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Acta Geologica Polonica, Vol. XX (202X), No. X, pp. xxx–xxx
DOI: 10.24425/agp.2020.134561
Marine fishes (Elasmobranchii, Teleostei)
from the Glendon Limestone Member
of the Byram Formation (Oligocene, Rupelian)
at site AWa-9, Washington County, Alabama, USA,
including a new species of gobiid (Gobiiformes: Gobiidae)
JUN A. EBERSOLE1, DAVID J. CICIMURRI2 and GARY L. STRINGER3
1 McWane Science Center, 200 19th Street North, Birmingham, AL 35203, USA.
E-mail: jebersole@mcwane.org
2 South Carolina State Museum, 301 Gervais Street, Columbia, SC 29201, USA.
E-mail: dave.cicimurri@scmuseum.org
3 Museum of Natural History, 708 University Avenue, University of Louisiana at Monroe, Monroe, LA 71209, USA.
E-mail: stringer@ulm.edu
ABSTRACT:
Ebersole, J.A., Cicimurri, D.J. and Stringer, G.L. XXXX. Marine fishes (Elasmobranchii, Teleostei) from the
Glendon Limestone Member of the Byram Formation (Oligocene, Rupelian) at site AWa-9, Washington County,
Alabama, USA, including a new species of gobiid (Gobiiformes: Gobiidae). Acta Geologica Polonica, XX (X),
xxx−xxx.
The Oligocene (Rupelian) Byram Formation (Vicksburg Group) in Alabama, USA, is divided into three
members, including (in ascending order) the Glendon Limestone, unnamed marl, and the Bucatunna Clay.
The Oligocene marine units in Alabama have been historically under-investigated, but bulk samples re-
cently obtained from Glendon Limestone Member exposures at site AWa-9 in Washington County yielded
20 unequivocal elasmobranch and teleost taxa. This surprisingly diverse paleofauna, based on isolated teeth,
bones and otoliths, includes the new taxon, Gobiosoma? axsmithi sp. nov., as well as “Aetomylaeus” sp.,
Albula sp., Aplodinotus gemma Koken, 1888, Ariosoma nonsector Nolf and Stringer, 2003, Balistidae indet.,
Citharichthys sp., Myliobatoidei indet., Diretmus? sp., Hemipristis sp., Negaprion aff. N. gilmorei (Leriche,
1942), Pachyscyllium sp., Paralbula sp., Physogaleus sp., Preophidion meyeri (Koken, 1888), Sciaena pseu-
doradians (Dante and Frizzell in Frizzell and Dante, 1965), Sciaenops? sp., Sparus? elegantulus Koken, 1888,
Sphyraena sp., and Syacium sp. Additional remains were recovered but could not be identified beyond unde-
termined Elasmobranchii or Teleostei. All these taxa represent first occurrences within the Glendon Limestone
Member in Alabama, and the “Aetomylaeus” sp., Pachyscyllium sp., Paralbula sp., and Sciaenops? sp. spec-
imens represent the first occurrences of each in the Oligocene of the Gulf Coastal Plain of the USA. We also
report the first record of Oligocene Paralbula Blake, 1940 teeth, and the first occurrence of an Oligocene
member of the Balistidae in the Western Hemisphere. This marine vertebrate assemblage indicates that the
Glendon Limestone Member at site AWa-9 represented a subtropical to temperate, middle shelf paleoenviron-
ment with a paleowater depth interpreted as 30–100 m.
Key words: Chondrichthyes; Teleostei; Otoliths; Paleogene; Gulf Coastal Plain; Vicksburg
Group.
2 JUN A. EBERSOLE ET AL.
INTRODUCTION
Surface exposures in southwest Alabama, USA,
preserve a nearly complete Oligocene marine se-
quence that includes, in ascending order, the Rupelian
Bumpnose Limestone, Red Bluff Clay, Forest Hill
Sand, Marianna Limestone, and Byram Formation
(including Glendon Limestone, unnamed marl, and
Bucatunna Clay members), and the Chattian Chicka-
sawhay Limestone and Paynes Hammock Sand (Szabo
et al. 1988). The Bumpnose Limestone, Red Bluff Clay,
Forest Hill Sand, Marianna Limestone, and Byram
Formation are all part of the Vicksburg Group, whereas
the Chickasawhay Limestone and Paynes Hammock
Sand are not assigned to a geologic group (Raymond et
al. 1988). These Oligocene formations have long been
known for their abundance of marine invertebrates
(see Cooke 1918; Howe 1942; Glawe 1967, 1969), but
very little research has been conducted in Alabama on
the vertebrates from these units.
The first scientific study of Oligocene vertebrate
remains from the state was by Koken (1888), who
described 23 otolith-based taxa from Eocene and
Oligocene strata in both Alabama and Mississippi,
USA. Cooke (1926) noted the occurrence of fossil
otoliths in the Byram Formation of Alabama, but
none of these were described or figured. Campbell
(1929a) figured many of Koken’s (1888) Alabama
taxa in a Journal of Paleontology article, and later
that year he republished the same specimens in his
Bibliography of Otoliths (Campbell 1929b).
Frizzell and Lamber (1962) described several
congrid otoliths from the Oligocene Red Bluff Clay
in Alabama. The specimens they reported were col-
lected from the Lone Star Cement Company Quarry
at St. Stephens in Washington County, Alabama, the
same locality (albeit a different unit) from which the
material forming the basis of the current study was
obtained. As part of their work on Cenozoic fishes of
the Gulf Coast, Frizzell and Dante (1965) listed the
St. Stephens quarry, and specifically the Red Bluff
Clay, as one of the localities from which their sam-
ple of otoliths was derived. A few years later, Salem
(1971), a graduate student of Frizzell, studied the Red
Bluff Clay otoliths from the St. Stephens quarry, and
several of these specimens were later reported by
Nolf (1985, 2013).
Whetstone and Martin (1978) described a fossil
sirenian from the Bucatunna Clay Member of the
Byram Formation at the St. Stephens quarry. This
specimen represented the first non-otolith vertebrate
to be described from any Oligocene unit in Alabama
and was the first vertebrate to be described from
any Oligocene unit other than the Red Bluff Clay.
Whetstone and Martin (1978) also provided a list
of vertebrates that were associated with the sire-
nian, which included crocodilian osteoderms, spines
and dental plates of myliobatid rays, teeth from the
shark genera Odontaspis, Galeocerdo, and Isurus,
and vertebrae and otoliths of teleost fishes. These
specimens were reported to be in the collections of
the Geological Survey of Alabama in Tuscaloosa,
USA, but they now reside in the collections of the
Alabama Museum of Natural History in Tuscaloosa.
Thurmond and Jones (1981, p. 6) discussed the lack
of studies on Oligocene vertebrates in Alabama but
mentioned that the “Oligocene Red Bluff Formation
has produced at least one specimen.” It is unclear,
however, if the specimen in question was one of the
otolith taxa described earlier by Frizzell and Lamber
(1962), Frizzell and Dante (1965), or Salem (1971), or
a different taxon altogether. The fossil fishes of the
Bucatunna Clay are in need of further investigation.
A more recent note of chondrichthyan taxa from
the Oligocene Vicksburg Group in Alabama was by
Manning (2003), who made mention of (but did not
describe) several species. Later, Ehret and Ebersole
(2014) noted the presence of carcharhinid, gingly-
mostomatid, and myliobatid teeth from Oligocene de-
posits in Alabama, but these specimens were also not
figured or described. Lastly, Stringer et al. (2020a)
reported the first occurrence of the enigmatic otolith
taxon, Equetulus silverdalensis (Müller, 1999), in the
state. This specimen was collected from the Chattian
Paynes Hammock Sand in Washington County,
Alabama.
To date, no vertebrate specimens have been de-
scribed f rom the Rupelian Glendon Limestone Member
of the Byram Formation in Alabama. Verte brate fos-
sils have, however, been described from this litho-
stratigraphic unit in neighboring Mississippi, includ-
ing fossil otoliths that were reported by Salem (1971),
Daly (1992), Phillips and Stringer (2007), and Stringer
and Starnes (2020). Daly (1992, p. 10) also listed
‘shark’ and ‘fish’ among the vertebrate taxa occurring
in the Glendon Limestone, and DeVries (1963, p. 40)
noted that “a few fish vertebrae and shark teeth can
be found in the Glendon Formation in Jasper County.”
Two years later, Moore (1965, p. 72) made mention of a
“shark-tooth locality in the Lower Oligocene Glendon
Formation” located south of Jackson, Mississippi.
Dockery and Thompson (2016) reported and fig-
ured an articulated skeleton of the fossil squirrelfish,
Holocentrites ovalis Conrad, 1941, that was collected
from the Glendon Limestone in Rankin County, and
this taxon has subsequently been confirmed from
FISHES FROM THE OLIGOCENE OF ALABAMA 3
the same unit in Smith County (James Starnes, pers.
comm.). Although Dockery and Manning (1986) re-
ported the occurrence of Otodus (Carcharocles) au-
riculatus (de Blainville, 1818) from the Vicksburg
Group and Byram Formation in Mississippi, this taxon
was not confirmed specifically from the Glendon
Limestone. Additionally, Manning (1997) reported
terrestrial mammals from the Byram Formation in
Mississippi. It is important to note that in Mississippi
the Byram Formation is considered a distinct lithologic
unit that overlies the Glendon Limestone, whereas in
Alabama the Glendon Limestone represents the lowest
member of the Byram Formation. Furthermore, the
Byram Formation in Mississippi is stratigraphically
equivalent to the unnamed marl member of the Byram
Formation in Alabama (Raymond et al. 1988; Dockery
and Thompson 2016).
Herein we provide the first account of marine
vertebrate taxa occurring in the Oligocene Glendon
Limestone Member of the Byram Formation in
Alabama, USA. We provide detailed descriptions
and figures of the various remains, discuss the tax-
onomic issues and paleobiogeographic significance
of the taxa, and comment on the paleoenvironment in
Alabama during this interval in the Rupelian.
GEOLOGICAL SETTING
The vertebrate specimens described herein were
derived from bulk samples collected from the top
of the Glendon Limestone Member of the Byram
Formation exposed along a hillside at St. Stephens
Historical Park in Washington County, Alabama,
USA (locality designation AWa-9, Text-fig. 1). This
historical locality, once a prominent landmark known
as St. Stephens Bluff, was purchased in the early 20th
century by the Lone Star Cement Company (LSCC),
which started quarrying operations at the site in 1928.
Over its many years of operation, LSCC quarrying
activity at the locality revealed the most complete
and continuously exposed Oligocene marine section
in Alabama, down to the Eocene/Oligocene contact
(Glawe 1967). Today this locality resides within the
boundaries of St. Stephens Historical Park.
In Alabama, the Byram Formation represents the
uppermost lithostratigraphic unit of the Oligocene
(Rupelian) Vicksburg Group. The formation is di-
vided into three members, including the Glendon
Limestone Member at the base, an unnamed marl
member, and the Bucatunna Clay Member at the top
(Raymond et al. 1988). The Byram Formation con-
formably overlies the Marianna Limestone, also of
the Vicksburg Group, and is conformably overlain
by the Chickasawhay Limestone. All of these lith-
ologic units are exposed in sequence at site AWa-9
(Glawe 1967, 1969). Hopkins (1917) credited the
name Glendon Limestone to Charles Wythe Cooke,
who described the unit in an earlier unpublished
manuscript. Hopkins (1917) originally designated
the Glendon Limestone Member as the top member
of the Marianna Limestone, but Cooke (1923) later
elevated the unit to formation rank. MacNeil (1944)
and Monroe (1954) disagreed with the latter assess-
ment and recognized the Glendon Limestone as the
basal member of the Byram Formation. According
to Dockery and Thompson (2016), this attribution
was logical because both the Glendon Limestone and
Byram Formation occur within the Lepidocyclina su-
pera large-foraminifera zone of Gravell and Hanna
(1938) and the Pecten perplanus byramensis pecten
zone of Glawe (1969). Following MacNeil (1944) and
Monroe (1954), the Geological Survey of Alabama
currently recognizes the Glendon Limestone as the
lower member of the Byram Formation (Raymond
et al. 1988), and this interpretation is followed here.
In Mississippi, however, the Mississippi Office of
Geology has chosen to follow Cooke (1923) in recog-
nizing the Glendon Limestone as a distinct lithologi-
cal formation (Dockery and Thompson 2016).
In Alabama, exposures of the Glendon Limestone
Member crop out in Choctaw, Clarke, and Washing ton
counties in the southwestern part of the state (Szabo
et al. 1988). The unit extends into eastern Mississippi
Text-fig. 1. Approximate location of site AWa-9 and Oligocene sur-
face exposures in Washington County, Alabama, USA. Modified
from Stringer et al. (2020a).
4 JUN A. EBERSOLE ET AL.
where it thickens and becomes more detrital, with
surface exposures occurring in at least seven coun-
ties (Dockery and Thompson 2016). Although the
Glendon Limestone has also been reported in Georgia
and northern Florida (Cooke 1923), further investiga-
tion by Huddlestun (1993) showed that this unit is ex-
posed at only a single locality in central Georgia, and
the reported occurrences in Florida instead represent
other time-equivalent units.
The type section for the Glendon Limestone
Member is located at Glendon, a flag station of
Southern Railway situated between Jackson and
Walker Springs in Clarke County, Alabama (Hopkins
1917; MacNeil 1944). Although the Glendon Lime-
stone type locality is in a different county than
site AWa-9, it is located only 15 km to the east and
the Glendon Limestone exposures at St. Stephens
Historical Park are considered a reference section for
the unit (Hopkins 1917). The geologic exposures at
site AWa-9 were extensively investigated by Glawe
(1967, 1969) and Mancini and Copeland (1986).
Glawe (1967) thoroughly mapped and described the
hillside outcrops at our specific collecting site, and
he recognized five distinct lithologic units within
the exposed section that he designated beds 8–12.
The samples examined herein were collected from
an exposure of the Glendon Limestone Member lo-
cated at the top of bed 9 of section 2 of Glawe (1967;
Text-fig. 2. Geologic section of site AWa-9 showing the sampled interval. Section modified from Glawe (1967). NP Zones follow Mancini and
Tew (1992).
FISHES FROM THE OLIGOCENE OF ALABAMA 5
Text-fig. 2), and this particular bed is comprised of
three irregularly indurated coquinoid and crystalline
limestone ledges that weather into indurated rocks
containing large tubular cavities (known locally as
‘horse bone’ weathering). The limestone ledges are
interbedded with cream-colored fossiliferous, argil-
laceous and glauconitic marl that weathers brown
and contains a shell hash composed of the macro-for-
aminiferan Lepidocyclina Gümbel, 1870 and bivalves
like Ostrea vicksburgensis (Conrad, 1848), Chlamys
duncanensis (Mansfield, 1934), Chlamys anatipes
(Morton, 1833), Pecten perplanus poulsoni Morton,
1834 (in the lower beds), P. perplanus byramensis
Gardner, 1945 (in the upper beds), and P. howeri
mariannensis Glawe, 1969 (Glawe 1967; Szabo et
al. 1988; Mancini and Tew 1992), and occasional
vertebrate remains (this report). The specimens we
examined were collected in situ from the top lime-
stone ledge of this unit. Although the lower ledges are
visible at the base of this section, they are exposed as
part of sheer 20 m high cliff that cannot be sampled
without the aid of rappelling equipment.
The age of the Glendon Limestone Member has
been disputed. Siesser (1983a, b) placed the upper
part of the Marianna Limestone and the entirety of
the Byram Formation (including all three members)
within calcareous nannoplankton Zone NP22 of
Martini (1971) based on the absence of Cyclo cocco-
lithus formosus Kamptner, 1963 and the presence of
Lanternithus minutus Stradner, 1962. Later, Hazel
et al. (1980) placed the upper part of the Glendon
Limestone Member, the unnamed marl member,
and the lower part of the Bucatunna Clay Member
within zones NP22 and 23 of Martini (1971). Mancini
and Tew (1992) placed the entirety of the Marianna
Limestone and Byram Formation within zones NP22
and 23, and this latter interpretation is followed here
(Text-fig. 2). Mancini and Tew (1992) also placed
these units within the Pseudohastigerina micra
planktonic foraminiferal interval zone and inter-
preted the fossiliferous limestones of the Glendon
Limestone Member to represent a TO1.1 type 2 dep-
ositional sequence (sensu Mitchum et al. 1977; Baum
and Vail 1988) and highstand systems tract regressive
deposits.
Huddlestun (1993) documented two separate fora-
miniferal populations within the Glendon Limestone,
with those in Alabama generally representing moder-
ately deep-water assemblages, whereas the flanking
Georgia and Mississippi populations are character-
ized by more shallow-water foraminifera. This latter
interpretation is corroborated by the recent discovery
of sea grasses within the lower Glendon Limestone
in Rankin County, Mississippi, which were likely
deposited after the Vicksburg highstand (James Star-
nes, pers. comm.).
METHODS
Four bulk samples weighing approximately
10–15 kg each were collected by two of the authors
(JAE and DJC) from site AWa-9, Washington County,
AL, USA (Text-fig. 1) during the summer and fall
of 2019 and the winter of 2020. The samples were
collected from the base of a hillside outcrop at site
AWa-9, and specifically from an unconsolidated lens
exposed at the top of the Glendon Limestone Member
of the Byram Formation at the top of bed 9 of sec-
tion 2 of Glawe (1967; Text-figs 1 and 2). The bulk
field samples were processed in the laboratories at
McWane Science Center (MSC) in Birmingham, AL,
USA and the South Carolina State Museum (SC) in
Columbia, USA, where they were wet screened down
to a 0.25 mm mesh (No. 60 USA Standard Testing
sieves) to ensure the recovery of both macro- and
micro-vertebrate remains. The resulting concen-
trates were dried, and fossils were hand-picked us-
ing magnification. Figured specimens that exceeded
0.5 cm in greatest dimension were photographed
with a Nikon D80 camera with Tamron macro lens.
Specimens smaller than 0.5 cm were photographed
using an AmScope MU1000 camera mounted
to an AmScope 3.5x–90x stereo microscope and
10MB TIFF images were captured using AmScope
Toupview software version 3.7. All photographs were
rendered in Photoshop CC 2017 software as part of
the production of the presented figures. Whenever
possible, vertebrate remains were compared directly
to comparative skeletal remains housed at MSC,
SC, and the otolith collection of one of the authors
(GLS) in West Monroe, Louisiana, USA. All spec-
imens have been permanently deposited within the
scientific collections at either MSC or SC. Unless
otherwise specified, higher taxonomic rankings
used herein follow that of Nelson et al. (2016). In the
classification of the otoliths, ordinal names typically
follow Wiley and Johnson (2010), while the fami-
ly-group names and authors of Recent fishes follow
Van der Laan et al. (2014). Authors for genera and
species depend greatly upon Eschmeyer’s Catalog
of Fishes: Genera, Species, References (Fricke et al.
2019). Information from Froese and Pauly (2019) was
also utilized. Elasmobranch tooth terminology fol-
lows that of Shimada (2002) and Cappetta (2012),
and tooth group terminology follows that of Siverson
6 JUN A. EBERSOLE ET AL.
(1999). Teleost tooth terminology follows that of
Ebersole et al. (2019), and otolith terminology largely
follows that of Smale et al. (1995) and Nolf (2013).
SYSTEMATIC PALEONTOLOGY
Class Chondrichthyes Huxley, 1880
Subclass Euselachii Hay, 1902
Infraclass Elasmobranchii Bonaparte, 1838
Division Selachii Cope, 1871
Superorder Galeomorphi (sensu Nelson, Grande
and Wilson, 2016)
Order Carcharhiniformes Compagno, 1973
Family Scyliorhinidae Gill, 1862
Subfamily Premontreinae Cappetta, 1992
Genus Pachyscyllium Reinecke, Moths, Grant and
Breitkreuz, 2005
Pachyscyllium sp.
(Text- f ig . 3A)
MATERIAL: SC2019.61.30 (anterior tooth).
DESCRIPTION: The specimen is a complete anterior
tooth measuring 2 mm in total height and 1.6 mm in
crown width. The crown consists of a very tall, nar-
row main cusp f lanked by a single pair of large lateral
cusplets. The main cusp is sharply tapered and very
slightly distally inclined. The labial face is weakly
convex and smooth, whereas the lingual face is very
convex but also smooth. The crown foot is thickened
and overhangs the root. The lateral cusplets are large,
with the mesial cusplet noticeably wider and shorter
than the distal one. The cusplet faces are equally
convex, and very faint vertical plications occur on
the lingual face. The cutting edge is sharp, smooth,
and complete from the main cusp apex to the lateral
base of the cusplets. The root is low, particularly in
labial view, and bilobate. The lobes are very short and
diverging, with a sub-triangular attachment surface
that is weakly convex. A lingual nutritive groove
is long and narrow, with a central foramen. A large
margino-lingual foramen occurs on each side of a
large lingual protuberance.
REMARKS: Four Oligo-Miocene species of Pachy-
scyllium are currently recognized (see Reinecke et
al. 2005, 2011; Cappetta 2012; Collareta et al. 2020)
including P. albigensis Reinecke, Moths, Grant and
Breitkreuz, 2005, P. braaschi Reinecke, Moths, Grant
and Breitkreuz, 2005, P. dachiardii (Lawley, 1876),
and P. distans (Probst, 1879). The Glendon Limestone
tooth appears to differ from these other taxa by hav-
ing a labial crown base that overhangs the root to
a lesser degree than on P. albigensis, P. braaschi,
or P. dachiardii, and it lacks the labial crown orna-
mentation observed on P. distans. Although these
characteristics might suggest the Glendon Limestone
specimen represents a unique species, this is difficult
to ascertain based on a single specimen.
Pachyscyllium has been described from numer-
ous Paleogene and Neogene localities in Europe (see
Cappetta 2012), however North American occur-
rences appear extremely rare. The only such published
occurrence appears to be by Case (1980, pl. 5, figs. 1,
2), who described two P. distans teeth (as Scyliorhinus
distans) from the Trent Formation in North Carolina,
USA. Although Case (1980) reported these specimens
as being derived from the lower Miocene (Aquitanian)
Trent Formation, this formational name was previ-
ously abandoned (see Ward et al. 1978), and the de-
scribed deposits are part of the River Bend Formation,
which is regarded as early Oligocene in age (Rossbach
and Carter 1991). Nevertheless, the teeth figured by
Case (1980) differ from the Glendon Limestone tooth
by having robust enameloid folds at the labial crown
foot. With only a single, incompletely preserved tooth
available to us, specimen SC2019.61.30 is here not
speciated. However, the lack of labial ornamenta-
tion on this specimen does suggests the presence of a
second species of Pachyscyllium in the Oligocene of
North America.
Family Hemigaleidae Hasse, 1878
Genus Hemipristis Agassiz, 1835
Hemipristis sp.
(Text- f ig . 3B)
MATERIAL: MSC 43055 (lower left anterior tooth).
DESCRIPTION: The tooth measures approximately
11.8 mm in total height. The main cusp is tall and
curved lingually and has a slight distal curvature.
Both labial and lingual faces of the main cusp are
strongly convex and smooth. The mesial and distal
cutting edges are restricted to the upper 25% of the
crown to where, in labial or lingual views, the upper
portion of the crown is flared mesio-distally and is
spade-shaped. Two conical cusplets are situated near
the distal crown base that are medially curved, and
the more apical cusplet is twice the height of the more
basally situated cusplet. A single small cusplet is pre-
served on the mesial edge; however, a small ridge-like
FISHES FROM THE OLIGOCENE OF ALABAMA 7
denticle is present below the mesial cusplet that is
likely analogous to a secondary cusplet. The mesial
cusplet is comparable in size to the more basal distal
cusplet. The root is tall, and lingually is nearly equal
in height to the main cusp. The lingual protuberance
on the root is robust and is incised by an elongated
nutritive groove that extends basally to a U-shaped in-
terlobe area. The root is mesiodistally compressed and
the root lobes are short and only slightly divergent.
REMARKS: MSC 43055 was directly compared
with the dentition of a Recent Hemipristis elongata
(Klunzinger, 1871) (MSC 42327), and the specimen
compares very favorably to that of a lower left ante-
rior tooth. In Alabama, teeth belonging to Hemipristis
curvatus Dames, 1883 have been confirmed from
the Bartonian Gosport Sand (Ebersole et al. 2019),
and this species is also present within the Priabonian
(upper Eocene) Yazoo Clay and Rupelian Marianna
Text-fig. 3. Scyliorhinidae, Hemigaleidae, and Carcharhinidae teeth from the Oligocene of site AWa-9, Washington County, Alabama.
A – Pachyscyllium sp.; SC2019.61.30, tooth in (1) labial, (2) distal, (3) lingual and (4) basal views. Scale bar = 1 mm. B – Hemipristis sp.;
MSC 43055, lower tooth in (1) labial, (2) lingual and (3) distal views. Scale bar = 5 mm. C, D – Negaprion aff. N. gilmorei (Leriche, 1942);
C – MSC 43057, lower tooth in (1) labial, (2) distal and (3) lingual views. Scale bar = 4 mm; D – SC2019.61.31, upper tooth in (1) labial and
(2) lingual views. Scale bar = 5 mm. E –Physogaleus sp.; MSC 43056, tooth in (1) labial and (2) lingual views. Scale bar = 5 mm.
8 JUN A. EBERSOLE ET AL.
Limestone (JAE, unpublished data). Additionally,
teeth belonging to Hemipristis serra Agassiz, 1843
have been confirmed from Pliocene Graham Ferry
Formation equivalent deposits in southwest Alabama
(Ebersole et al. 2017; Stringer et al. 2020b). Although
these species have morphologically similar teeth,
those of H. curvatus are smaller in overall size and
have fewer mesial denticles. Traditionally, H. curvatus
teeth have been reported from upper Eocene deposits,
whereas H. serra has been documented from the late
Oligocene through the early Pleistocene (Adnet et al.
2007; Cicimurri and Knight 2009; Cappetta 2012).
Unfortunately, we are unable to determine with
confidence if the single small specimen available
to us represents H. curvatus, H. serra, or a transi-
tional form between the two (i.e., Adnet et al. 2007).
Additional teeth, especially those from the upper lat-
eral files, will be needed to ascertain which species of
Hemipristis is present within the Glendon Limestone
Member of Alabama.
Family Carcharhinidae Jordan and Evermann, 1896
Genus Negaprion Whitley, 1940
Negaprion aff. N. gilmorei (Leriche, 1942)
(Text-fig. 3C, D)
MATERIAL: MSC 43057 (lower right lateral tooth),
SC2019.61.31 (upper left lateral tooth).
DESCRIPTION: Specimen MSC 43057 is small,
measuring 5 mm in width and 3 mm in total height.
The main cusp is short and triangular and has a very
slight distal inclination. The main cusp has a slightly
convex labial face and is strongly convex lingually.
The crown is lingually bent in mesial and distal
views, and the enameloid is smooth. The crown base
is strongly sloped and extends onto elongated mesial
and distal shoulders, but does not extend to the me-
sial and distal edges of the root. The root lobes are
rounded and strongly divergent. In lingual view, the
height of the root is slightly shorter than the height of
the crown. The root has a rounded lingual protuber-
ance that is bisected by a shallow nutritive groove.
The interlobe area is wide and shallow, and a small
distal notch is present.
SC2019.61.31 shows the effects of root etching.
The specimen measures 6.5 mm in width and 5 mm
in total height. The crown consists largely of a wide-
based, inclined cusp and low distal heel. The mesial
cutting edge is oblique, elongated, slightly sinuous,
and appears to be smooth. There is an inconspicuous
notch that we believe represents the transition to a
mesial heel, which is continuous with the apical part
of the mesial edge. The distal cutting edge is much
shorter, nearly vertical, and forms a pointed apex with
the mesial edge. The low distal heel is elongated and
forms a nearly 90º angle with the distal cutting edge.
The distal cutting edge is evidently also smooth. The
labial crown face is flat, whereas the lingual face is
very convex. The root is bilobate, with very thin and
widely separated lobes. A nutritive groove is visible
on the medial lingual root face.
REMARKS: Although the Glendon Limestone spec-
imens are small, they appear to be conspecific with
Negaprion gilmorei teeth that have been reported
from the Claibornian (late Ypresian to Bartonian) of
Alabama (Ebersole et al. 2019). This taxon has also
been confirmed from the Chattian Chickasawhay
Limestone in section 3 of Glawe (1967) at site AWa-9
(JAE and DJC, unpublished data). In contrast, upper
teeth of the similar Carcharhinus elongatus (Leriche,
1910) from the early (Rupelian) and late (Chattian)
Oligocene of Europe (Baut and Génault 1999;
Reinecke et al. 2001, 2005; Génault 2012) and C. gib-
besii (Woodward, 1889) from the Chattian of South
Carolina (Cicimurri and Knight 2009) have clearly
separated mesial and distal heels that are moderately to
coarsely serrated. The Glendon Limestone specimens
are tentatively assigned to Negaprion gilmorei due to
their diminutive size, poor preservation, and the small
number of teeth in our sample (n=2).
Genus Physogaleus Cappetta, 1980a
Physogaleus sp.
(Text- f ig . 3E)
MATERIAL: MSC 43056 (lateral tooth).
DESCRIPTION: Specimen MSC 43056 is weath-
ered, but as preserved measures 6.8 mm in width and
5.2 mm in greatest height. The main cusp is short,
triangular, and distally inclined. The mesial cutting
edge is elongated, slightly concave, and extends al-
most to the mesial edge of the root. Poorly preserved
denticulation occurs on the basal part of the mesial
edge, but otherwise the edge is smooth. The distal
cutting edge is smooth, slightly concave, and forms
an oblique angle with the distal heel. The distal heel
is short and separated from the distal cutting edge by
a notch. The distal edge of the crown extends to the
distal edge of the root. Two pronounced cusplets are
FISHES FROM THE OLIGOCENE OF ALABAMA 9
present on the distal heel. The root is bilobate with
divergent and rounded lobes, and the interlobe area is
shallow and U-shaped. An indistinct nutritive groove
occurs on a low lingual root protuberance.
REMARKS: Although specimen MSC 43056 is
poorly preserved, salient features visible on the spec-
imen include a slightly concave mesial cutting edge,
an upturned cusp apex, two distal cusplets, and irreg-
ular and poorly defined denticulations on the lower
half of the mesial cutting edge. This suggests that
MSC 43056 was from a lower jaw tooth file, as upper
teeth of Physogaleus tend to have a more convex
or sigmoidal mesial cutting edge, a distally directed
cusp apex, and often have more defined mesial den-
ticulation (see Pharisat 1991; Reinecke et al. 2001;
Ebersole et al. 2019).
Three species of Physogaleus have previously
been confirmed from Paleogene deposits in the Gulf
Coastal Plain of the USA, including the Eocene P.
alabamensis (Leriche, 1942), P. americanus Case,
1994, and P. secundus (Winkler, 1874) (see Case
1994; Ebersole et al. 2019). When compared to the
lower teeth of these taxa, MSC 43056 differs from
P. americanus by lacking the single, pronounced,
mesial denticle as seen on the type specimens (Case
1994, figs 214−218). MSC 43056 also differs from P.
alabamensis, which can have up to 12 distal cusplets
and has well-defined mesial denticulations that often
extend to nearly two-thirds the height of the me-
sial cutting edge (see Ebersole et al. 2019). Although
MSC 43056 appears to fall within the morpholog-
ical range of P. secundus, assignment to this taxon
is problematic because its Rupelian age is well out-
side the known stratigraphic range for this taxon.
Physogaleus secundus is generally regarded as a
middle Eocene taxon (see Cappetta 2012), a notion
that is supported by the Alabama fossil record, where
this species is common in the lower to middle Eocene
Claiborne Group (Ebersole et al. 2019) but appears
to be absent from Priabonian deposits in the state
(JAE, unpublished data). Although several Oligocene
Physogaleus species have been named from else-
where, meaningful comparisons to MSC 43056 are
difficult to conduct due to the poor preservation of
the tooth. With only a single specimen represented in
our sample, MSC 43056 is herein not speciated.
Division Batomorphi Cappetta, 1980b
Order Myliobatiformes Compagno, 1973
Suborder Myliobatoidei Compagno, 1973
Family Incertae sedis
Gen. et sp. indet.
(Text-fig. 4A)
MATERIAL: SC2019.61.32 (tooth crown).
DESCRIPTION: The root is not preserved on speci-
men SC2019.61.32, but the crown measures 1.3 mm
in width (mesio-distal) and 1 mm in length (la-
bio-lingual). The crown is roughly diamond-shaped
in occlusal view and the labial crown margin is
somewhat angular, whereas the lingual margin is
uniformly convex. The crown is low, with an oblique
labial face that is overall weakly concave. The basal
rim is thickened and forms a distinctive rim around
the concave surface along with the transverse crest.
Of note is a thin transverse ridge located on the
lower half of the labial face, which does not connect
with the lateral crown margins. The transverse crest
is massive, thick and flat-topped (as preserved), and
it divides the crown into nearly equal labial and
lingual halves. The lingual crown face is convex
mesio-distally but concave apico-basally, and the
enameloid is smooth. Although the root is not pre-
served, a profile view shows that it was located near
the lingual crown margin.
REMARKS: Herman et al. (2000) and Hovestadt
and Hovestadt-Euler (2010) noted that several ex-
tant Myliobatoidei taxa, within at least two fami-
lies, the Dasyatidae and Urolophidae, can have teeth
bearing a secondary labial transverse crest in addi-
tion to the primary transverse crest. Of the extant
genera they examined, which included Himantura
Müller and Henle, 1837a, Pteroplatytrygon Fowler,
1910, Ta eni ura Müller and Henle, 1837b, Urolophus
Müller and Henle, 1837a, Urolophoides Lindberg,
1930, and Urobatis Garman, 1913, Hovestadt and
Hovestadt-Euler (2010) observed that only the teeth
of Himantura and Urobatis lacked strongly devel-
oped labial ornamentation. Although the lack of la-
bial ornamentation, combined with an occlusal out-
line consisting of an angled labial margin and broadly
rounded lingual margin, appears to ally SC2019.61.32
with Urobatis (see Herman et al. 2000; Hovestadt
and Hovestadt-Euler 2010), additional specimens are
needed to further elucidate the identity and paleobi-
ology (i.e., gynandric heterodonty) of the Glendon
Limestone taxon.
Family Myliobatidae Bonaparte, 1835
Subfamily Myliobatinae Bonaparte, 1835
Genus Aetomylaeus Garman, 1908
10 JUN A. EBERSOLE ET AL.
“Aetomylaeus” sp.
(Text-fig. 4B, C)
MATERIAL: MSC 43053 (upper symphyseal tooth),
MSC 43061 (upper symphyseal tooth).
DESCRIPTION: The two specimens are incomplete.
As preserved, MSC 43053 measures 13.9 mm in me-
siodistal width and 3.7 mm in labiolingual length,
whereas MSC 43061 measures 11.1 mm in mesiodis-
tal width and 3.8 mm in labiolingual length. In oral
view the lateral angles are seen to be obtuse and lo-
cated closer to the labial margin. The occlusal crown
surface of both teeth is convex, and in labial view the
crown is thickest medially and thins toward the me-
sial and distal edges. The labial crown face is nearly
vertical and straight, and the crown foot slightly over-
hangs the root. The lingual face is lingually inclined
and straight. Both the labial and lingual crown faces
are ornamented with a fine network of reticulated
ridges basally, transitioning into irregular and bifur-
cating longitudinal ridges that extend to the occlu-
sal surface. The lingual crown foot is marked by
a thin transverse ridge. The root is polyaulocorhize
and consists of a series of thin lamellae that are sep-
arated by shallow nutritive grooves. The lamellae are
ablated, but they extend at least to the lingual crown
margin.
REMARKS: We interpret that both specimens were
from the upper dentition because their occlusal sur-
faces are convex, and the crown foot is straight. Teeth
from the lower dentition typically have a straight oc-
clusal surface, but a medially convex crown foot. The
labial crown face on MSC 43061 is unusually curved,
suggesting this tooth was at the very front of the tooth
plate.
Historically, fossil myliobatid teeth exhibiting the
morphology described above have been identified
as Myliobatis Cuvier, 1816. The Glendon Limestone
teeth can be differentiated from extant Myliobatis
and fossil specimens attributed to this genus in that
the lateral angles are obtuse and located closer to the
labial crown margin, the lingual transverse ridge is
thin and sharp, and the ornamentation on crown faces
consists of reticulated ridges (at least basally, but of-
ten on the entire surface). In contrast, the Myliobatis
and Myliobatis-like teeth generally have lateral an-
gles that are about 90º, the lingual transverse ridge is
thick and rounded, and the crown faces have longitu-
dinal ridges. The morphology also differs from spec-
imens attributed to Rhinoptera Cuvier, 1829 in the
same respects. The gross morphology of the Glendon
Limestone specimens is most similar to teeth of ex-
tant Aetomylaeus.
Although Hovestadt and Hovestadt-Euler (2013)
referred many fossil specimens previously identified
as Myliobatis to Pteromylaeus Garman, 1913, other
analyses have shown Pteromylaeus to be a junior
synonym of Aetomylaeus (Naylor et al. 2012; White
2014). The latter assignment was followed by Ebersole
et al. (2019), who demonstrated that the presence of
reticulated crown ornamentation on fossil myliobatid
teeth is a generic identifier for Aetomylaeus-like taxa.
Recent phylogenetic analyses by Villalobos-Segura
and Underwood (2020) indicated that the radiation
of Recent myliobatid genera occurred much more re-
cently than previously thought, with an estimated di-
vergence during the early to middle Miocene. From
a taxonomic perspective, this data suggests that the
usage of extant myliobatid generic names in the fossil
record should be restricted to Neogene and younger
fossils, but morphologically similar Paleogene taxa
should be referred to different genera within the mylio-
batid lineage. Because the number of fossil taxa within
this newly established ghost lineage are currently un-
known, MSC 43053 and MSC 43061 are herein as-
signed to “Aetomylaeus” sp. with the understanding
that they may belong to an undescribed Paleogene rep-
resentative within the Aetomylaeus lineage.
Infraclass Elasmobranchii indet.
(Text-fig. 4D, E)
MATERIAL: SC2019.61.33 (placoid scale), SC2019.
61.34 (placoid scale).
DESCRIPTION: Two scale morphotypes have been
recovered. The first morphotype (SC2019.61.33)
consists of a massive rhomboidal crown measuring
1.2 mm in width and 1.1 mm in length. The anterior
margin is thick and bears seven robust ridges on the
anterior face. The dorsal surface is smooth and flat.
The posterior edge is thin and sharp. The base is
not preserved but appears to have been only slightly
smaller in area than the crown.
The second morphotype (SC2019.61.34) consists
of a very thin, ovate crown measuring 1.1 mm in
greatest dimension. The crown has an even thick-
ness, with marginal faces being convex and the upper
surface being flat, and the enameloid is smooth. The
base is imperfectly preserved, but appears to have
been rather narrow and medially located.
REMARKS: SC2019.61.33 is very similar to material
FISHES FROM THE OLIGOCENE OF ALABAMA 11
described by Laurito Mora (1999) from the Miocene
of Costa Rica that was referred to Galeocerdidae.
However, the identification of taxa based on isolated
placoid scales is complex, as studies of extant chon-
drichthyans has shown that scale morphology can
vary greatly depending on gender, location on the
Text-fig. 4. Myliobatiformes teeth and Elasmobranchii indet. placoid scales from the Oligocene of site AWa-9, Washington County, Alabama.
A – Myliobatoidei indet.; SC2019.61.32, tooth in (1) occlusal, (2) basal, (3) labial and (4) profile views. Scale bar = 0.5 mm. B, C – “Aeto-
mylaeus” sp.; B – MSC 43053, tooth in (1) lingual, (2) profile, (3) labial, (4) basal and (5) occlusal views. Scale bar = 5 mm; C – MSC 43061,
tooth in (1) lingual, (2) profile, (3) labial, (4) basal and (5) occlusal views. Scale bar = 5 mm. D, E – Elasmobranchii indet. placoid scales in
(1) outer view. Scale bar = 0.5 mm. D – SC2019.61.33; E – SC2019.61.34.
12 JUN A. EBERSOLE ET AL.
body, and ontogenetic stage (Reif 1985; Cappetta
2012). As a result, the Glendon Limestone placoid
scales are not identified beyond the infraclass level.
Class Osteichthyes Huxley, 1880
Subclass Actinopterygii (sensu Goodrich, 1930)
Unranked Neopterygii Regan, 1923
Infraclass Holostei Müller, 1845
Division Teleosteomorpha Arratia, Scasso and
Kiessling, 2004
Subdivision Teleostei Müller, 1845
Supercohort Teleocephala de Pinna, 1996
Cohort Elopomorpha Greenwood, Rosen, Weitzman
and Myers, 1966
Order Elopiformes Sauvage, 1875
Family Phyllodontidae Dartevelle and Casier, 1943
Subfamily Paralbulinae Estes, 1969
Genus Paralbula Blake, 1940
Paralbula sp.
(Text-f ig. 5A, B)
MATERIAL: SC2019.61.45 (tooth), SC2019.61.46
(tooth).
DESCRIPTION: Specimens SC2019.61.45 and
SC2019.61.46 measure less than 1 mm in greatest
diameter. The teeth have a circular occlusal outline
with smooth enameloid, and have a conspicuous cin-
gulum around the crown margin. In profile view the
teeth have a flat-topped occlusal surface and convex
lateral edges. The teeth are low-crowned in that the
overall tooth height is less than their greatest occlu-
sal diameter. The enameloid does not extend to the
tooth base, exposing an irregular basal ring of den-
tine around both teeth. In basal view the teeth have a
small medially located pulp cavity that is framed by a
thick wall of dentine.
REMARKS: Specimens SC2019.61.45 and SC2019.
61.46 differ from other similar teeth in our sam-
ple by having the combination of a low and smooth
crown with enameloid cingulum, and small basal
pulp cavity. These characteristics suggest that the
specimens are referable to Paralbula, a phyllodontid
taxon that has been reported from various Paleocene
and Eocene localities around the world, including
Alabama (Weiler 1929; Blake 1940; Arambourg
1952; Estes 1969; Weems 1999; Schein et al. 2011;
Ebersole et al. 2019). Four species of Paralbula have
been recognized, including P. casei Estes, 1969, P.
marylandica Blake, 1940, P. salvani (Arambourg,
1952), and P. stromeri (Weiler, 1929). The smooth
crown enameloid of the Glendon Limestone teeth is
comparable to the condition seen on P. stromeri and
many P. marylandica teeth, whereas P. casei and P.
salvani teeth bear crown ornamentation consisting of
granulation and concentric ridges. Although the teeth
of P. marylandica can also have a granular ornamen-
tation, it is much less apparent than on P. casei or
P. salvani, and many P. marylandica teeth can lack
ornamentation altogether (Blake 1940; Estes 1969;
Ebersole et al. 2019).
Text-fig. 5. Phyllodontidae and Albulidae teeth from the Oligocene
of site AWa-9, Washington County, Alabama. A, B – Paralbula sp.;
A – SC2019.61.45, tooth in (1) occlusal, (2) basal and (3) profile
views. Scale bar = 1 mm; B – SC2019.61.46, tooth in (1) occlusal,
(2) basal and (3) profile views. Scale bar = 0.5 mm. C – Albula sp.;
SC2019.61.4, tooth in (1) occlusal and (2) profile views. Scale bar
= 1 mm.
FISHES FROM THE OLIGOCENE OF ALABAMA 13
Paralbula marylandica is the only smooth-
crowned species to be reported in North America,
but the stratigraphic range of this taxon is thus far
only known to extend from the lower Paleocene to
middle Eocene (Blake 1940; Estes 1969; Weems
1999; Schein et al. 2011; Ebersole et al. 2019).
Paralbula stromeri is the only member of the genus
known to have a range that extends into the upper
Eocene (Priabonian), but this species has thus far
only been confirmed from Egypt and the United
Kingdom (Weiler 1929; Estes 1969). Estes (1969)
noted the morphological similarities between the
teeth of P. marylandica and P. stromeri, but was
able to differentiate the two taxa based on their ba-
sibranchial plates (skeletal features). The Glendon
Limestone specimens represent the first Oligocene
record of Paralbula, but with only two incomplete
teeth in our sample, we cannot determine if they rep-
resent new records of P. marylandica or P. stromeri,
or an undescribed Oligocene species.
Order Albuliformes Greenwood, Rosen, Weitzman
and Myers, 1966
Family Albulidae Bleeker, 1849
Subfamily Albulinae Bleeker, 1849
Genus Albula Scopoli, 1777
Albula sp.
(Text-fig. 5C)
MATER IAL: SC2019.61.44 (tooth).
DESCRIPTION: The specimen has a sub-circular
occlusal outline and measures slightly under 1 mm
in greatest diameter. The occlusal surface is flat and
lacks enameloid due to in vivo usage. In profile view
the edges of the tooth taper basally. The crown mar-
gins bear a thin layer of smooth enameloid. In basal
view, there is a very small, circular, medially located
pu lp cavit y.
REMARKS: The morphology of the specimen is
similar to that of Albula oweni (Owen, 1845), a taxon
that occurs in lower to middle Eocene Claibornian
strata of Alabama (Ebersole et al. 2019). The mor-
phological similarity to Albula oweni is based on the
basally tapering sides of the tooth. This characteristic
differentiates SC2019.61.44 from all other Paleogene
teleost teeth that have been reported from Alabama,
including Albula eppsi White, 1931, which has teeth
with evenly convex lateral edges (Ebersole et al.
2019). Due to the poor preservation of the single tooth
in our sample, specimen SC2019.61.44 is conserva-
tively referred only to Albula sp.
Order Anguilliformes Goodrich, 1909
Suborder Congroidei Kaup, 1856
Family Congridae Kaup, 1856
Subfamily Bathymyrinae Böhlke, 1949
Genus Ariosoma Swainson, 1838
Ariosoma nonsector Nolf and Stringer, 2003
(Text-fig. 6A)
MATERIAL: MSC 43054.2 (sagitta), MSC 43059.9
(sagitta), SC2019.61.3 (sagitta), SC2019.61.4 (sagitta),
SC2019.61.28 (incomplete sagitta).
DESCRIPTION: The sagittae are primarily oval in
outline (sensu Smale et al. 1995), but a very prom-
inent dorsal dome results in a more rounded shape.
The height/length ratios range from 0.80–0.86. The
margins are typically smooth, and the anterior margin
is rounded, but not broadly. The anterodorsal margin
is convex with a slight concavity occurring just be-
fore the prominent dorsal dome. The dorsal margin
consists of a high, rounded dome, which is primarily
medially located. The posterodorsal margin is gener-
ally slightly concave. The posterior margin is typically
tapered, often pointed, and the ventral rim is usually
broadly rounded and marked by a distinctive, nearly
medially located angle. The inner face is smooth and
convex, except for some irregular depressions in the
upper portion of the dorsal area. The sulcus is wide,
only slightly incised, and extends from near the ante-
rior margin to near the posterior margin (about 85% of
the length of the sagitta). The anterior portion of the
sulcus is lower than the posterior portion. Except for
the dorsal extremity of the ostial channel, the sulcus is
filled entirely with colliculum, and the ostial channel
curves backward. There is no conspicuous division
of the ostial and caudal portions of the sulcus, and
the posterior end of the sulcus is broadly tapered and
somewhat widened ventrally. There are no indications
of a ventral furrow. The outer face is smooth and con-
vex except for an area near the posterior end, where a
shallow and dorsoventrally oriented depression occurs.
REMARKS: The Oligocene specimens assigned to
Ario soma nonsector have several characteristics in
common with Ariosoma as defined by Schwarzhans
(2019). These include an S-shaped sulcus (see
Schwarz hans 2019, fig. 2), a middorsal expansion (i.e.,
dorsal dome), and a backward curving ostial channel.
14 JUN A. EBERSOLE ET AL.
The Eocene specimens assigned to Ariosoma nonsec-
tor by Nolf and Stringer (2003) have the backward
curving ostial channel and the middorsal expansion.
However, the sulcus is not S-shaped as seen in the
specimens illustrated by Schwarzhans (2019). The
Eocene specimens are believed to be a precursor to the
Oligocene specimens of Ariosoma nonsector. It should
be noted that the Oligocene specimens may represent
a new species of Ariosoma, but we feel this determi-
nation is not warranted based on the preservation and
limited number of specimens in our sample. Large
adult specimens of Ariosoma nonsector and the coeval
Paraconger sector Koken, 1888 are rather distinctive,
but smaller adult and particularly juvenile specimens
can be difficult to distinguish. In fact, Koken’s (1888)
type suite of Otolithus (Platessae) sector is actually a
mixture of these two taxa (see his pl. 17, fig. 14 for P.
sector, and figs 15 and 16 for A. nonsector). However,
P. sector has a greater length compared to its height
(i.e., is more elongate) and a has a narrower sulcus than
on A. nonsector. The latter taxon was separated from
P. sector by Nolf and Stringer (2003) based on their
examination of over 5,500 late Eocene otoliths from
the Yazoo Clay in Louisiana, USA.
Unranked Clupeocephala Patterson and Rosen, 1977
Cohort Eutelostei Rosen, 1985
Superorder Acanthopterygii Greenwood, Rosen,
Weitzman and Myers, 1966
Series Berycida (sensu Nelson, Grande and Wilson,
2016)
Order Trachichthyiformes (sensu Nelson, Grande
and Wilson, 2016)
Suborder Anoplogastroidei (sensu Nelson, Grande
and Wilson, 2016)
Family Diretmidae Gill, 1893
Genus Diretmus Johnson, 1864
Diretmus? sp.
(Text-fig. 6B)
MATERIAL: MSC 43059.10 (sagitta).
DESCRIPTION: The sagitta is unique by being tall
(sensu Smale et al. 1995). The height is approximately
twice its length, and the dorsal and ventral margins
are both conspicuously tapered. The margins appear
fairly smooth, probably due to erosion, but there is ev-
idence of several lobes on some margins. The anterior
margin is short, nearly vertical, and characterized by
the sulcus opening. The anterodorsal margin is steep
and outwardly curved. The dorsal margin is very short
and pointed, and the posterodorsal margin is steep and
slightly convex (and possibly incurved). The posterior
margin is short, slightly rounded, and nearly vertical.
The posteroventral margin is steep, nearly straight,
and slants from the posterior to the ventral margin.
The ventral margin is represented by a short, rounded
point, whereas the anteroventral margin is steep and
slightly outwardly curved. There is a prominent sulcus
extending across almost 90% of the otolith length.
The details of the sulcus are difficult to discern to due
to erosion, but the ostium is taller than the cauda and
ventrally expanded. The cauda is approximately twice
the length of the ostium. There is a distinct circular de-
pressed area above the cauda. The outer face is slightly
convex and appears to have been highly sculptured
(erosion has obliterated the features).
REMARKS: The shape of the sagitta of Diretmus?
sp. is unusual and uncommon, especially for otoliths
of the Gulf Coastal Plain. This species was originally
described by Müller (1999) as “genus Caproidarum”
serratus, but Nolf (2013) described it as “Diretmida”
serratus and placed it in the family Diretmidae.
Ebersole et al. (2019) described the only other fossil
diretmid specimen known from Alabama, a single
sagitta of Diretmus cf. D. serratus (Müller, 1999)
recovered from the Eocene (Lutetian and Bartonian)
Text-fig. 6. Congridae, Diretmidae , and Ophidiidae otoliths from
the Oligocene of site AWa-9, Washington County, Alabama. A –
Ariosoma nonsector Nolf and Stringer, 2003; SC2019.61.3, right
sagitta in (1) inner and (2) dorsal views. Scale bar = 5 mm. B –
Diretmus sp.; MSC 43059.10, right sagitta in (1) inner view. Scale
bar = 1 mm. C – Preophidion meyeri (Koken, 1888); MSC 43054.5,
right sagitta in (1) inner and (2) dorsal views. Scale bar = 3 mm.
FISHES FROM THE OLIGOCENE OF ALABAMA 15
Lisbon Formation. Unfortunately the poor preser-
vation of the Glendon Limestone specimen inhibits
more specific taxonomic identification.
Series Percomorpha (sensu Nelson, Grande and
Wilson, 2016)
Subseries Ophidiida (sensu Nelson, Grande and
Wilson, 2016)
Order Ophidiiformes (sensu Nelson, Grande and
Wilson, 2016)
Suborder Ophidioidei (sensu Nelson, Grande and
Wilson, 2016)
Family Ophidiidae Rafinesque, 1810
Subfamily Incertae sedis
Genus Preophidion Frizzell and Dante, 1965
Preophidion meyeri (Koken, 1888)
(Text- f ig . 6C )
MATERIAL: MSC 43054.3 (sagitta), MSC 43054.5
(sagitta), MSC 43054.6 (sagitta), MSC 43059.4 (sa-
gitta), MSC 43067.1 (sagitta), MSC 43067.6 (sagitta),
SC2019.61.9 (sagitta), SC2019.61.10 (sagitta), SC2019.
61.11 (sagitta), SC2019.61.12 (sagitta), SC2019.61.13
(sagitta), SC2019.61.17 (sagitta).
DESCRIPTION: The sagitta is oblong to elliptic in
shape (sensu Smale et al. 1995) with height/length
ratios ranging from approximately 0.44–0.53. Adult
specimens measure up to 6 mm in length, and both
adult and juvenile specimens tend to have smooth
margins. The inner face of the sagitta is smooth
and convex. The anterior margin is bluntly pointed,
whereas the anterodorsal margin is long and slightly
arched. A rounded anterodorsal dome is usually vis-
ible, and the dorsal margin is short and almost hor-
izontal. The posterodorsal margin is very slightly
arched and longer than the anterodorsal margin. The
posterior margin is thinly pointed, and the ventral
margin is evenly and broadly rounded. The sulcus is
lightly impressed, divided, completely enclosed, and
marked by incised lines. The ostium is about equal in
length and height to the cauda and has sides that are
nearly parallel. The anterior end of ostium is sharply
pointed and extends almost to the anterodorsal mar-
gin. The ostium is not excavated but is filled with
colliculum. With the exception of a slight ventral
constriction near the junction with the ostium, the
sides of the cauda are nearly parallel. The intersection
of the cauda and ostium is marked by a thin, slightly
inclined, incised line. The cauda is filled with col-
liculum and is not excavated. The posterior end of
the cauda is bluntly rounded and separated from the
posterior margin by a distinct, narrow border. There
is a shallow, elongated and irregularly depressed area
located medially, above the sulcus. A crista superior
is weakly developed, a crista inferior is either lack-
ing or very weakly developed, and a ventral furrow
is typically absent. The outer face is convex, with
the dorsal portion more strongly so, and is strongly
sculptured to undulated.
REMARKS: The otolith-based fossil cusk-eel genus
Preophidion was established by Frizzell and Dante
(1965). Preophidion is particularly widely distrib-
uted and often abundant in Eocene strata of the Gulf
Coastal Plain of the USA (Ebersole et al. 2019), but
the genus has also been reported from Oligocene
strata (Stringer et al. 2001; Stringer and Miller 2001).
In the Gulf Coastal Plain, Preophidion is known to
occur in Alabama, Georgia, Louisiana, Mississippi,
and Texas (Frizzell and Dante 1965; Breard and
Stringer 1995; Stringer and Miller 2001; Green and
Stringer 2002; Nolf and Stringer 2003; Stringer 2016;
Ebersole et al. 2019).
Subseries Gobiida Betancur-R et al., 2013
Order Gobiiformes (sensu Nelson, Grande and
Wilson, 2016)
Family Gobiidae Cuvier, 1816
Subfamily Gobiinae Cuvier, 1816
Genus Gobiosoma Girard, 1858
Gobiosoma? axsmithi sp. nov
(Text-fig. 7A, B)
urn:lsid:zoobank.org:act:C659AEB2-8B52-484D-
AD8D-04A3E606FA25
TYPE MATERIAL: Holotype (SC2019.61.18, sa-
gitta) and paratype (SC2019.61.19, sagitta).
OTHER MATERIAL: SC2019.61.171 (broken sa-
gitta), SC2019.61.172 (broken sagitta).
TYPE LOCALITY: Site AWa-9, St. Stephens
Historical Park, Washington County, Alabama, USA,
base of hillside at the top of section 2 of Glawe (1967).
TYPE STRATUM: Unconsolidated section at the
top of bed 9, section 2 of Glawe (1967), Glendon
Lime stone Member of the Byram Formation, lower
Oligocene (Rupe lian), zones NP22/23, Pseudo hasti-
gerina micra plank tonic foraminiferal interval zone.
16 JUN A. EBERSOLE ET AL.
DERIVATION OF NAME: Named in honor and
memory of Brian J. Axsmith, an American paleo-
botanist, paleoecologist, and professor of biology at
the University of South Alabama, Mobile, Alabama,
USA.
DIAGNOSIS: Sagittae are small (1 mm length), es-
sentially square in outline, with a height/length ratio
of 1.04. Specimens are almost plano-convex, with a
rounded dorsal margin and a ventral margin that is
almost straight and horizontal. The sulcus is small,
medially positioned, and inclined at about 15° to-
wards the anteroventral. The ostium is slightly ta-
pered and the cauda slightly shorter in length and
rounded posteriorly, with a raised, oblong-shaped
subcaudal iugum. The cauda length/ostium length
ratio is approximately 0.83 and the ostium height/
cauda height ratio 1.67. An oblong subcaudal iugum
is present. There is a prominent dorsal depression and
ventral furrow. The inner face very slightly convex to
almost flat, but the outer face is broadly convex.
DESCRIPTION: The outline of the sagitta is primar-
ily square (sensu Smale et al. 1995). The inner face
is somewhat convex and generally smooth (except
for the dorsal depression). The margins are typically
smooth and may be somewhat sharp in transversal
view. The anterior margin ranges from nearly straight
to slightly incurved, and the anterodorsal margin is
unevenly rounded and distinguished by a predorsal
projection. The dorsal margin is broadly rounded and
characterized by a weakly developed obtuse medial
angle. The posterodorsal margin is rounded, with a
postdorsal projection that is similar to the predorsal
projection. The posterior margin is incurved, and the
posteroventral margin is characterized by a slight
posteroventral projection or angle. The ventral mar-
gin is almost straight and horizontal, whereas the
anteroventral margin has a slight anteroventral pro-
jection or angle. There is a well-defined, somewhat
excavated, divided sulcus that slants approximately
15º from the posterodorsal margin to the anteroven-
tral margin. There appears to be a small ridge-like
crista superior located above the sulcus, and the os-
tium is slightly longer and higher than the cauda. The
ostium is well separated from the anterior margin,
and is tapered and somewhat pointed. There is an
indication of a slight ostial lobe. The slightly smaller
cauda ends well before the posterior margin and is
distinguished by an evenly rounded posterior tip.
A diagnostic raised, oblong-shaped subcaudal iugum
is present. There is a prominent and rather deep dor-
sal depression that appears rounded or irregular. The
ventral furrow is very distinct and curves from the
anterior to the posterior of the sulcus. The outer face
is thickest near the center, slightly convex, and has
smooth, rounded structures.
REMARKS: Nolf et al. (2006) reported a gobioid sa-
gitta from Eocene (Ypresian) strata of India that thus
far represents the oldest known member of the order.
Gobioid otoliths are very rare in all known teleost
assemblages until the late Eocene. Of the few speci-
mens that have been reported, Ebersole et al. (2019)
described a single sagitta from the Lisbon Formation
(Lutetian to Bartonian) of Alabama, and Nolf and
Stringer (2003) reported 57 specimens (as “genus
Gobiidarum” vetustus) from the upper Eocene Yazoo
Clay of Louisiana. Schwarzhans (pers. comm.) noted
that the ostial and caudal colliculi are always fused
in the gobies, and the ventral furrow curves around
the back of the cauda and coalesces with the dorsal
depression (or fades).
As the diversity and abundance of Eocene and
Oligocene gobioid otoliths is very low in the Gulf
Coastal Plain, younger fossil gobioid otoliths and
Recent specimens were compared to the Glendon
Text-fig. 7. Gobiosoma? axsmithi sp. nov. from the Oligocene of site
AWa-9, Washington County, Alabama, right sagittae in (1) inner, (2)
outer and (3) dorsal views. A – SC2019.61.18, holotype. Scale bar
= 1 mm. B – SC2019.61.19, paratype. Scale bar = 0.5 mm.
FISHES FROM THE OLIGOCENE OF ALABAMA 17
Limestone specimens. Critical comparisons were
made of the predorsal angle of projection, postdorsal
projection, postventral angle, preventral projection,
and the presence/absence (and characteristics) of the
subcaudal iugum. The most extensive and compre-
hensive comparative study of fossil gobioids is that
of Schwarzhans et al. (2020) for the middle Miocene
of the Czech Republic, Slovakia, and Poland. Their
work included the gobiid groups of Aphia Risso, 18 27,
Priolepis Valenciennes in Cuvier and Valenciennes,
1837, Gobius Lin n æus , 1758 , Thorogobius Miller,
1969, and Pomatoschistus Gill, 1863b, all of which
were compared to the Alabama gobiids. The Alabama
specimens were also compared to Recent and fossil
gobiids reported by Arellano et al. (1995), Nolf (2013,
pls 316–325), and Gut et al. (2020). In a study of early
Oligocene otoliths from Japan, Schwarzhans et al.
(2017) illustrated additional Recent gobiid sagittae
from Chaenogobius Gill, 1859 and Gymnogobius
Gill, 1863b. The fossil forms, however, were placed
in a separate and new genus and species.
Based on the aforementioned critical character-
istics, we believe that the Alabama otoliths compare
most favorably to the Recent genus Gobiosoma,
which has a very similar plesiomorphic sagitta.
Comparisons of the Glendon Limestone specimens
to sagittae of Recent species of Gobiosoma, such as
G. aceras (Ginsburg, 1939), G. bosc (de Lacépède,
1798), G. chiquita (Jenkins and Evermann, 1889),
G. robustum Ginsburg, 1933, G. schultzi (Ginsburg,
1944), G. seminudum (Günther, 1861), and G. yu-
catanum Dawson, 1971, indicate that the Alabama
Gobiosoma represents a new fossil species and is
herein described as such. The addition of the “?” at
the end of the generic name indicates that, although
the fossil species compares well to the extant genus, it
is possible that the form represents an unknown fossil
genus. This can only be determined with certainty
through the discovery of skeletal remains that are
associated with in situ otoliths.
Subseries Ovalentaria Smith and Near
in Wain wright et al., 2012
Order Istiophoriformes Betancur-R et al., 2013
Family Sphyraenidae Rafinesque, 1815
Genus Sphyraena Artedi, 1793
Sphyraena sp.
(Text- f ig . 8A–C)
MATERIAL: SC2019.61.35 (tooth), SC2019.61.36
(tooth apex), SC2019.31.37 (laniary tooth apex).
DESCRIPTION: SC2019.61.35 is the most complete
specimen, consisting of an ablated crown measuring
8 mm in total height. The crown is lanceolate in labial
view, with sharp, smooth, convex anterior and pos-
terior carinae. The labial face is nearly flat, but the
lingual face is convex, particularly near the crown
base. In anterior/posterior view there is a slight me-
dial curvature. In basal view there is a small medi-
ally located pulp cavity. Specimens SC2019.61.36 and
SC2019.31.37 are represented by tooth apices that are
similar to that described above. SC2019.31.37, how-
ever, is not symmetrical in labial or lingual views,
indicating it is a laniary tooth.
REMARKS: The Glendon Limestone Member teeth
are similar to several fossil taxa previously described
from the Paleogene of Alabama, like Palaeo cybium
Monsch, 2005 and Scomberomorus de Lacépède,
1801 (Ebersole et al. 2019). However, the Glendon
Limestone Sphyraena teeth differ from Palaeocybium
by being more labio-lingually compressed and by
having less pronounced and pointed carinae in pro-
file view. These teeth also lack the basal thicken-
ing seen on Paleogene Scomberomorus teeth, and
the carinae extend to the base of the tooth (they do
not on Scomberomorus; see Ebersole et al. 2019).
Furt hermore, both Palaeocybium a nd Scomberomorus
lack the laniary tooth morphology that occurs in the
dentition of Sphyraena.
The Glendon Limestone teeth were compared di-
rectly to those of the extant Sphyraena barracuda
(Edwards in Catesby, 1771) (MSC 43215, SC2018.3.1)
and Sphyraena borealis DeKay, 1842 (MSC 43076).
Because the teeth have unserrated carinae, they
appear better aligned with the smaller members of
the genus, such as S. borealis, as opposed to the
much larger and serrated-toothed S. barracuda.
Furthermore, based on the size and shape of the
teeth, both SC2019.61.35 and SC2019.61.36 appear
to have been derived from the palatine or dentary.
SC2019.61.37, on the other hand, has a sharp anterior
carina and a posterior barb, features which can occur
on Sphyraena laniary teeth. Unfortunately, all of the
teeth in our sample are incomplete and we cannot
provide a more precise taxonomic assignment.
Order Pleuronectiformes Bleeker, 1859
Suborder Pleuronectoidei (sensu Nelson, Grande and
Wilson, 2016)
Superfamuly Pleuronectoidea Rafinesque, 1815
Family Paralichthyidae Regan, 1910
Genus Citharichthys Bleeker, 1862
18 JUN A. EBERSOLE ET AL.
Text-fig. 8. Sphyraenidae, Paralichthyidae, Sciaenidae, Sparidae, and Balistidae elements from the Oligocene of site AWa-9, Washington
County, Alabama, except when stated otherwise. A–C – Sphyraena sp.; A – SC2019.61.35, tooth in (1) labial and (2) profile views. Scale bar
= 5 mm; B – SC2019.61.36, tooth in (1) labial and (2) profile views. Scale bar = 1 mm; C – SC2019.61.37, laniary tooth in (1) labial and (2)
profile views. Scale bar = 1 mm. D – Citharichthys sp.; SC2019.61.22.1, right sagitta (reversed for comparison) in (1) inner and (2) dorsal
views. Scale bar = 1 mm. E – gen. et sp. indet.; SC2019.61.27, lapillus in (1) macular, (2) anti-macular and (3) side views. Scale bar = 0.5 mm. →
FISHES FROM THE OLIGOCENE OF ALABAMA 19
Citharichthys sp.
(Text- f ig . 8D)
MATERIA L: SC2019.61.22 (14 sagitt ae), SC2019.61.23
(43 sagitt ae).
DESCRIPTION: Sagittae of this taxon are typically
no larger than 3.0–3.5 mm in length, but the Glendon
Limestone Member specimens are much smaller,
with many of them only around 1.0 mm. The sagittae
tend to be relatively thin, and the outline is primarily
a compressed pentagon (sensu Schwarzhans 1999).
The margins range from smooth to occasionally un-
dulated, and the rims of the margins are usually
sharp. The anterior margin has an obtuse median an-
gle with no rostrum. The anterodorsal margin ranges
from nearly straight to concave. In contrast, the dor-
sal margin is almost horizontal and characterized by
a predorsal projection and postdorsal angle, which
is usually bent slightly outward. The posterodor-
sal margin is weakly concave and located between
the postdorsal angle and the posterior margin. The
posterior margin is distinguished by a pointed, out-
wardly projected posterior tip that occupies a slightly
supramedian position. The ventral margin is deeply
curved and has an obtuse medioventral angle. The
inner face is moderately convex and smooth, and
bears a very distinctive fusiform sulcus. The sulcus
is medially located, almost horizontal, and widens
just behind the centerline, with the anterior and pos-
terior portions being narrower. Distinction of the
ostium and cauda cannot be made because the col-
luculi are completely fused. If present, the ventral
furrow is indistinct. There are dorsal and ventral
depressions that nearly continuously connect around
the sulcus, forming a circumsulcal depression. The
outer face is generally flat to slightly concave, with
little ornamentation.
REMARKS: Citharichthys otoliths appear to be
rare in Alabama, as none were reported from the
lower-to-middle Eocene (Ypresian to Bartonian)
Claiborne Group in Alabama (Ebersole et al. 2019)
and only two Citharichthys macrops Dresel, 1885
sagittae were documented from Pliocene (Zanclean
to Piacenzian) Graham Ferry Formation equivalent
deposits in Alabama (Stringer et al. 2020b). The large
number of Citharichthys sagittae in the Glendon
Limestone Member assemblage (nearly 50% of the
total number of specimens) is unusual and may be
related to unique or unusual paleoenvironmental con-
ditions.
Citharichthys? sp.
(Text- f ig . 8E)
MATERIAL: SC2019.61.27 (2 lapilli).
DESCRIPTION: The utricular otoliths (or lapilli)
are very small and approximately 0.6 mm in length.
Both specimens appear to have an oval outline (sensu
Smale et al. 1995), although one is not as well pre-
served. The macular side (ventral side sensu Assis
2005) is noticeably convex, and it is smooth except
for the well-defined gibbus macula and the promi-
nentia marginalis. Although well defined, the gibbus
macula is not large relative to the size of the lapillus.
The linea basalis appears to be a single lobe. The
anti-macular side (dorsal side sensu Assis 2005) is
essentially flat and featureless.
REMARKS: The two Glendon Limestone Member
lapilli appear to be conspecific. Due to their unre-
markable morphology, lapilli are generally not uti-
lized in the study of fossil otoliths. However, there
are exceptions, including taxa within families like
Ariidae (sea catfishes) and Sciaenidae (drums and
croakers) (Nolf 2013; Schwarzhans et al. 2018;
Stringer and Bell 2018; Schwarzhans and Stringer
2020). Although the lapilli for many species of A riidae
and Sciaenidae are known (Chao 1978; Aguilera et al.
2020), the Glendon Limestone lapilli do not appear to
conform to any of these taxa. Our two specimens are
similar to lapilli of Recent Citharichthys arctifrons
Goode, 1880 (see Campana 2004, p. 204), and it is in-
teresting to note that Citharichthys comprises almost
50% of the otoliths within the Glendon assemblage.
Therefore, there is a relatively good possibility that
the lapilli represent this genus, and we tentatively
assign them to Citharichthys.
F – Syacium sp.; SC2019.61.21, right sagitta in (1) inner and (2) dorsal views. Scale bar = 1 mm. G, H – Sciaena pseudoradians (Dante and
Frizzell in Frizzell and Dante, 1965); G – MSC 43059.1, right sagitta (reversed for comparison) in (1) inner and (2) dorsal views. Scale bar =
5 mm; H – MSC 43507, from the Glendon Limestone of Mississippi (included for comparison), right sagitta (reversed for comparison) in (1) in-
ner and (2) dorsal views. Scale bar = 5 mm. I, J – Aplodinotus gemma Koken, 1888; I – SC2019.61.1, right sagitta (reversed for comparison) in
(1) inner and (2) dorsal views. Scale bar = 5 mm; J – MSC 43506, from the Glendon Limestone of Mississippi (included for comparison), right
sagitta (reversed for comparison) in (1) inner and (2) dorsal views. Scale bar = 5 mm. K – Sciaenops? sp.; SC2019.61.43, tooth in (1) profile
and (2) basal views. Scale bar = 1 mm. L – Sparus? elegantulus (Koken, 1888); MSC 43067.5, right sagitta (reversed for comparison) in (1) in-
ner and (2) dorsal views. Scale bar = 2 mm. M – Balistidae indet.; SC2019.61.38, tooth in (1) occlusal and (2) profile views. Scale bar = 1 mm.
20 JUN A. EBERSOLE ET AL.
Genus Syacium Ranzani, 1842
Syacium sp.
(Text-fig. 8F)
MATERIAL: SC2019.61.20 (sagitta), SC2019.61.21
(sagitta), SC2019.61.25 (sagitta).
DESCRIPTION: The sagittae are primarily square
(sensu Smale et al. 1995). The inner face is smooth
and weakly convex. The margins are generally
smooth, but can be variable and irregular. The an-
terior margin ranges from nearly vertical to slightly
outward slanted from the anteroventral margin to the
anterodorsal margin. There is no rostrum because the
sulcus does not extend to the anterior margin. The an-
terodorsal margin is distinguished by a slightly acute,
prominent angle, and the dorsal margin is irregular
and almost horizontal. The posterodorsal margin is
distinguished by a posterodorsal projection that is
conspicuous, but not very large. The posterior mar-
gin is typically straight, but it can be nearly vertical
or slightly slanted toward the posteroventral margin.
The posteroventral margin is characterized by a dis-
tinct, slightly rounded angle that joins with the pos-
terior margin. The fairly smooth ventral margin is
nearly straight and horizontal, and the anteroventral
margin is slightly rounded and forms a distinct an-
gle with the anterior margin. The inner face bears a
highly specialized, fusiform sulcus that slants down-
ward from the posterodorsal margin, almost to the
anteroventral margin. Additionally, the fusiform sul-
cus widens just behind its midline, with the anterior
and posterior portions being narrower. Although the
sulcus extends across approximately 75% of the in-
ner face, it is narrow and represents only about 20%
of the of the height of the inner face. The sulcus is
divided into ostial and caudal areas, with the ostium
located near the lower portion of the anterior and
anteroventral margins, but not reaching the margins.
The ostium is narrow and constricted at the anterior
and posterior, and the anterior portion is tapered and
almost pointed. The cauda is longer and wider than
the ostium, and the anterior of the cauda is tapered,
whereas the center portion is enlarged. The poste-
rior of the cauda is tapered with a somewhat pointed
tip, and the cauda appears to be excavated slightly
deeper than the ostium. The ostium and cauda are
conspicuously connected or fused, and colliculum
occurs within. There is a marked circumsulcal de-
pression extending from above the ostium to around
the cauda, ending below the ostium. This extended
depression forms an elevated flattened area for the
fusiform sulcus. There is no visible ventral furrow.
The outer face is relatively flat on the dorsal and ven-
tral areas, but slightly convex in the center.
REMARKS: The fossil record of Syacium in Ala-
bama was previously represented by a single sagitta
(Stringer et al. 2020b), and the genus appears to be
rare in all reported assemblages across the Gulf and
Atlantic coastal plains of the USA. With the exception
of the Alabama specimens and a single Syacium oto-
lith reported by Stringer (1992) from the Mississippi
River mudlump islands (late Pleistocene−early
Holocene), very few other specimens are known (Nolf
and Stringer 2003; Nolf 2013; Stringer et al. 2017;
Stringer and Bell 2018; Ebersole et al. 2019; Stringer
and Shannon 2019; Stringer and Hulbert 2020).
Family Paralichthyidae indet.
MATERIAL: SC2019.61.26 (eroded sagitta).
REMARKS: The specimen attributed to Para lich thyi-
dae indet. is significantly eroded, but the discernible
features align it with this family. The outline and gen-
eral morphology of the specimen is very similar to
taxa like Citharichthys or Syacium, both of which are
present within our Glendon Limestone Member sam-
ple. The position and the shape of the fusiform sulcus
is also comparable to members of the Paralichthyidae.
Unfortunately, the preservation of SC2019.61.26 does
not allow for a more refined identification.
Order Acanthuriformes (sensu Nelson, Grande and
Wilson, 2016)
Suborder Sciaenoidei Betancur-R et al., 2013
Family Sciaenidae Cuvier, 1829
Genus Sciaena Linnæus, 1758
Sciaena pseudoradians (Dante and Frizzell in
Frizzell and Dante, 1965)
(Text-fig. 8G, H)
MATERIAL: MSC 43054.13 (sagitta), MSC 43059.6
(sagitta), MSC 43059.1 (sagitta), MSC 43059.11 (sa-
gitta).
DESCRIPTION: The sagittae are generally oval to
subrectangular in outline (sensu Smale et al. 1995),
and have approximate height/length ratios ranging
from 0.75–0.82 (however the ratios are affected by
the appreciable erosion of the anterior margin). The
FISHES FROM THE OLIGOCENE OF ALABAMA 21
inner face is convex and smooth. The margins are
primarily smooth, with the anterior margin being very
broadly and evenly rounded. The anterodorsal margin
is slightly convex and the dorsal margin slopes gently
downward anteriorly and posteriorly from a subtle ob-
tuse central angle. There is a conspicuous posterodor-
sal angle on almost all specimens. The posterior mar-
gin is almost straight and vertical, whereas the ventral
margin is broadly rounded. There is a very prominent
sulcus (heterosulcoid type) that extends across almost
95% of the inner face. The ostium extends for almost
50% the length of the sulcus, and the height of the
cauda is only about 25% the height of the ostium. The
ventral portion of the ostium is much more expanded,
and the ostium is filled with colliculum. The anterior
of the ostium is even with the anterior margin of the
sagitta, and the dorsal and ventral margins of the os-
tium tend to be constricted anteriorly. The cauda is
long and narrow, and it has a horizontal portion and
a sharply downturned portion. The horizontal portion
of the cauda is slightly shorter than the downturned
portion, and the angle between the two is close to 90°.
The downturned portion is tapered but still somewhat
rounded, and it almost reaches the posteroventral
margin. The outer face is generally weakly concave
and sculptured.
REMARKS: Unfortunately, all four of the specimens
we assigned to Sciaena pseudoradians are consider-
ably eroded, and they would have been more elongate
if not for abrasion and attrition on the thin edges of
the anterior margin. Additionally, the ventral edge
of the ostium would be longer and remain essen-
tially horizontal. The posterior margin is usually al-
most vertical, as is illustrated by Nolf (2013, pl. 277).
These features allow one to differentiate S. pseudo-
radians from the very similar and coeval Aplodinotus
gemma Koken, 1888 (see below). Additionally, al-
though larger specimens of S. pseudoradians develop
lower height/length ratios (i.e., more elongated), the
height/length ratios of larger specimens of A. gemma
tend to have higher height/length ratios (i.e., are more
rounded). These characteristics are well-illustrated
in figures of A. gemma in Nolf (2013, pl. 269), who
also noted the similarity of small A. gemma (around
5 mm) sagittae to those of small S. pseudoradians
specimens (see Nolf 2013, pl. 277). Because it is
typical for very small (juvenile) sciaenid otoliths to
be plesiomorphic and more difficult to identify, we
figure two well-preserved adult specimens from a
different locality (but same stratigraphic unit) for
comparison with our less adequately preserved spec-
imens from site AWa-9.
Genus Aplodinotus Rafinesque, 1819
Aplodinotus gemma (Koken, 1888)
(Text-fig. 8I, J)
MATERIAL: MSC 43059.2 (sagitta), SC2019.61.1
(sagitta), SC2019.61.2 (sagitta).
DESCRIPTION: The sagittae range from somewhat
oval to an almost elongated discoid outline (sensu
Smale et al. 1995), and generally have height/length
ratios ranging from 0.90–0.95. The inner face is con-
vex and smooth, with the greatest thickness in the
area between the ostium and the cauda. The mar-
gins are primarily smooth. The anterior margin is
broadly and evenly rounded, and the anterodorsal
margin is slightly convex. The dorsal margin slopes
gently downward anteriorly and posteriorly from a
very subtle obtuse central angle. There is a conspic-
uous posterodorsal angle on almost all specimens.
The posterior margin is almost straight and slants
slightly inward to varying degrees. The ventral mar-
gin is broadly rounded and a very prominent sulcus
(heterosulcoid type) extends across almost 95% of
the inner face. The ostium extends for over 50% of
the length of the inner face, and the height of the
cauda is only about 20% of the height of the ostium.
The ventral portion of the ostium is greatly expanded
toward the ventral margin, especially on large spec-
imens. The ostium is filled with colliculum, and its
anterior portion is even with the anterior margin of
the sagitta. The dorsal and ventral margins of the os-
tium generally constrict anteriorly. The cauda is long
and narrow and divided into a horizontal portion and
a sharply downturned portion. The horizontal por-
tion is slightly shorter than the downturned portion,
and the angle between the two is close to 90º. The
downturned portion of the cauda is tapered, some-
what rounded, extends almost to the posteroventral
margin, and is usually curved toward the posterior of
the ostium. The outer face is usually slightly concave
and sculptured.
REMARKS: According to Nolf (2003), this spe-
cies is known from the Oligocene of Mississippi
(Mint Spring Formation, Byram Formation, and
Chickasawhay Limestone) and Louisiana (Rosefield
Formation), USA. Nolf (2003) postulated that pres-
ent-day freshwater genera (for example, Aplodinotus
grunniens Rafinesque, 1819 of the Mississippi River
drainage system in the USA) are descendants of ma-
rine ancestors like A. gemma and related species from
the USA and Central and South America. Juvenile
22 JUN A. EBERSOLE ET AL.
specimens of this species are more difficult to iden-
tify because of their plesiomorphic morphology. The
similarity of A. gemma sagittae to Sciaena pseudora-
dians, especially among smaller specimens (< 6 mm),
is outlined above. The ontogenetic changes observed
in A. gemma include the curving of the distal portion
of the cauda toward the posteroventral margin of the
ostium. This feature is highly characteristic and not
observed on S. pseudoradians (Nolf 2013, pl. 269, top
specimen of A. gemma) otoliths. The curving of the
distal part of the cauda toward the ostium is not seen
on the specimen of A. gemma from site AWa-9 due to
its small size (approximately 5 mm).
Genus Sciaenops Gill, 1863a
Sciaenops? sp.
(Text- f ig . 8K)
MATER IAL: SC2019.61.43 (tooth).
DESCRIPTION: Specimen SC2019.61.43 is a high-
crowned tooth, with an overall height (1.2 mm) that
exceeds its occlusal diameter (0.7 mm). The tooth
has smooth crown enameloid and a circular occlusal
outline. In profile view the edges of the crown are
straight and gently tapered apically, and the occlusal
surface is evenly convex. The enameloid covers the
upper two-thirds of the tooth, whereas the lower third
is comprised of exposed dentine. The upper half of
the enameloid crown is a darker in color than the
lower half. The edges of the dentine base are parallel
and straight, and slightly inset from the crown base.
The tooth base is cylindrical and has a wide, deep,
and medially positioned pulp cavity.
REMARKS: SC2019.61.43 appears to be congeneric
with the pharyngeal teeth of a Recent Sciaenops ocel-
latus (Lin næus, 1766) specimen i n the MSC collection
(MSC 42611). The pharyngeal bones of MSC 42611
exhibit two distinct tooth morphologies, including
those that are recurved with a pointed apex and those
that are erect with a blunt and evenly convex apex.
Specimen SC2019.61.43 compares very favorably to
the latter morphology. The specimen differs from
other teeth in our Glendon Limestone sample by be-
ing high-crowned and having a more convex apex
and deeper pulp cavity. Although SC2019.61.43 is
similar in size and proportion to teeth of Albula eppsi
that have been reported from the Eocene of Alabama
(see Ebersole et al. 2019, fig. 60a–c), teeth of that
taxon differ by having evenly convex lateral edges
and crown enameloid that extends almost to the base
of the tooth.
Near et al. (2013) provided molecular evidence in-
dicating that S. ocellatus and Micropogonias undula-
tus (Linnæus, 1766) diverged from a sciaenid ances-
tor sometime during the Oligocene. Lo et al. (2015)
further postulated that S. ocellatus originated in the
western Atlantic Ocean during the early Miocene,
roughly 15.8 Ma. Specimen SC2019.61.43 is therefore
conservatively assigned to Sciaenops? sp. because it
is unclear if the tooth represents a basal representa-
tive of the genus or an altogether different and unde-
scribed Oligocene sciaenid within the Sciaenops and
Micropogonias lineage.
Order Spariformes (sensu Nelson, Grande and
Wilson, 2016)
Family Sparidae Rafinesque, 1818
Genus Sparus Linnæus, 1758
Sparus? elegantulus (Koken, 1888)
(Text- f ig . 8L)
MATERIAL: MSC 43054.4 (sagitta), MSC 43054.7
(sagitta), MSC 43059.7 (sagitta), MSC 43067.2 (sagitt a),
MSC 43067.3 (sagitta), MSC 43067.4 (sagitta), MSC
43067.5 (sagitta), SC2019.61.5 (sagitta), SC2019.61.6
(sagitta), SC2019.61.7 (sagitta), SC2019.61.8 (sa-
gitta), SC2019.61.14 (sagitta), SC2019.61.15 (sagitta),
SC2019.61.24 (sagitta).
DESCRIPTION: The sagittae are generally small,
with most ranging from 3 to 4 mm in length. The sa-
gitta outline is oval (sensu Smale et al. 1995), and the
height/length ratios range from 0.64–0.70. The inner
face is slightly convex and smooth, and the margins
are variable in shape, with irregular lobes common.
The anterior margin is characterized by the rostrum
and antirostrum, and the anterodorsal margin is con-
vex and often irregular. The dorsal margin is convex
and usually irregular, and the posterodorsal margin is
convex and steeper than the anterodorsal margin. The
posterior margin is tapered and may be somewhat
pointed, and the ventral margin is broadly rounded. A
prominent sulcus (heterosulcoid type) extends across
85% of the inner face, and the ostium is about one-
half the length of the cauda. The height of the cauda
is less than the height of the ostium. A slight ventral
expansion of the ostium is filled with colliculum. The
anterior portion of the ostium extends onto the ros-
trum, and the ventral margin of the ostium is essen-
tially horizontal. The cauda is elongated, with a long
FISHES FROM THE OLIGOCENE OF ALABAMA 23
horizontal portion and a short downturned portion.
The angle of the horizontal and downturned portions
is approximately 45º, and the downturned portion is
slightly tapered and significantly separated from the
posteroventral margin. An irregular depressed area
occurs above the sulcus, largely above the cauda.
A ventral furrow is present, and the outer face is
slightly concave.
REMARKS: This species was first reported from
the Gulf Coast of the USA by Koken (1888), and it
has since been documented from the upper Eocene
Yazoo Clay of Louisiana (as “genus Sparidarum” el-
egantulus) where it was reported as common (Nolf
and Stringer 2003). The assignment of the taxon to
Sparus? is based on Janssen (2012), who recom-
mended the use of the type genus for the family fol-
lowed by a question mark in the event the family and
species is known, but the genus is unknown. This
convention indicates that the species belongs to a
known or possibly undescribed genus in the family.
Order Tetraodontiformes Berg, 1940
Suborder Balistoidei Rafinesque, 1810
Family Balistidae Rafinesque, 1810
Gen. et sp. indet.
(Text- f ig . 8M )
MATERIAL: SC2019.61.38 (worn tooth).
DESCRIPTION: This tooth measures just over 1 mm
in length and width. The labial two-thirds of the oc-
clusal surface of the tooth is represented by a large
and flat wear facet, and the anterior portion of the
main cusp is not preserved. The lingual one-third of
the crown is lingually projected and strongly convex.
The labial portion of the tooth is much wider than the
lingual portion, giving the tooth a tear-drop shaped
occlusal outline. The mesial, distal, labial, and lin-
gual edges are rounded. The crown enameloid does
not extend to the base of the tooth, revealing a basal
concentric ring of exposed dentine. The basal edge of
the tooth is abraded and irregular.
REMARKS: Specimen SC2019.61.38 appears to
be comparable to teeth assigned to the Balistidae
that were recovered from middle Eocene depos-
its in Alabama (see Ebersole et al. 2019, fig. 67).
The combination of tear-drop shaped occlusal out-
line and rounded lateral edges is unique among
the Glendon Limestone Member teeth described
herein, and separates this taxon from other similar
Paleogene taxa like Eotrigonodon Weiler, 1929 (see
Ciobanu 2011, figs 5–10). The morphology is consis-
tent with extant balistid incisiform teeth, but a more
refined identification is currently not possible due to
the incomplete preservation of the single specimen
available. The only described Paleogene balistid is
the middle Eocene taxon, Gornylistes prodigiosus
Bannikov and Tyler, 2008, a species known only
from a complete skeleton. However, as the dentition
of that species has not been described and no mean-
ingful comparisons can therefore be made with the
Glendon Limestone tooth, specimen SC2019.61.38
is assigned only to the Balistidae. Although simi-
lar teeth occur in the Bartonian Gosport Sand, and
they are relatively common in the Priabonian Yazoo
Clay in Alabama (JAE, unpublished data) and the
Parkers Ferry Formation of South Carolina (DJC,
unpublished data), specimen SC2019.61.38 extends
the temporal range of this morphology into the
Rupelian.
Teleostei indet.
MATERIAL: MSC 43054.9 (sagitta), MSC 43059.8
(sagitta), SC2019.61.16 (sagitta), SC2019.61.29 (sa-
gitta), SC2019.61.39 (tooth), SC2019.61.40 (tooth),
SC2019.61.41 (tooth), SC2019.61.42 (tooth), SC2019.
61.47 (worn tooth), SC2019.61.48 (dorsal fin spine),
SC2019.61.49 (dorsal fin spine), SC2019.61.50 (dorsal
fin spine), SC2019.61.51 (dorsal fin spine), SC2019.
61.52 (dorsal fin spine), SC2019.61.53 (dorsal fin
spine), SC2019.61.54 (dorsal fin spine), SC2019.61.55
(2 fin spine fragments), SC2019.61.56 (fin spine),
SC2019.61.57 (fin spine), SC2019.61.58 (fin spine),
SC2019.61.59 (fin spine), SC2019.61.60 (left quad-
rate), SC2019.61.61 (left quadrate), SC2019.61.62
(right premaxilla), SC2019.61.63 (left dentary),
SC2019.61.64 (dentary?), SC2019.61.65 (jaw frag-
ment), SC2019.61.66 (atlas vertebra), SC2019.61.67
(atlas vertebra), SC2019.61.68 (vertebra), SC2019.61.
69 (atlas vertebra), SC2019.61.70 (vertebra), SC2019.
61.71 (vert ebra), SC2019.61.72 (vertebra), SC2019.61.
73 (vertebra), SC2019.61.74 (vertebra), SC2019.61.75
(vertebra), SC2019.61.76 (ver tebra), SC2019.61.77
(vertebra), SC2019.61.78 (ver tebra), SC2019.61.79
(vertebra), SC2019.61.80 (vertebra), SC2019.61.81
(vertebra), SC2019.61.82 (vertebra), SC2019.61.83
(vertebra), SC2019.61.84 (bone fragment), SC2019.
61.85 (bone fragment), SC2019.61.86 (bone frag-
ment), SC2019.61.87 (bone fragment), SC2019.61.88
(bone fragment), SC2019.61.89 (bone fragment),
SC2019.61.90 (epihyal?), SC2019.61.91 (bone frag-
24 JUN A. EBERSOLE ET AL.
ment), SC2019.61.92 (epihyal?), SC2019.61.93 (bone
fragment), SC2019.61.94 (bone fragment), SC2019.
61. 95 (bone fragment) SC2019.61.96 (bone fragment),
SC2019.61.97 (preoperculum fragment), SC2019.61.98
(bone fragment), SC2019.61.99 (cranial? element),
SC2019.61.100 (bone fragment), SC2019.61.101 (bone
fragment), SC2019.61.102 (scale), SC2019.61.103
(scale), SC2019.61.104 (scale), SC2019.61.105 (scale),
SC2019.61.107 (bone fragment), SC2019.61.168
(2 scales), SC2019.61.169 (scale), SC2019.61.170 (scale).
REMARKS: Our Glendon Limestone sample con-
tains a plethora of micro-teleost remains that cannot
be identified, including ablated sagittae and teeth,
dorsal and pectoral fin spines, vertebral centra, scales,
miscellaneous cranial and jaw elements, and unidenti-
fied bone fragments. The otoliths listed in this section
are too poorly preserved to be assigned to any lower
taxonomic ranking, but they likely belong to one of
the taxa described herein. A majority of the teeth are
represented by broken or incomplete tooth caps that
are too worn or abraded for further identification. One
well preserved tooth, SC2019.61.39, differs by being
slightly recurved and cone-shaped. Unfortunately the
lack of adequate comparative specimens did not allow
us to further identify this specimen.
Three morphologies of dorsal and pectoral fin
spines are present in our sample. These include: 1)
spines that are ornamented with fine or coarse lateral
longitudinal ridges, but with no anterior or posterior
denticulation; 2) smooth spines lacking anterior and/
or posterior denticulation; and 3) a single spine or-
namented with coarse lateral ridges and two rows
of posterior denticles, but no denticles occur on the
anterior margin. As the denticulation and lateral or-
nament on this last spine morphology (SC2019.61.56)
is reminiscent of spines occurring on members of the
Ariidae (sea catfishes), the specimen was directly
compared to those of Recent Ariopsis felis (Linnæus,
1766) and Bagre marinus (Mitchill, 1815) (MSC
43216 and MSC 43217, respectively). Although the
ridged ornamentation is similar between all three
spine morphologies, SC2019.61.56 lacks anterior den-
ticulation but has two rows of posterior denticles that
flank a posterior furrow. This morphology differs
from the dorsal and pectoral spines of A. felis and B.
marinus, which exhibit a single row of both anterior
and posterior denticles, with those at the posterior re-
siding within a longitudinal furrow. The other orna-
mented and unornamented spine morphologies in the
Glendon Limestone sample share characteristics with
a variety of Recent bony fishes, and all are therefore
left in open nomenclature.
The remaining elements, including vertebrae,
scales, cranial elements, and bone fragments, are also
left in open nomenclature. Unfortunately the lack
of available comparative material of small marine
fishes and the paucity of prior studies on Oligocene
micro-teleosts have not allowed us to further identify
these specimens.
DISCUSSION
There is evidence that the vertebrate and inver-
tebrate paleofaunas within the Glendon Limestone
Member in Alabama and the Glendon Limestone (for-
mation) in Mississippi vary appreciably with regard
to paleoecological preferences (Mumma 1965; Fisher
and Ward 1984; Huddlestun 1993; Fluegeman et al.
2019; Stringer and Starnes 2020), and this probably
also occurs at site AWa-9. Only a single lithostrati-
graphic horizon at the site was investigated, and the
paleoecological interpretations provided herein are
therefore limited to the sampled interval at the local-
ity from which the vertebrate remains were recov-
ered, and may not apply to the sub- and superjacent
strata. The species diversity of the Glendon Limestone
Member assemblage was evaluated based on all of
the recovered vertebrates (teeth, skeletal remains, oto-
liths) that could be identified to at least the family
level. Regarding species diversity, both the number
of species (diversity or richness) and the percentage
of each species (relative abundance) were considered.
Twenty unequivocal taxa were identified in the as-
semblage, including six cartilaginous fishes and four-
teen bony fishes (nine based on otoliths and five based
on osteological remains). The assemblage is strongly
skewed in the percentage each taxon comprises, with
one bony fish, Citharichthys, constituting over 49%
of the total number of identifiable otolith specimens.
Furthermore, three taxa of bony fishes, Citharichthys,
Sparus? elegantulus, and Preophidion meyeri, account
for over 60% of the identifiable otolith assemblage.
The remains of cartilaginous fishes are very lim-
ited within the Glendon Limestone Member, both in
number of specimens (n=8) and species (n=6). This
greatly influences their contribution to an under-
standing of the paleoenvironment. Likewise, a large
percentage of the osteological remains of the bony
fishes were not identifiable beyond anatomical ele-
ment, which greatly inhibits their application in pa-
leoenvironmental reconstruction. Despite these lim-
itations, we were able to glean paleoenvironmental
data based on the otoliths and osteological remains
we could identify.
FISHES FROM THE OLIGOCENE OF ALABAMA 25
The families of fishes represented by otoliths were
utilized to ascertain the general paleoenvironmental
parameters represented by the Glendon Limestone
Member assemblage, following methods that have
been effectively applied to similar assemblages in
both the Gulf and Atlantic coastal plains (Stringer
and Bell 2018; Ebersole et al. 2019; Stringer and
Shannon 2019; Stringer and Hulbert 2020; Stringer
et al. 2020b). This was accomplished by compar-
ing the ecological ranges of extant families of fishes
represented by the fossil otoliths (Cohen et al. 1990;
Hoese and Moore 1998; Nelson et al. 2016; Snyder
and Burgess 2016; Froese and Pauly 2019). We in-
clude here the caveat that there are limitations and
considerations inherent to this application of biolog-
ical uniformitarianism, which are compounded in
temporally much older assemblages, especially in the
Mesozoic (Schwarzhans et al. 2018; Stringer et al.
2018, 2020c; Schwarzhans and Stringer 2020).
As is shown in Table 1, three of the families
identified by otoliths are restricted to marine wa-
ters (Congridae, Diretmidae, and Ophidiidae), and
these comprise about 17% of the total specimens.
Additionally, there are four families that are known to
inhabit fresh, brackish, and marine waters. However,
there are factors to consider regarding these four fam-
ilies, as the Gobiidae are chiefly brackish and marine,
whereas the Paralichthyidae and Sparidae are chiefly
marine but rare in freshwater, and very rare in fresh
and brackish waters (respectively). The only family
equally abundant in freshwater, brackish, and ma-
rine environments, the Sciaenidae, comprises only
approximately 6% of the total otolith sample. There
are no families present in the Glendon Limestone
Taxa identified from otoliths Quantity % of total Ecology Climate
ANGUILLIFORMES
Congridae M Trop. – temp.
Ariosoma nonsector 6 5.17
TRACHICHTHYIFORMES
Diretmidae M Trop. – temp.
Diretmus? sp. 1 0.86
OPHIDIIFORMES
Ophidiidae M Trop. – temp.
Preophidion meyeri 12 10.34
GOBIIFORMES
Gobiidae F. B. M1Trop. – subtrop.
Gobiasoma? axsmithi sp. nov. 4 3.45
PLEURONECTIFORMES
Paralichthyidae F. B. M2Trop. – temp.
Syacium sp. 3 2.58
Citharichthyes sp. 57 49.13
Citharichthyes? sp. (lapilli) 2 1.72
Paralichthyidae indet. 1 0.86
ACANTHURIFORMES
Sciaenidae F. B. M Trop. – temp.
Sciaena pseudoradians 4 3.45
Aplodinotus gemma 3 2.58
Sciaenidae indet. 1 0.86
SPARIFORMES
Sparidae F. B. M3Trop. – temp.
Sparus? elegantulus 14 12.06
ORDER INDET.
Family indet.
Teleostei indet. 8 6.89
Total 116 100
Table 1. Taxa represented by otoliths from the Glendon Limestone Member of the Byram Formation (Oligocene, Rupelian), site AWa-9, Wash-
ington County, Alabama, USA. Ecologic and climatic information based on families is represented. Abbreviations: B – brackish; F – freshwater;
M – marine; Temp. – temperate; Subtrop. – subtropical; Trop. – tropical. Ecology data superscripts as follows: 1 – chiefly marine and brackish
water; 2 – chiefly marine and rare in fresh water; 3 – chiefly marine and very rare in fresh- and brackish water. Ecologic and climatic data derived
from Hoese and Moore (1998), Nelson et al. (2016), Froese and Pauly (2019), and World Register of Marine Species Editorial Board (2020).
26 JUN A. EBERSOLE ET AL.
Member otolith assemblage that represent exclusively
freshwater or brackish environments. As the otolith
assemblage is largely comprised of families that have
marine representatives, it is likely that the Glendon
Limestone Member represents marine deposition,
with little evidence of brackish or freshwater influ-
ence (i.e., probably not very close to shore).
The Glendon Limestone Member otoliths also
furnish information regarding paleo-water depths. Of
the specimens we identified, none represent families
that are indicative of deep water (i.e., greater than 200
m). Nolf and Brzobohaty (1994) stated that marine as-
semblages that lack or contain very few Myctophidae
(lanternfish) likely indicate a neritic environment
with little open oceanic inf luence. Although mycto-
phids are known from as early as the Eocene (Nolf
2013), none are present in our Glendon Limestone
Member assemblage. The assemblage also does not
contain any Macrouridae (grenadiers), a family of
typically bathybenthic fishes living below 200 m,
thus providing strong evidence for water depths less
than 200 m (i.e., outer shelf or less).
An analysis of the most abundant otolith taxa in
our sample offers additional information on paleoen-
vironmental conditions. At a minimum, the data pre-
sented above indicates that the Glendon Limestone
Member at site AWa-9 represents a marine inner to
outer shelf-depth paleoenvironment. Citharichthys is
by far the most abundant bony fish in our sample,
representing nearly 50% of the total number of iden-
tified specimens. This high percentage would there-
fore indicate that the paleoecological conditions at
the site must have been optimum for the survival and
proliferation of Citharichthys. Page et al. (2013) re-
ported six species of Citharichthys in the present-day
Atlantic Ocean and Gulf of Mexico of the USA, and
according to Froese and Pauly (2019) many of these
species occur at relatively shallow depths, although
some occur in much deeper water (i.e., Citharichthys
dinoceros Goode and Bean, 1886 as deep as 2000
m). Three Citharichthys species are not found at
depths of less than approximately 35 m, but two spe-
cies occur in shallow coastal waters (sounds, bays,
lagoons). As there is no evidence that the Glendon
Limestone Member paleowater depth was 200 m or
more, the deep-water Citharichthys are not consid-
ered. Additionally, the presence of very shallow-wa-
ter species (Carpenter et al. 2015) is not supported
by the balance of the Glendon Limestone Member
assemblage. Thus, the remaining Citharichthys spe-
cies would support a paleowater depth of no less than
30 m (i.e., shallow middle shelf). Of course, it must
be considered that the preceding paleowater depth
determination is based on the present distribution of
Citharichthys species in the modern Gulf of Mexico,
and could be affected or altered by various factors.
However, when taken in conjunction with other lines
of evidence, it appears to be feasible.
Sparus? elegantulus is the second most abundant
species in the Glendon Limestone Member assem-
blage, constituting 12.06% of the total identified spec-
imens. Janssen (2012) recommended using the type
genus for the family followed by a question mark in
the event that the family and species known, but the
genus is unknown. This convention indicates that the
species could belong to a known or possibly unde-
scribed genus in that family. Therefore, for the spec-
imens identified as Sparus? elegantulus we only uti-
lize ecological information as it applies to the familial
level. Members of the Sparidae can be found in fresh,
brackish, and marine waters (Froese and Pauly 2019),
although Nelson et al. (2016) noted that this group
rarely occurs in freshwater or brackish environments.
Sparids are demersal inhabitants of the continental
shelf and slope, but usually most common along the
shore, from shallow to deeper water (Iwatsuki and
Heemstra 2015). Therefore, the presence of Sparus?
elegantulus indicates a marine environment, but this
taxon is not a useful paleoenvironmental indicator.
The third most abundant species in the otolith
assemblage is the ophidiid (cusk-eel) Preophidion
meyeri, which represents 10.34% of the assemblage.
Preophidion meyeri is a fossil otolith-based genus
and species, and its relationship to modern ophidiid
taxa is unclear. However, extant ophidiids only occur
in marine waters (Nelson et al. 2016; Froese and Pauly
2019), and these fish are therefore a good indication
that the Glendon Limestone Member paleoenviron-
ment was not freshwater or brackish. Nolf (2013) as-
signed P. meyeri to the subfamily Neobythitinae, and
according to Nelson et al. (2016), representatives of
this subfamily range from littoral to abyssal (> 2000
m). However, during the Paleogene the ophidiids
were represented by a very rich neritic fauna living
mainly on soft and muddy substrates, and these are
some of the most common and most speciose groups
of teleosts recovered from shallow-marine deposits
(Nolf 1985, 2013; Stringer 1986; Breard and Stringer
1995; Stringer and Miller 2001; Nolf and Stringer
2003; Schwarzhans and Bratishko 2011).
Two Sciaenidae taxa represent roughly 6% of the
total number of otolith specimens in our Glendon
Limestone Member sample. Although Recent sciae-
nids are found in fresh, brackish, and marine waters,
it is unlikely that the Glendon Limestone sciaenids
represent freshwater forms because, as Nolf (2003)
FISHES FROM THE OLIGOCENE OF ALABAMA 27
has noted, all Paleogene sciaenids known from the
USA Gulf Coast deposits are associated with marine
assemblages. Although some taxa may have been
derived from strata representing nearshore paleoen-
vironments, none can be qualified as freshwater or
even lagoonal. Although many modern sciaenids uti-
lize estuaries as nurseries (Barbieri 1993; Barbieri
et al. 1994; Pattillo et al. 1997; Snyder and Burgess
2016), the otoliths are usually very small (i.e., larval
and juveniles) and abundant (Stringer and Shannon
2019, and references therein). The Glendon Limestone
sciaenids do not meet either of these criteria, as the
size of these otoliths indicate they all represent adult
individuals, and very few specimens were recovered
in our sample. Based on these observations, there are
no indications that the Glendon Limestone sciaenids
were living in freshwater, brackish water, or estuaries.
In addition to Citharichthys, a second paralich-
thyid, Syacium, was recovered during our investiga-
tion. Although only two specimens were identified,
the presence of Syacium offers particular insight
into the Glendon Limestone Member paleoenviron-
ment. According to Page et al. (2013) and Froese
and Pauly (2019), only three species of Syacium are
currently found in the Atlantic Ocean and Gulf of
Mexico [S. gunteri Ginsburg, 1933, S. micrurum
Ranzani, 1842, and S. papillosum (Linnæus, 1758)],
and these taxa usually occur in waters ranging from
27–95 m, 0–100 m, and 27–95 m in depth, respec-
tively. Therefore given the modern distribution of
these extant species, the presence of Syacium in the
Glendon Limestone Member could reflect a paleowa-
ter depth of up to 100 m.
The families of fishes represented by otoliths
also provide a general indication of climatic condi-
tions. Six of the seven families in our sample are
found in tropical to temperate waters, including the
Paralichthyidae (which are represented by the largest
number of specimens). The genus Gobiosoma of the
Gobiidae, represented by four specimens (3.45% of
the total sample), currently inhabits tropical to sub-
tropical waters of the Americas. Thus, the majority
of the otolith taxa we identified are indicative of a
climatic setting that was tropical to temperate, based
on the modern distribution of fish families (Hoese
and Moore 1998; McEachran and Fechhelm 1998,
2005; Nelson et al. 2016; Froese and Pauly 2019;
World Register of Marine Species Editorial Board
2020). Most of the otolith taxa in our sample repre-
sent fishes that preferred soft substrates like sand or
mud. Furthermore, only one specimen (Gobiosoma)
was recovered that exhibited invertebrate settlement
indications (a very small boring in the dorsal depres-
sion, Text-fig. 7B), which could be an indication of
very little surface residence time.
In addition to the otoliths, several of the chon-
drichthyan fossils in our sample provide clues to
the Glendon Limestone Member paleoenvironment
(Table 2). For example, Hemipristis elongata (Klun-
zinger, 1871), a modern analogue for Hemi pris-
tis sp., is a tropical marine taxon that prefers water
depths of between 1–130 m (Froese and Pauly 2019).
Interestingly, Müller (1999) indicated that H. serra
was common in warm water during the Neogene,
but during the Rupelian in Europe the absence of
Hemipristis was taken to be related to the colder wa-
ter conditions that existed during that time (von der
Hocht 1978). Furthermore, Negaprion brevirostris
(Poey, 1868), an extant analogue for the extinct N.
gilmorei, is a marine/brackish subtropical taxon with
depth preferences of between 1–92 m (Froese and
Pauly 2019). The depth ranges for both of these gen-
era are well within the 30–100 m Glendon Limestone
Member paleowater depth as indicated by the oto-
lith taxa, and they also corroborate the tropical/sub-
tropical to temperate conditions indicated. Another
elasmobranch in our sample, Pachyscyllium sp., is
an extinct genus within Scyliorhinidae (catsharks).
According to Collareta et al. (2020), extant scyliorh-
inids inhabit moderately deep waters in tropical
to temperature seas worldwide. Yet another taxon,
Physogaleus sp., is an extinct requiem shark in the
Carcharhinidae, one of the largest families of ex-
tant sharks that includes both coastal and offshore
taxa that inhabit tropical to temperate marine waters
(Castro 1983; Villafaña et al. 2020). Although these
latter two extinct taxa contribute little to our under-
standing of the paleowater depth, they do indicate
tropical/subtropical to temperate climatic conditions.
Two teeth assigned to “Aetomylaeus” sp. were iden-
tified in our Glendon Limestone sample. Froese and
Pauly (2019) recognized seven extant members of
this genus, with all but one of them preferring tropi-
cal waters and depths of 117 m or less. The only ex-
ception is Aetomylaeus bovinus (Geoffroy St. Hilaire,
1817), a subtropical taxon with water depth prefer-
ences of 150 m or less (Froese and Pauly 2019). Thus,
the presence of “Aetomylaeus” sp. in the Glendon
assemblage suggests the maximum paleowater depth
likely did not exceed 117 m.
Several of the bony fishes identified from oste-
ological remains provide some paleoecological data
on the Glendon Limestone Member (Table 2). For
example, a single tooth assigned to Albula sp. was
identified in our sample. Extant members of the fam-
ily primarily inhabit marine waters but are extremely
28 JUN A. EBERSOLE ET AL.
rare in fresh or brackish environments (Froese and
Pauly 2019), and they primarily occur in the tropics
(Nelson et al. 2016). We also identified Sphyraena
(barracuda) teeth in our sample. Extant representa-
tives of the genus generally inhabit tropical to sub-
tropical marine waters and prefer water depths of
between 3–30 m, although some species have been
reported as deep as 100 m (Nelson et al. 2016; Snyder
and Burgess 2016; Froese and Pauly 2019). A sin-
gle Sciaenops? tooth in our sample is comparable to
the Recent Sciaenops ocellatus, a species in which
juveniles are found in bay shores, open waters of es-
tuaries, and secondary bays in depths up to 3 m, but
adults can occur in nearshore waters off beaches and
at depths as great as 40–70 m (Pattillo et al. 1997 and
references cited therein). The single Balistidae tooth
indicates a marine environment (Froese and Pauly
2019), and representatives of the family are found
primarily on the shelf, although there are oceanic
species (Snyder and Burgess 2016).
Comparison of the paleoenvironmental parame-
ters based on the osteological remains (cartilaginous
and bony fishes) to those indicated by the teleostean
otoliths reveal interesting similarities. The paleowa-
ter depths indicated by the osteological remains of
the sharks and bony fishes corroborates the paleowa-
ter depths determined based on the otoliths. There are
no taxa based on skeletal remains that are completely
outside the range indicated by the taxa identified by
otoliths. Furthermore, the taxa represented by osteo-
Taxa identifed from osteological remains Quantity % of total Ecology Climate
CARCHARHINIFORMES
Scyliorhinidae M Trop. – temp.
Pachyscyllium sp. 1 1.15
Hemigaleidae M Trop. – temp.
Hemipristis sp. 1 1.15
Carcharhinidae F, B, M Trop. – temp.
Negaprion aff. N. gilmorei 2 2.29
Physogaleus sp. 1 1.15
MYLIOBATIFORMES
Myliobatoidei – –
Gen. et sp. indet. 1 1.15
Myliobatidae F, B Trop. – temp.
“Aetomylaeus” sp. 2 2.29
ELOPIFORMES
Phyllodontidae – –
Paralbula sp. 2 2.29
ALBULIFORMES
Albulidae F, B, M1Tropical
Albula sp. 1 1.15
ISTIOPHORIFORMES
Sphyraenidae M Trop. – subtrop.
Sphyraena sp. 3 3.44
ACANTHURIFORMES
Sciaenidae F, B, M Trop. – temp.
Sciaenops? sp. 1 1.15
TETRADONTIFORMES
Balistidae M Trop. – subtrop.
Gen. et sp. indet. 1 1.15
ORDER INDET.
Family indet. – –
Elasmobranchii indet. 2 2.29
Teleostei indet. 69 79.31
Total 87 100
Table 2. Taxa represented by osteological remains from the Glendon Limestone Member of the Byram Formation (Oligocene, Rupelian), site
AWa-9, Washington County, Alabama, USA. Ecologic and climatic information based on families is represented. Abbreviations: B – brackish;
F – freshwater; M – marine; Temp. – temperate; Subtrop. – subtropical; Trop. – tropical. Ecology data superscripts as follows: 1 – chiefly marine
and very rare in fresh- and brackish water. Ecologic and climatic data derived from Hoese and Moore (1998), Nelson et al. (2016), Froese and
Pauly (2019), Collareta et al. (2020), and World Register of Marine Species Editorial Board (2020).
FISHES FROM THE OLIGOCENE OF ALABAMA 29
logical remains reflect the same climatic conditions
as the taxa indicated by the otoliths, and osteological
remains do not contradict any of the paleoecologi-
cal assertions based on the otoliths. In summary, the
skeletal remains of the cartilaginous and bony fishes
corroborate in a general sense (i.e., less detailed but
still supportive) the paleoenvironment as evidenced
by the bony fish otoliths, which agrees with the find-
ings of Miller (2000) and Stringer and Miller (2001).
CONCLUSIONS
Bulk samples collected from the Glendon Lime-
stone Member exposures at site AWa-9 in Washington
County, Alabama, USA yielded an u nexpectedly large
number and diverse assortment of marine vertebrate
remains, and 20 unequivocal elasmobranch (n=6) and
teleost (n=14) taxa were identified. Each of these taxa
represent the first occurrence within the Oligocene
(Rupelian) Glendon Limestone Member in Alabama,
USA, and one new species, Gobiosoma? axsmithi
sp. nov., is recognized. Several other taxa, including
“Aetomylaeus” sp., Pachyscyllium sp., Paralbula sp.,
and Sciaenops? sp., represent the first occurrences of
each in the Oligocene of the Gulf Coastal Plain of the
USA. Furthermore, the Glendon Paralbula teeth rep-
resent a slight range extension for this genus from the
Priabonian (upper Eocene) into the Rupelian (lower
Oligocene). Finally the Balistidae indet. tooth in our
sample represents the first Oligocene occurrence of
this family in the Western Hemisphere.
The compilation of the environmental data ob-
tained from the vertebrate assemblage in the Glendon
Limestone Member at site AWa-9 indicates that pa-
leowater depth was at least 30 m (shallow middle
shelf). This is shallower than the minimum depth
for the Glendon Limestone of Mississippi postulated
by Mumma (1965), which was 75 m (deep middle
shelf). However, this minimum depth is very similar
to the deep inner shelf (approximately 20 m) depth
indicated by foraminifera (Fisher and Ward 1984)
from the Glendon Limestone in Warren County,
Mississippi. The maximum depth is more difficult
to ascertain, but there is evidence that the maximum
water depth did not exceed 200 m (outer shelf) based
on the presence and absence of diagnostic fish spe-
cies. Although not a strong indicator, the presence of
“Aetomylaeus” and Syacium could indicate a max-
imum paleowater depth of 100 m and tropical/sub-
tropical to temperate climatic conditions. The fishes
represented by otoliths attest to soft substrate of mud,
sand, or a mixture of the two. The lack of invertebrate
settlement on the otolith specimens indicates short
surface residence-time (fairly rapid burial).
As Oligocene units in Alabama are highly fos-
siliferous but historically understudied, future ex-
amination of these strata will undoubtedly yield ad-
ditional vertebrate taxa. Thus, the need for further
investigations in the Alabama Oligocene is certainly
warranted and encouraged.
Acknowledgements
We graciously thank Jennifer Faith, Director of St. Stephens
Historical Park (Washington County, AL, USA), for granting
us access and permission to collect samples from the site.
James Starnes of the Mississippi Office of Geology (Jackson,
USA) is thanked for assisting JAE with reidentifying the
lithologic units mapped by Glawe (1967) at site AWa-9 and
for his helpful discussions regarding the Glendon Limestone
Formation in Mississippi. Drew Gentry of McWane Science
Center (Birmingham, AL, USA) is thanked for his assistance
with collecting bulk samples from the locality. K.A. Johnson
(National Marine Fisheries Service, Southeast Fisheries
Science Center, Pascagoula, MS, USA), R. Taylor (formerly of
the Florida Fish and Wildlife Conservation Commission, Fish
and Wildlife Research Institute, St. Petersburg, FL, USA), J.R.
Hendon (Center for Fisheries Research and Development, Gulf
Coast Research Laboratory, University of Southern Mississippi,
Ocean Springs, MS, USA), and D. Nolf (Institut Royal des
Sciences Naturelles de Belgique, Brussels, Belgium) gener-
ously provided Recent fishes and otoliths. W. Schwarzhans
(Natural History Museum of Denmark, Zoological Museum,
Copenhagen, Denmark) made valuable suggestions regarding
the identity and taxonomy of otoliths, especially the gobiids and
paralichthyids. F.H. Mollen (Elasmobranch Research, Belgium)
and W. Schwarzhans are thanked for their insightful review
comments on an earlier version of this work. Finally, we thank
handling editor Anna Żylińska for her editorial comments and
assistance throughout the publication of this study.
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Manuscript submitted: 25th November 2020
Revised version accepted: 29th January 2021
