A new extinct shad from Poland in the light of clupeiform diversity and distribution within the Paratethys during the Oligocene

Abstract

The Order Clupeiformes Bleeker, 1859 comprises herrings, anchovies, sprats, sardines, and shads. The fossil record of this group is rich within the Paratethys. Here we describe a new clupeiform fish, †Sanalosa janulosa gen. et sp. nov., from the Lower Oligocene of the Carpathian Basin, Poland. This new genus has a unique combination of characters (lower jaw articulation located under the posterior part of the orbit; abdominal scutes well developed with 3 to 5 in the gular region, 11–14 prepelvic scutes associated with ribs, 11–12 postpelvic scutes; several striae on the frontals; an opercle with 6–12 thin radial ridges; a horizontal ramus of the preopercle shorter than the vertical one; 42–44 vertebrae; 8–10 supraneurals; a dorsal fin with 18–22 rays, and an anal fin with 21–23 rays), supporting recognition of a new genus and species within the Family Alosidae Svetovidov, 1952. Similarities and differences between fossil and extant genera of the Clupeiformes are discussed to shed more light on their relationship. Moreover, the palaeobiogeography, diversity and distribution of Oligocene clupeiform fishes in the Paratethys are presented and discussed.

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© 2024 Malgorzata Bienkowska-Wasiluk, Mateusz Granica and Oleksandr Kovalchuk. This is an open access
article under the CC BY license (http://creativecommons.org/licenses/by/4.0/), which permits use, distribution,
and reproduction in any medium, provided that the article is properly cited.
Acta Geologica Polonica, Vol. 74 (2024), No. 3, e23
DOI: 10.24425/agp.2024.151753
A new extinct shad from Poland in the light of clupeiform
diversity and distribution within the Paratethys
during the Oligocene
MALGORZATA BIENKOWSKA-WASILUK1, MATEUSZ GRANICA1 and OLEKSANDR KOVALCHUK2,3
1 University of Warsaw, Faculty of Geology, Żwirki i Wigury 93, 02-089 Warszawa, Poland;
e-mails: m.wasiluk@uw.edu.pl; m.granica2@uw.edu.pl
2 University of Wrocław, Faculty of Biological Sciences, Department of Palaeozoology,
Sienkiewicza 21, 50-335 Wrocław, Poland;
3 National Museum of Natural History of the National Academy of Sciences of Ukraine, Department
of Palaeontology, Bohdana Khmelnytskoho 15, 01054 Kyiv, Ukraine;
e-mail: biologiest@ukr.net
ABSTRACT:
Bienkowska-Wasiluk, M., Granica M. and Kovalchuk, O. 2024. A new extinct shad from Poland in the light
of clupeiform diversity and distribution within the Paratethys during the Oligocene. Acta Geologica Polonica,
74 (3), e23.
The Order Clupeiformes Bleeker, 1859 comprises herrings, anchovies, sprats, sardines, and shads. The fossil
record of this group is rich within the Paratethys. Here we describe a new clupeiform fish, †Sanalosa janulosa
gen. et sp. nov., from the Lower Oligocene of the Carpathian Basin, Poland. This new genus has a unique
combination of characters (lower jaw articulation located under the posterior part of the orbit; abdominal scutes
well developed with 3 to 5 in the gular region, 11–14 prepelvic scutes associated with ribs, 11–12 postpelvic
scutes; several striae on the frontals; an opercle with 6–12 thin radial ridges; a horizontal ramus of the preopercle
shorter than the vertical one; 42–44 vertebrae; 8–10 supraneurals; a dorsal fin with 18–22 rays, and an anal fin
with 21–23 rays), supporting recognition of a new genus and species within the Family Alosidae Svetovidov,
1952. Similarities and differences between fossil and extant genera of the Clupeiformes are discussed to shed
more light on their relationship. Moreover, the palaeobiogeography, diversity and distribution of Oligocene
clupeiform fishes in the Paratethys are presented and discussed.
Key words: Teleostei; Alosidae; New genus; Paratethys; Oligocene; Menilite Formation.
https://zoobank.org/References/db413df2-7645-4deb-a1ab-9b74378493fa
INTRODUCTION
The large and taxonomically diverse Order Clupei-
formes Bleeker, 1859 includes more than 400 extant
species of herrings, anchovies, sprats, sardines, shads,
and menhadens (Fricke et al. 2024). Representatives of
this group have a wide (mostly tropical) distribution,
and they are one of the most intensely commercially
exploited fishes worldwide (FAO 2022). Clupeiforms
are primarily marine, although some of them are
freshwater and anadromous (Nelson et al. 2 016). T hey
are medium-sized fishes, usually in the 150–250 mm
length range (e.g., Whitehead 1985). Most clupeiform
species form schools and swim near the surface, usu-
ally in coastal waters, feeding on plankton (Whitehead
1985; Nelson et al. 2016).
There is no consensus regarding the classification
of the Clupeiformes, although the classification of

2 MALGORZATA BIENKOWSKA-WASILUK ET AL.
Wang et al. (2022) used herein seems to fill a signif-
icant gap in taxonomic issues albeit some taxa of this
group are not defined by morphological characters.
Although the fossil skeletal record of the Clupei-
formes is rich (e.g., Daniltshenko 1960, 1980; Grande
1985; Murray et al. 2005; Baykina 2012, 2013a, b;
Marramà and Carnevale 2015a, b, 2018; Baykina and
Schwarzhans 2017a, b; Kovalchuk et al. 2020; Granica
et al. 2024), our knowledge on extinct representatives
is still insufficient, and their evolutionary history and
past diversity remain ambiguous and poorly under-
stood. The commonly used classification of clupei-
form fossils by Grande (1985) differs significantly
from that proposed by Wang et al. (2022), for exam-
ple in the understanding of the scope of the Family
Clupeidae Cuvier, 1817. It is a large and diverse fam-
ily in Grande (1985), but reduced to seven genera
in Wang et al. (2022), including only four genera of
the Subfamily Clupeinae Cuvier, 1817 sensu Grande
(1985) and three others representing the subfamilies
Alosinae Svetovidov, 1952 sensu Grande (1985) and
Pellonulinae Whitehead, 1985. Wang et al. (2022) rec-
ognised the Family Alosidae Svetovidov, 1952 with
four genera: Alosa Linck, 1790; Brevoortia G ill, 1861;
Sardina Antipa, 1904; and Sardinops Hubbs, 1929,
but without any subfamilies. The genera Alosa and
Brevoortia were classified by Grande (1985) to the
Alosinae, while the genera Sardina and Sardinops
– to the Clupeinae. Understanding the interrelation-
ships of fossil taxa in the light of current taxonomy is
challenging and needs a thorough revision.
The Oligocene deposits of the Menilite Formation
from the Outer Carpathians of Poland hold a unique
fossil fish record including extremely numerous re-
mains of clupeiforms usually represented by com-
plete or fragmented skeletons and isolated scales. In
this study, a new clupeiform fish is described from
the Oligocene deposits of Poland. The diversity and
palaeobiogeography of the Clupeiformes during the
Oligocene are discussed. Our investigation sheds
new light on the evolution, distribution, and diversity
of this group in the Paratethys.
LOCALITIES AND THEIR STRATIGRAPHIC
POSITION
The specimens were collected from five localities
situated in the Podkarpackie Voivodeship, southern
Poland, from the Oligocene deposits of the Silesian
and Skole units or nappes of the Outer Carpathians
(Text-fig. 1). Four localities (Dobra Góra, Hermanowa,
Jamna Dolna, and Średnia) expose the Skole Unit,
whereas a single locality (Jasienica Rosielna) lies
within the Silesian Unit.
The Dobra Góra locality (DG in Kotlarczyk et
Text-fig. 1. Location maps. A – Study area within Central Europe;
B – Study area within simplified geological map of the Outer
Carpathians (modified from Kováč et al. 1998); C – Localities
where the specimens were collected (black dots) within simplified
geological map of the Polish part of the Outer Carpathians (modi-
fied from Żytko et al. 1989).

A NEW EXTINCT SHAD FROM THE OLIGOCENE OF POLAND 3
al. 2006) lies 15 km north-east of Sanok city. The
Hermanowa locality (HE in Kotlarczyk et al. 2006;
Přikryl et al. 2016) is located 10 km south of Rzeszów
city. The Jamna Dolna locality (JD1 in Kotlarczyk et
al. 2006; Bienkowska-Wasiluk 2010) is 20 km south-
west of Przemyśl city. Średnia (SR in in Kotlarczyk
et al. 2006) is 15 km west of Przemyśl city. In these
four localities, the specimens were obtained from
the Rudawka Tractionite Member Unit, ichthyofaunal
zone IPM 2. The Jasienica Rosielna locality (Wasiluk
2013) is 30 km south of Rzeszów city. In this local-
ity the specimens come from the upper part of the
Menilite Formation, ichthyofaunal zone IMP2. The
ichthyofaunal zone IPM2 is correlated with the cal-
careous nannoplankton Biozone NP23 (Kotlarczyk
et al. 2006).
MATERIAL AND METHODS
The material used in this study is housed in the
Museum of the Faculty of Geology, University of
Warsaw (MWGUW, Muzeum Geologiczne im.
Stanisława Józefa Thugutta) and in the Department
of Palaeozoology, University of Wrocław, Wrocław
(ZPALWr.), Poland. It consists of six complete and al-
most complete articulated skeletons in the MWGUW
collection and one complete skeleton in the ZPALWr.
collection. The specimens were studied under a ste-
reomicroscope NIKON SMZ1000 at the Scanning
Electron and Optical Microscopy Laboratory at the
Faculty of Geology of the University of Warsaw.
All fishes were measured as standard length (SL),
which is the length of a specimen measured from the
anterior tip of the snout to the posterior margin of
the hypurals. The osteological terminology follows
Grande (1985), and Whitehead and Teugels (1985).
All extinct taxa are marked with a dagger (†) preced-
ing their name. Comparative information about the
Clupeiformes was mostly derived from Daniltshenko
(1960, 1980), Grande (1985), Whitehead (1985),
Murray et al. (2005), Baykina (2012, 2013a, b, 2015),
Marramà and Carnevale (2015a, b, 2018), Baykina
and Schwarzhans (2017a, b), Kovalchuk et al. (2020),
Kevrekidis et al. (2021), Fricke et al. (2024), Froese
and Pauly (2024), and Granica et al. (2024).
SYSTEMATIC PALAEONTOLOGY
Subdivision Teleostei Müller, 1846 sensu Arratia,
1999
Order Clupeiformes Bleeker, 1859 sensu Wang,
Dizaj, Huang, Sarker, Kevrekidis, Reichenbacher,
Esmaeili, Straube, Moritz and Li, 2022
Suborder Clupeoidei Bleeker, 1859
Family Alosidae Svetovidov, 1952
Genus †Sanalosa, gen. nov.
TYPE SPECIES: †Sanalosa janulosa sp. nov.
DIAGNOSIS: Lower jaw articulation located under
the posterior part of the orbit; abdominal scutes well
developed (including 3 to 5 scutes in the gular region,
11–14 prepelvic, associated with ribs, and 11–12 post-
pelvic scutes); several striae on the frontals; opercle
with 6–12 thin radial ridges; horizontal ramus of the
preopercle shorter than the vertical one; 42–44 verte-
brae; 8 to 10 supraneurals; dorsal fin with 18–22 rays,
and anal fin with 21–23 rays.
DERIVATION OF NAME: In reference to the San
River (close to which the fossils considered were
found) added to the Latin word Alosa meaning ‘shad’.
†Sanalosa janulosa gen. et sp. nov.
(Tex t-f igs 2 –9)
TYPE MATERIAL: The holotype, MGWUW
ZI/57/215/a–b, is a part and counterpart of a well pre-
served, ne arly complete articulate d skeleton. Paratypes
include: MGWUW ZI/57/214/a–b, ZI/57/171/1/a–b, as
part and counterpart, and ZI/57/182 in a single plate
(three specimens).
TYPE LOCALITY: Jamna Dolna near Bircza,
Podkarpackie Voivodeship (Subcarpathian Province)
of south-eastern Poland, Outer Carpathians, Poland.
TYPE HORIZON: Rudawka Tractionite Member of
the Menilite Formation, Lower Oligocene, Rupelian,
nannoplankton zone NP23.
Table 1. Specimens of †Sanalosa janulosa gen. et sp. nov. and
localities from which they were obtained.
Number of specimen Type of
material Locality
MWGUW ZI/57/215/a-b holotype Jamna Dolna
MWGUW ZI/57/182 paratype Dobra Gora
MWGUW ZI/57/171/1/a-b paratype Hermanowa
MWGUW ZI/57/214/a-b paratype Jasienica Rosielna
MWGUW ZI/57/133 material Jasienica Rosielna
MWGUW ZI/57/219 material Podkarpackie
Voivodeship
ZPALWr. N/6407 material Srednia

4 MALGORZATA BIENKOWSKA-WASILUK ET AL.
DERIVATION OF NAME: Named in honour of the
Polish poet Janusz Szuber (1947–2020) from Sanok
city, located close to the type locality, added to the
reduced word Alosa.
DIAGNOSIS: Same as for genus.
ADDITIONAL MATERIAL: MWGUW ZI/57/133,
ZI/57/219; ZPALWr. N/6407 (see Table 1).
Text-fig. 2. †Sanalosa janulosa gen. et sp. nov. from the Oligocene of the Outer Carpathians, SE Poland. A – holotype, MWGUW ZI/57/215/a;
B – paratype, MWGUW ZI/57/171/1/a; C – MWGUW ZI/57/219.

A NEW EXTINCT SHAD FROM THE OLIGOCENE OF POLAND 5
MEASUREMENTS: See Table 2.
DESCRIPTION: Small fishes with a moderately
high, elongated and laterally compressed body (Text-
fig. 2); the smallest specimen is 46 mm standard
length (SL) and the largest is 96 mm SL. The head
is triangular in lateral view, its length is 23–40%
SL. The mouth is small and terminal. The lower jaw
articulation is located under the posterior part of the
orbit (Text-fig. 3). The belly is moderately convex.
The abdominal scutes form a very distinctive keel.
Both pre- and postpelvic scutes are well-developed,
present from the coracoid to almost the beginning
of the anal fin and situated along the ventral margin.
Neurocranium. The neurocranium is elongated
and triangular in lateral outline. Paired frontals are
Text-fig. 3. †Sanalosa janulosa gen. et sp. nov., skull and its details. A, B – Skull, paratype, MWGUW ZI/57/171/1/a; photo and superim-
posed interpretative drawing. C, D – Frontal, paratype, MWGUW ZI/57/182; photo and superimposed interpretative drawing. E, F – Lower
jaw, paratype, MWGUW ZI/57/182, photo and superimposed interpretative drawing, anterior to the right. G, H – 1st infraorbital, paratype,
MWGUW ZI/57/182, photo and superimposed interpretative drawing, anterior to the right. I, J – Quadrate and symplectic, paratype, MWGUW
ZI/57/171/1/a, photo and superimposed interpretative drawing. K, L – Opercle, holotype, MWGUW ZI/57/214/a, photo and interpretative
drawing. Abbreviations: aa – anguloarticular; ao – antorbital; ch – ceratohyal; d – dentary; ect – ectopterygoid; f – frontal; hh – hypohyal; hym
– hyomandibular; io – infraorbital; iop – interopercle; enpt – endopterygoid; le – lateral ethmoid; me – mesethmoid; mtp – metapterygoid; mx
– maxilla; na – nasal; op – opercle; pas – parasphenoid; pl – palatine , pmx – premaxilla; pop – preopercle; q – quadrate; smx – supramaxilla;
so – supraorbital; sop – subopercle; sph – sphenotic; sym – symplectic.

6 MALGORZATA BIENKOWSKA-WASILUK ET AL.
the largest bones in the skull roof. They are pointed
anteriorly, wider posteriorly and narrow anteriorly;
the bones are curved with the descending anterior
part. In the posterior part, the frontals are sculptured
with several slightly curved frontoparietal striae
(Text-fig. 3C, D). Most of the frontals in the speci-
mens studied are slightly compressed dorsoventrally,
which allows us seeing both right and left bone simul-
taneously. The parasphenoid is long, thin, straight in
its central part, being slightly curved posterodorsally
and anteroventrally. The parietal, the supraoccipital
and the epioccipital are poorly visible posteriorly
to the posterior margin of frontals. The dorsal mar-
gin of the parietal articulates with the ventral mar-
gin of frontals. The pterotic is not preserved in the
studied material, and the sphenotic region is poorly
preserved. The orbitosphenoid is a moderately long,
descending anteriorly and extending from above the
central part of the orbit. The pterosphenoid is slightly
dorsally convex. The triangular lateral ethmoid artic-
ulates with the anterior part of the frontals.
Circumorbital series. The nasal is small and
moderately elongated. The supraorbital extends from
above the central part of the orbit. The infraorbitals
are poorly preserved; the first of them, ventral to the
anterior part of the orbit, appears to be the largest
bone of the series. The bone margins are poorly pre-
served, but they cover a large portion of the skull.
The sclerotic ring is poorly preserved, its posterior
part is not preserved but the anterior part appears to
have a crescent moon shape.
Oral jaws and dentition. The premaxilla is tooth-
less and slightly curved in lateral view. The maxilla
is narrow anteriorly and high posteriorly. Its ventral
margin is slightly convex and toothless. The ante-
rior, narrow part is moderately long and only slightly
curved. Two supramaxillae are present. The anterior
part of the asymmetrical second supramaxilla is nar-
row and straight, while its posterior part is robust.
Dorsal and ventral margins of the posterior part of
the bone are convex and rounded. The first supra-
maxilla appears to be a thin, straight bone. The hy-
pomaxilla is absent. The lower jaw is articulated with
the skull beneath the posterior part of the orbit, and
this articulation does not reach the vertical of the
posterior margin of the orbit. The anterior edge of the
lower jaw is moderately protruding. The lower jaw
(toothless dentary together with the anguloarticular)
is subtrapezoid (Text-fig. 3E, F), its ventral margin
is straight, and the dorsal margin is slightly convex.
The anteroventral edge of the jaw is rounded. The
anguloarticular has a moderately developed articular
process.
Suspensorium. The palatine seems to be long and
narrow. The ectopterygoid forms an obtuse angle
in its mid-length. The metapterygoid articulates an-
teriorly with the quadrate. The latter is triangular,
with a thick ventral margin; its articulation with the
lower jaw is located on the anteroventral corner. The
symplectic (Text-fig. 3I, J) is thin and gracile. The
hyomandibula is poorly preserved in the material
studied, it is almost parallel to the vertical posterior
margin of the orbit.
Opercular region. The preopercle is low, its
horizontal ramus is considerably shorter and wider
than the vertical one. The dorsal margin of the ver-
tical ramus reaches the middle of the orbit. The
angle between the preopercular rami is consider-
ably greater than 90º. The preopercle has a smooth
surface except for the canal-bearing ridges in the
central part of the bone between the rami. Margins
of the bone are rounded. The opercle is high and
moderately wide, sculptured with 6–12 thin radial
ridges (Text-fig. 3K, L). The ridges almost reach the
ventral and caudal margins of the bone. The anterior
margin is straight and slightly convex in the middle
part. Parallel to the margin, there is a thick slightly
convex ridge. The dorsal margin is convex, with
a descending posterodorsal corner. The posterior
margin is convex, with a slight incision in its upper
part. The ventral margin is straight anteriorly and
considerably rounded posteriorly. The subopercle
Table 2. Morphometric characteristics of †Sanalosa janulosa gen. et sp. nov. Measurements are given in millimetres (mm) and as a percentage
of the standard length, SL (in parentheses).
Morphometric character
MWGUW
ZI/57/215/a
holotype
MWGUW
ZI/57/171/1/b
paratype
MWGUW
ZI/57/182
paratype
MWGUW
ZI/57/214/a
paratype
MWGUW
ZI/57/133
MWGUW
ZI/57/219
ZPALWr.
N/6407 Studied material
Standard length [SL] 55 46 – 65 50 96 62 50–96
Head length 19 (35) 13 (28) 13 (–) 15 (23) 20 (40) 28 (29) 17 (27) 13–28 (23–40)
Maximum body depth 18 (33) 11 (24) 13 (–) 19 (29) 10 (20) 34 (35) 18 (30) 10–34 (20–35)
Predorsal distance 28 (51) 23 (50) 23 (–) 30 (46) 26 (52) 44 (46) 29 (48) 23–44 (46–52)
Prepelvic distance 31 (56) 26 (57) – 31 (48) – 49 (51) 30 (49) 26–49 (48–57)
Preanal distance 41(75) 34 (74) – 53 (82) – 74 (77) 48 (77) 34–74 (74–82)

A NEW EXTINCT SHAD FROM THE OLIGOCENE OF POLAND 7
articulates with the ventral margin of the opercle; its
ventral margin is rounded and convex. The suboper-
cle process is not visible. The interopercle appears
to be long and slightly curved.
Hyoid and branchial arches. There are 6–7 bran-
chiostegal rays (Text-fig. 4). Both anterior and pos-
terior rays are long, with thin and delicate anterior
ones and posterior ones being wider. The last ray is
higher than the others. The urohyal is feather-shaped,
with a narrow and long anterior part. The bone is
the highest in its central part. The ventral margin of
the anterior part of the urohyal is prolonged onto the
posterior part in a form of a ridge. The posterior part
is rounded with the posterodorsal corner ascending.
The dorsal margin appears to be slightly curved in
the anterior part and straight in the posterior part.
The hyoid bar and the margin between the dorsal and
ventral hypohyals are poorly preserved in the mate-
rial studied.
Vertebral column, ribs, and intermuscular bones.
The vertebral column consists of 42–44 vertebrae
including 17–18 caudal ones. Three anterior abdom-
inal vertebrae are covered with an opercle. The first
preural centrum is triangular in lateral view. In the
caudal region, neural spines are slightly curved and
positioned at approximately 45º to the vertebrae cen-
trum; haemal spines are positioned similarly. There
are 21–22 pairs of thin and long ribs reaching the
dorsal margin of abdominal scutes. At least two se-
ries of intermuscular bones are visible throughout the
abdominal part of the spine, including two ones in the
caudal region. Intermuscular bones are delicate and
curved, especially in the abdominal region; one se-
ries is short and close to the vertebrae centra, clearly
Text-fig. 4. †Sanalosa janulosa gen. et sp. nov., hyoid and branchial arches. A, B – Branchiostegal rays, paratype, MWGUW/57/182, photo
and superimposed interpretative drawing. C, D – Urohyal, paratype, MWGUW/57/171/1/a, photo and superimposed interpretative drawing.
E, F – Urohyal, paratype, MWGUW/57/182, photo and superimposed interpretative drawing. Abbreviations: br – branchiostegal rays; ch –
ceratohyal; hh – hypohyal; uh – urohyal.

8 MALGORZATA BIENKOWSKA-WASILUK ET AL.
visible in the abdominal region. The series near the
neural spines are closer to the vertebrae column than
the series near the haemal spines.
Eight to ten supraneurals are present, they are
curved and nail-shaped with the dorsal part wider and
narrowing right after the wide dorsal margin (Text-
fig. 5). They are positioned between the posterior
margin of the skull and beginning of the dorsal fin.
The latter is located slightly anteriorly to the middle of
the body, originating above the 14th to 18th vertebrae,
and terminating above the 23rd to 25th vertebrae.
Dorsal fin. The dorsal fin is triangular and con-
sists of 18–22 rays, the first anterior ray is the short-
est (Text-fig. 6). There are at least 17 pterygiophores.
The last pterygiophore is modified to a slender, hori-
zontally oriented stay, which has the length of at least
3 vertebrae.
Paired fins and girdles. The posttemporal is elon-
gate and subtriangular. The pectoral fins are long and
positioned just slightly above the abdominal outline.
They consist of 17–21 rays. The first rays are the lon-
gest. Two rod-like postcleithra are present. The su-
pracleithrum is curved posteriorly; margin between
the supracleithrum and cleithrum is below the verte-
bral column. The S-shaped cleithrum is the longest
bone in the pectoral girdle; it reaches the anterior
margin of the coracoid which is subquadrate in lat-
eral view (Text-fig. 7).
The pelvic fins are positioned beneath the middle
or anterior part of the dorsal fin with the length of
5–6 vertebrae. They originate below the 18th–19th
vertebrae. The pelvic bone is triangular in lateral
view with the length of 4–5 vertebrae, pointing an-
teriorly but poorly visible because of the abdominal
scutes. The pelvic fin consists of 8 rays.
Anal fin. The anal fin consists of 21–23 rays and
has 20–22 pterygiophores. It originates below the
27th–33rd vertebrae and terminates above the 38th–
42nd vertebrae. Rays closer to the caudal fin are dis-
placed. The first ray is shorter than the others. The
last two rays are not elongated.
Caudal fin and skeleton. The caudal fin is forked
and deeply notched. Six hypurals are present and two
epurals are visible (Text-fig. 8). The second hypural
Text-fig. 5. †Sanalosa janulosa gen. et sp. nov., supraneurals. A, B – Holotype, MWGUW/57/215/a, photo and superimposed interpretative
drawing.
Text-fig. 6. †Sanalosa janulosa gen. et sp. nov., dorsal fin. A, B – Holotype, MWGUW/57/215/a, photo and superimposed interpretative draw-
ing. Abbreviations: dfs – dorsal fin stay; fr – fin rays; p – pterygiophores.

A NEW EXTINCT SHAD FROM THE OLIGOCENE OF POLAND 9
seems to be fused with the first ural centrum. The
first preural vertebrae bears a short and thin neural
plate. The parhypural is long and higher in the ante-
rior part. The fin has 20 principal rays (I, 9 + 9, I) and
about 11 procurrent rays.
Squamation. Large cycloid scales with parallel
grooves (Text-fig. 9A–F).
Abdominal scutes. A continuous series originates
anterior to the pectoral fins and terminates anterior to
the anal fin. Scutes in the gular region, prepelvic and
postpelvic ones are well-developed (Text-fig. 9G, H).
The prepelvic scutes are larger. The ventral margin
of each scute is descending posteriorly and forms a
distinctive keel. There are at least 3 to 5 scutes along
the gular region (free prepelvic scutes), 11–14 prepel-
vic scutes associated with the ribs and at least 11–12
postpelvic ones. The uncertainty in the meristic data
regarding the abdominal scutes is because the pec-
toral fins might cover some of the scutes, and scutes
near the anal fin might have been displaced. The
pelvic scute is the largest as compared to the others.
DISCUSSION
Systematic discussion
The osteological and meristic data (Table 3) sup-
port the assignment of the examined specimens from
the Oligocene of the Polish Outer Carpathians to a
new genus and species of the Family Alosidae in the
Order Clupeiformes.
Text-fig. 8. †Sanalosa janulosa gen. et sp. nov., caudal skeleton. A, B – Details of the caudal skeleton, paratype, MWGUW ZI/57/214/a, photo
and superimposed interpretative drawing. Abbreviations: ep – epural; hyp – hypural; np – neural plate; phy – parhypural; pu – preural centrum;
un – uroneural.
Text-fig. 7. †Sanalosa janulosa gen. et sp. nov., pectoral girdle. A–D – Holotype, MWGUW/57/215/a, photo and superimposed interpretative
drawing. Abbreviations: cl – cleithrum; cor – coracoid; ptt – posttemporal, sc – scapula; scl – supracleithrum.

10 MALGORZATA BIENKOWSKA-WASILUK ET AL.
Table 3. Summary of selected morphological characters used to discriminate selected genera and species of the Order Clupeiformes. *Abdominal
scutes formula: [prepelvic scutes; pospelvic scutes] unassociated free scutes in the gular region; rib–associated prepelvic scutes scutes; scutes
between the coracoid and pelvic fin; scutes behind the pelvic fin. Comparative information is derived from Daniltshenko (1960, 1968, 1980),
Grande (1982, 1985), Whitehead (1985), Murray et al. (2005), Baykina (2012, 2013), Marramà and Carnevale (2015a, b, 2018), Baykina and
Schwarzhans (2017a, b), Kovalchuk et al. (2020), Kevrekidis et al. (2021), Fricke et al. (2024), Froese and Pauly (2024), and Granica et al. (2024).
Taxon
Frontoparietal striae
Opercle
Branchiostegal rays
Supraneurals
Dorsal scutes
Hypomaxilla
Dorsal-fin pterygiophores
or rays (rays)
Anal-fin pterygiophores
or rays (rays)
Pectoral-fin rays
Pelvic-fin rays
Vertebrae
Abdominal scutes *
†Sanalosa janulosa present striations 6–7 8–10 0 17
(18–22)
20–23
(21–23) 17–21 8 42–44 [15–17; 11–12]
3–5; 11–14; 11–12
Alosa (extant) striations 7–8 9–13 0–1 15–22 15–27 14–18 9–11 47–60
Alosa algeriensis striations (18–22) (20–25) 9 53–57 33–39 [19–23;
13–16]
Alosa fallax striations (16–22) (19–26) 15–17 9 49–59 32–41 [18–23;
12–18]
†Alosa genuina striations (15–17) (17–18) 14–15 9 39–40 [12; 8]
†Alosa sculptata striations (14) (18–19) 17 7–8 44 [15; 14]
†Alosa cf. sagorensis striations (15) (20) 15–16 8 39–41 22–24
†Beksinskiella 4+ smooth 6–7 8–10 0 absent 19–20 17–22 18–21 8–10 44–48 0; 12–14; 8+
†Bolcaichthys 10–14 smooth 5–6 8 0 absent 15–16 15–16 14–18 8 40–42 0; 10–11; 10–11
Brevoortia striations 7 10–12 17–24 18–24 7 45–48 about 30–32
†Chasmoclupea smooth 13 0 absent 12 7 40+ 4; 17; 5+
Clupea smooth 8 15–19 0 absent 17–18 15–18 8–10 52–57
Clupeoides smooth 2+ ? 0 absent 11–17 15–26 7 7–12; 6–10
Clupeonella smooth 7 11 0 absent 15 18–21 8 42 23–32
Dussumieria 12–17 21–22 19–22 14–18 12–15 8 55–56 0; 0; 0
†Eoalosa smooth 13+ 0 15 17 7 47 0; 12; 5
†Gosiutichthys smooth 7–8 6–7 12–13 absent 10–11 10–13 6–7 34–36 20–22
†Karaganops present smooth 7 10 0 absent 18–19 17–18 15 8–9 44–46 0; 13–15; 10
†Knightia present smooth 7–8 7–8 12–14 absent 11–14 13–17 11–14 7 37–39 about 21–23
†Maicopiella absent smooth 7 8–10 0 absent 19 17–18 17 8–9 42–45 0; 14–15; 10–11
†Moldavichthys present smooth 7–8 9–10 0 16–17 17–18 ? 8 39–44 [15–16; 8]
Opisthonema smooth 6 7–9 1 absent 18–19 18–22 ? 8 45–47
†Paretrumeus smooth (15–17) (7–8) 20–23 26–27 50–55 0; 0; 0
†‘Pomolobus’ striations 0 14–17 17–22 14–18 8–9 40–43 [10–13; 9–12]
†‘Pomolobus’ curtus striations (14–15) (19–20) 14–15 9 40–41 [11–12; 9–10]
†‘Pomolobus’ facilis striations (16–17) (20–22) 17–18 8–9 42–43 [10–11; 12]
†‘Pomolobus’ antiquus striations (14–15) (17–18) 17 9 42 [12–13; 9–10]
†Primisardinella smooth 9–10 0 absent 15–16 13–15 8 39–40 3–4; 10–11; 9–10
†Pseudohilsa present smooth 5 10–11 0 absent (10–17) (16–19) 15 8–9 36–42 4; 11–12; 10–11
†Rupelia present smooth 7 9 0 absent 20 16–18 19–20 9 48–50 0; 15; 10–11
Sardina striations 7 10–11 0 absent 17–18 17–19 8 50–51
†Sardina
necteodosciobanensis striations (16–18) (20) 18–19 9 46–47 [12–13; 12–13]
†Sardina tarletskovi striations 11 15–16
(14–15) 14 47–49 [17–18; 13]
Sardinella 7–14 smooth 5–7 8–10 0 absent 16–19 16–20 13–18 8–9 43–48 0; 15–20; 11–16
Sardinops striations 7–8 10 absent 18–19 17–18 8 50–52
†Sarmatella smooth 7 10–12 0 absent 15–20 13–17 16–17 8–9 44–54 0; 22–24; 10–12
Sprattus smooth 7 15–17 0 absent 17–18 16–19 7–8 45–48
†Trollichthys smooth 5–6 14–16 13 8 41–42 0; 0; 0

A NEW EXTINCT SHAD FROM THE OLIGOCENE OF POLAND 11
The presence of abdominal scutes (keeled scales
along the ventral midline) indicates that †Sanalosa
janulosa gen. et sp. nov. belongs to the Clupeiformes
(see Wang et al. 2022). The fusion of the second hy-
pural with the first ural centrum, a separated first
hypural, the fusion of the first uroneural with the first
Text-fig. 9. †Sanalosa janulosa gen. et sp. nov., scales and abdominal scutes. A, B – Details of a scale from the anterior median region of
the body, MWGUW/57/219, photo and superimposed interpretative drawing. C, D – Details of a scale from the anterior dorsal region of
the body, MWGUW/57/219, photo and superimposed interpretative drawing. E, F – Details of a scale from the median region of the body,
MWGUW/57/219, photo and superimposed interpretative drawing. G, H – Details of abdominal scutes, holotype, MWGUW/57/215/a, photo
and superimposed interpretative drawing. Abbreviations: fs – un-associated free scutes in the gular region; ps – pelvic scute; pps – postpelvic
scutes, scutes behind the pelvic fin; rs – rib-associated prepelvic scutes.

12 MALGORZATA BIENKOWSKA-WASILUK ET AL.
preural centrum, the size reduction of the first ural
centrum and the separation of the parhypural from the
first ural centrum indicate that the species belongs to
the Suborder Clupeoidei (see Grande 1985). The oper-
cle sculptured with radial ridges supports its inclusion
within the Family Alosidae (see Wang et al. 2022).
Comparison with extant genera
†Sanalosa gen. nov. differs from extant genera
of the Clupeiformes except of Alosa, Brevoortia,
Sardina, and Sardinops (see Grande 1985; Whitehead
1985; Wang et al. 2022) by the opercle sculptured
with radial ridges. It differs from Alosa, Brevoortia,
Sardina, and Sardinops (Grande 1985; Whitehead
1985; Froese and Pauly 2024) by having a lesser num-
ber of vertebrae (42–44 vs. 47–60, 45–48, 50–51 and
50–52, respectively). It differs in the number of rays
of the pelvic fin from Alosa and Brevoortia (8 vs.
9–11 and 7, respectively). †Sanalosa gen. nov. can
be differentiated from the genus Alosa by the posi-
tion of the lower jaw articulation with the skull – it
does not reach the vertical of the posterior margin
of the orbit whereas the lower jaw articulation in
Alosa is behind this vertical axis. †Sanalosa gen. nov.
has a smaller number of supraneurals compared to
Brevoortia, with the highest number the same as the
smallest number of supraneurals in Brevoortia (8–10
vs. 10–12). †Sanalosa gen. nov. differs from Sardina
in a higher number of pterygiophores of the anal fin
(20–23 vs. 17–19).
The urohyal of Sardinops (see Sato et al. 1988)
has a longer and higher anterior part of the bone than
that in †Sanalosa gen. nov. The respective bone of
Sardinops has a curved dorsal margin and a more
paddle-like outline. The urohyal of †Sanalosa gen.
nov. has a straighter dorsal margin and more feather-
like outline.
Comparison with extinct genera and species
†Sanalosa gen. nov. differs from extinct genera of
the Clupeiformes (e.g., †Beksinskiella Granica, Bien-
kowska-Wasiluk and Pałdyna, 2024; †Bolcaichthys
Marramà and Carnevale, 2015a, †Chasmoclupea
Mur ray, Simons and Attia, 2005, †Eoalosa Marramà
and Carnevale, 2018, †Gosiutichthys Grande, 1982,
†Karaganops Baykina and Schwarzhans, 2017a,
†Knightia Jo rdan, 1907, †Maicopiella (Menner, 1949),
†Paretrumeus Daniltshenko, 1980, †Primisardinella
Daniltshenko, 1968, †Pseudohilsa Menner, 1949,
†Rupelia Baykina and Kovalchuk in Kovalchuk et al.
2020, †Sarmatella Menner, 1949, and †Trollichthys
Marr amà and Car ne vale, 2015b except of †‘Pomolobus’
Rafinesque, 1820 and †Moldavichthys Baykina and
Schwarzhans, 2017b by the opercle sculptured with
radial ridges (see Table 3). †Sanalosa gen. nov. dif-
fers from the Eocene †Trollichthys Marramà and
Carnevale, 2015b, and †Paretrumeus Daniltshenko,
1980 in the presence of abdominal scutes.
The Miocene species †Moldavichthys switshens-
kae (Baykina and Schwarzhans, 2017b) differs from
†Sanalosa janulosa gen. et sp. nov. by fewer rays in
the dorsal fin (15–16 vs. 18–22), pectoral fins (14–16
vs. 17–21) and the anal fin (17–18 vs. 21–23), as well
as the smaller number of postpelvic abdominal scutes
(8 vs. 11–12). The new genus and species has a dif-
ferent morphology of the preopercle. The horizontal
ramus is slightly shorter, and the vertical ramus is
narrower. Another difference is the presence of the
teeth: the maxilla of †M. switshenskae is serrated and
there are teeth on the premaxilla and dentary. The
Oligocene †‘Pomolobus’ curtus Daniltshenko, 1960
differs from †S. janulosa gen. et sp. nov. by fewer
rays in the dorsal fin (14–15) and the pectoral fins
(14–15). †Sanalosa janulosa gen. et sp. nov. can be
differentiated from the Oligocene †‘Pomolobus’ fac-
ilis Daniltshenko, 1960 and †‘P.’ antiquus (Sm i rnov,
1936) by having more numerous rays in the dorsal
fin (18–22 vs. 16–17; 14–15, see Daniltshenko 1980).
Those three species also have fewer prepelvic ab-
dominal scutes (11–12; 10–11; 12–13 vs. 15–17 in-
cluding 3 to 5 in the gular region).
The Oligocene †Beksinskiella longimana (see
Granica et al. 2024) differs from †S. janulosa gen.
et sp. nov. by its different preopercle morphology
(both rami are similar in length vs. the horizontal
ramus is shorter than the vertical one). The urohyal
of †B. longimana becomes higher in the central part
of the bone at one point, while the height of this
bone in †Sanalosa gen. nov. changes gradually. The
posterior margin of the urohyal is fully rounded in
†B. longimana. The posterior part of the bone in †B.
longimana retains a similar height, while the height
in the urohyal of †Sanalosa gen. nov. in the poste-
rior part becomes smaller towards its dorsal margin.
†Beksinskiella longimana has poorly developed post-
pelvic scutes, and their number is smaller (about 8),
while †S. janulosa has more numerous (11–12) and
well-developed ones.
The Oligocene †Sardina necteodosciobanensis
Ciobanu, 1977 differs from †S. janulosa gen. et sp.
nov. by its having fewer rays in the dorsal fin (16–
18), more rays in pelvic fins (9) and more vertebrae
(46–47). The Miocene †Sardina tarletskovi Baykina,
2015 differs from †S. janulosa gen. et sp. nov. by its

A NEW EXTINCT SHAD FROM THE OLIGOCENE OF POLAND 13
having fewer rays in the anal fin (15) and in the dorsal
fin (14–15). †Sardina tarletskovi has more numerous
vertebrae (47–49). It also has a smaller head (22%
SL). †Sanalosa janulosa gen. et sp. nov. has more
dorsal fin rays (18–22) than the Oligocene–Miocene
†Alosa genuina Daniltshenko, 1960 (15–17), †Alosa
sculptata Weiler, 1928 (14) and †Alosa cf. sagoren-
sis Steindachner, 1863 (15) (see Weiler 1933, 1938;
Ciobanu 1977; Daniltshenko 1980). The sculpture on
the opercle in †S. janulosa covers a smaller area than
that in †A. cf. sagorensis (see Weiler 1933).
Distribution, diversity and palaeobiogeography of
the Oligocene Clupeiformes
The earliest clupeiform fish in the fossil record,
preserved as skeleton, has been reported from the
Lower Cretaceous of Brazil (De Figueiredo 2009),
dated to approximately 126–121 Ma (Barremian, cal-
ibrated Geological Time Scale after Gradstein et al.
2020).
During the Oligocene, representatives of this or-
der were abundant in marine ecosystems, which is
indicated by their rich fossil record and high percent-
age in fossil assemblages (Kotlarczyk et al. 2006;
Bienkowska-Wasiluk 2010; Přikryl et al. 2 016 ).
More than twenty clupeiform taxa, preserved as
skeletons, have been repor ted from the Czech Republic,
Egypt, Hungary, Poland, Romania, Russia, and Ukra-
ine (Weiler 1933; Ciobanu 1977; Daniltshenko 1980;
Murray et al. 2005; Kovalchuk et al. 2020; Granica
et al. 2024), although recent investigations (e.g.,
Kovalchuk et al. 2020) show that revisions of some
previously described taxa led to a reduced number of
species.
†Alosa sculptata has been recorded from Romania
(Ciobanu 1977) and Germany (Weiler 1928), although
some morphological characters of the species have
not been described (e.g., the number of supraneurals,
epurals and branchiostegal rays), therefore the valid-
ity of this species needs to be reconsidered.
†Alosa cf. sagorensis has been identified from
Hungary (Weiler 1933, 1938) and Poland (Szymczyk
1978), but the Polish specimens are fragmentary and
a complete skeleton has not been found yet. Similarly
to †A. sculptata, the validity of this species needs
to be reconsidered due to the absence of important
morphological characters in the original descrip-
tion. †Alosa sagorensis has been described from the
Miocene of Croatia (Steindachner 1863).
†Beksinskiella longimana (Heckel, 1850) has
been recorded from the Czech Republic, Poland, and
Ukraine (Granica et al. 2024)
†Chasmoclupea aegyptica Murray, Simons and
Attia, 2005 has been described from the freshwater
deposits of Egypt (Murray et al. 2005).
†Moldavichthys switshenskae from the Miocene
of Moldova (Baykina and Schwarzhans, 2017b) is
one of the earliest species of the Alosidae that has a
well-documented morphology. However, more than
6 taxa of putative Alosidae occurred earlier than
Moldavichthys in the Oligocene.
†‘Pomolobus’ antiquus, †‘P.’ curtus, and †‘P.’ fac-
ilis have been described from Russia (Daniltshenko
1960). Pomolobus is currently regarded as a synonym
of Alosa (Fricke et al. 2024). These three species
share some characters with Alosa but differ in having
fewer vertebrae and abdominal scutes. Their revision
would be desirable to clarify their taxonomic status.
†Rupelia rata (Daniltshenko, 1959) is known from
Russia (Kovalchuk et al. 2020), as well as the putative
Dussumieriidae, †Paretrumeus avitus Daniltshenko,
1980.
†Sardina necteodosciobanensis has been de-
scribed from Romania (Ciobanu 1977).
All the above species with the exception of †Ch.
aegyptica inhabited marine environments.
The Miocene fossil record of the Clupeiformes
within the Paratethys is represented mostly by spe-
cies which are different from the Oligocene ones,
with only one or two species (†Alosa cf. sagoren-
sis, †Beksinskiella longimana) found to be present in
both series. This could be a result of significant en-
vironmental changes in the Paratethys including sea-
level rise and drop, periodical isolation of its sub-ba-
sins and tectonic events (e.g., Kotlarczyk et al. 2006;
Kováč et al. 2016, 2017; Sachsenhofer et al. 2017).
Clupeiform fossils have been reported from the
Oligocene of Western, Central and Eastern Para-
tethys (e.g., Kotlarczyk et al. 2006; Maxwell et al.
2016; Kovalchuk et al. 2020). In the Western Para-
te thys, in the Upper Rhine Graben lived †Alosa
sculptata (see Weiler 1928). This species was present
also in the Central Paratethys in the Carpathian Basin
together with †Alosa cf. sagorensis, †Beksinskiella
longimana, †Sanalosa janulosa gen. et sp. nov., and
†Sardina necteodosciobanensis. †Alosa cf. sagoren-
sis lived also in the Hungarian Basin of the Central
Paratethys (see Weiler 1933). The clupeiform assem-
blage of Eastern Paratethys included †Paretrumeus
avitus, †‘Pomolobus’ antiquus, †‘P.’ curtus, †‘P.’
facilis, and †Rupelia rata (see Daniltshenko 1980;
Kovalchuk et al. 2020), all of which were absent in the
Central and Western Paratethys. †Chasmoclupea ae-
gyptica inhabited the rivers of North Africa (Murray
et al. 2005).

14 MALGORZATA BIENKOWSKA-WASILUK ET AL.
Although a number of fish taxa lived both in the
Central and Eastern Paratethys during Oligocene
(e.g., Bannikov 2010; Barkaszi and Kovalchuk 2021;
Kovalchuk and Barkaszi 2021; Přikryl et al . 2022), the
composition of the clupeiform assemblages differed
considerably in these parts of the Paratethyan realm
(Text-fig. 10). All Eocene genera from the Tethys,
i.e., †Bolcaichthys, †Eoalosa, and †Trollichthys (see
Marramà and Carnevale 2015a, b, 2018) were ab-
sent in the Paratethys during the Oligocene. No clu-
peiform genus has been recorded in the Peri-Tethys
during the Eocene (see Daniltshenko 1980). Only
Alosa and Sardina were present in the Paratethys
during the Oligocene and Miocene. †Alosa sculptata,
†A. cf. sagorensis, and †Sardina necteodosciobanen-
sis lived during the Oligocene (see Weiler 1928, 1933;
Ciobanu 1977), whereas †A. sagorensis, †A. genuina,
and †Sardina tarletskovi lived during the Miocene
(see Steindachner 1863; Daniltshenko 1980; Baykina
2015).
Other genera living in the Paratethys during
the Oligocene, i.e., †Beksinskiella, †Paretrumeus,
†‘P o m o l o b u s ’, †Rupelia, †Sanalosa gen. nov., and
S a r d i n a have not been recorded in the Miocene
of the Paratethys. †Karaganops, †Maicopiella,
†Moldavichthys, †Pseudohilsa, and †Sarmatella
originated in the Paratethys during the Miocene.
Although the clupeiform evolutionary history re-
mains to be explored further, it is clear that they
evolved rapidly during Eocene, Oligocene, and
Miocene, and endemism prevailed in the Paratethys.
Differences between the species of clupeiforms are
expressed in meristic and osteological characters, but
the taxa have a high number of shared characters.
This concerns both recent and Paleogene–Neogene
Clupeiformes in the northern part of the Tethys.
Therefore comparison of taxa and recognition of evo-
lutionary trends needs further comprehensive analy-
ses. We believe that future investigations of clupei-
form fossils from the former Tethys will improve our
knowledge on the evolutionary history of this group.
CONCLUSIONS
Osteological, morphometric and meristic analy-
ses of the clupeiform material from the Polish Outer
Carpathians revealed a new genus and species,
†Sanalosa janulosa. The description of †S. janulosa
gen. et sp. nov. provides a substantial improvement
to our knowledge of osteology of the Oligocene alo-
sids, documenting essential features such as abdom-
inal scutes, supraneurals, and urohyal. The newly
described species existed in the Central Paratethys
together with †Alosa sculptata, †Alosa cf. sagoren-
sis, †Beksinskiella longimana, and †Sardina nec-
Text-fig. 10. Paleobiogeography of representatives of the Order Clupeiformes in the Oligocene based on skeleton findings; palaeogeography
adopted from Popov et al. (2002).

A NEW EXTINCT SHAD FROM THE OLIGOCENE OF POLAND 15
teodosciobanensis. Clupeiform assemblages were
highly diverse in the basins of the Paratethyan realm,
showing a rapid, often endemic evolution. We believe
that our investigation will improve the knowledge
on the evolutionary history of clupeiform fishes and
can contribute to improving the palaeobiogeographic
reconstructions of the Paratethys.
Acknowledgements
We express our gratitude to Marcin Pałdyna for discus-
sions and collecting some specimens, Robert Szybiak for access
to his specimens, Radosław Wasiluk for helping with field-
work, Tomasz Praszkier and Krzysztof Dembicz for collecting
some specimens. We acknowledge Bettina Reichenbacher and
Werner Schwarzhans for their valuable comments and reviews
that significantly improved the quality of the manuscript. The
research of OK was supported by an individual grant ‘The role
of particular fish groups in the functioning of late Mesozoic and
Cenozoic ecosystems of Eastern Europe’ (No. 0123U102984)
from the National Academy of Sciences of Ukraine.
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Manuscript submitted: 13th May 2024
Revised version accepted: 16th September 2024