o he pe mission o s udy he ma e ials in hei ca e
and assis ance in he collec ions. S anisla Š ambe g (Museum o
Eas e n Bohemia, H adec K álo é, Czech Republic), F ede ik Spindle
(Dinosau ie Museum Al mühl al, Denkendo , Ge many) and A jan
Mann (Na ional Museum o Na u al His o y, Smi hsonian Ins i u ion,
Washing on, USA) a e hanked o he ui ul discussion on he ax-
onomic assignmen o he Nýřany ma e ial. We a e also hank ul o
Jakub Sakala and S anisla Opluš il (bo h Cha les Uni e si y, P ague,
Czech Republic), as well as Ja osla Zajíc (Czech Academy o
Sciences, P ague, Czech Republic) o he discussion on he palaeo-
en i onmen o he Nýřany locali y. Valuable commen s om wo e-
iewe s, Ad iana López-A ba ello (Ludwig-Maximilians-Uni e si ä
München, Ge many) and Sam Giles (Uni e si y o Michigan, USA),
helped o g ea ly imp o e he ini ial d a o he manusc ip . This s udy
was suppo ed by he Speci ic Resea ch P ojec MUNI/A/1261/2022
o he Facul y o Science a Masa yk Uni e si y in B no and he Czech
Science Founda ion (g an no. GA22-25799S).
Au ho con ibu ions’
PB, concei ed he s udy, conduc ed analyses, w o e he ex
and p epa ed igu es; MI, concei ed and supe ised he
s udy, w o e he ex ; ET, MO, conduc ed analyses; VN, con-
duc ed analyses and w o e he ex .
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