Historical Biology

An International Journal of Paleobiology

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A late Miocene (Hemphillian) freshwater fish

(Osteichthyes) fauna from Mobile County,
Alabama, USA

Jun A. Ebersole & Stephen J. Jacquemin

To cite this article: Jun A. Ebersole & Stephen J. Jacquemin (2018): A late Miocene (Hemphillian)
freshwater fish (Osteichthyes) fauna from Mobile County, Alabama, USA, Historical Biology, DOI:
10.1080/08912963.2018.1530995

To link to this article: https://doi.org/10.1080/08912963.2018.1530995

Published online: 09 Oct 2018.

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HISTORICAL BIOLOGY
https://doi.org/10.1080/08912963.2018.1530995

ARTICLE

A late Miocene (Hemphillian) freshwater fish (Osteichthyes) fauna from Mobile

County, Alabama, USA
Jun A. Ebersolea and Stephen J. Jacqueminb
a
Collections Department, McWane Science Center, Birmingham, AL, USA; bDepartment of Biological Sciences, Wright State University - Lake
Campus, Celina, OH, USA

ABSTRACT ARTICLE HISTORY

Systematic descriptions of North American Miocene freshwater fishes are extremely uncommon, with Received 23 July 2018
those from the Gulf Coastal Plain region being completely absent. In this study we examined 324 Accepted 28 September 2018
historically collected freshwater fish elements derived from the late Miocene (Hemphillian) Mauvilla KEYWORDS
fossil site located in Mobile County, Alabama, USA. Analysis of these elements confirmed the presence Mauvilla local fauna; Gulf
of three unequivocal taxa including Atractosteus sp. cf. A. spatula, Ictalurus spp., and Aplodinotus Coastal Plain; Ecor Rouge
grunninens. The preserved elements consisted of spines, pharyngeal bones, pharyngeal teeth, and Sand; Ictalurus; Aplodinotus;
scales, likely indicating a taphonomic bias towards large individuals with robust elements. The co- Atractosteus
occurrence of these three taxa indicate they were derived from a medium-to-large river fish assem-
blage, suggesting that one or more of the rivers that constitute the nearby Mobile Basin may have been
present in the region during the late Miocene. Molecular divergence estimates and morphological
similarity to modern representatives both support the presence of these extant taxa in the late Miocene,
and the examined remains represent some of the earliest fossil occurrences of these extant taxa. These
three taxa also represent the first biogeographic occurrence of each in the late Miocene of both
Alabama and the Gulf Coastal Plain of North America and helps to improve our understanding of
Miocene freshwater fish assemblages.

Introduction Taylor (1971). Later that year, Isphording and Lamb

Late Miocene (Hemphillian) vertebrate local faunas are
extremely uncommon along the Gulf Coastal Plain of
North America, and only one, the Mauvilla local fauna,
has been described from the northern Gulf of Mexico (see
Savage and Russell 1983; Figure 6–32; Hulbert and
Whitmore 2006; figure 1). The Mauvilla local fauna is
named for the town of Mauvilla in north Mobile County,
Alabama, USA, a municipality that is located 2.4 km from
the site where these vertebrate remains were derived
(Figure 1). The early history of the Mauvilla fossil site
was chronicled by Hulbert and Whitmore (2006), who
noted its discovery by amateur collectors James E. Davis,
Jr. and his father. In 1966, the Davis’ discovered a mam-
malian skull along a small creek that they took to the
University of South Alabama in Mobile County, AL, USA
for identification. Soon after, the specimen was shipped to
Frank C. Whitmore, Jr. at the United States National
Museum (USNM) in Washington, D.C., USA, who deter-
mined the specimen belonged to a mammal that was
Hemphillian in age. Being the only specimen of this age
from Alabama, and one of the few from the Gulf Coastal
Plain in North America, Whitmore recognised its impor-
tance and conducted a preliminary excavation of the site in
1967. Whitmore recovered a small number of vertebrate
remains during this 1967 excavation, one of which, a pro-
toceratid, Synthetoceras, was described by Patton and
(1971) provided the first preliminary list of vertebrate
taxa from the Mauvilla locality, which included fishes
(Scombroidei, mackerels, and related fish), reptiles
(Trionyx sp. and Crocodilia), and mammals (Hipparion
phosphorum, Nannippus sp. cf. N. lenticularis,
Pomatodelphis inaequalis, Teleoceras sp., Synthetoceras sp.
cf. S. tricornatus, and indeterminate camelid and cervid
remains). This preliminary list of mammalian remains
was later reiterated in Thurmond and Jones (1981).
Whitmore again visited the Mauvilla locality in November
and December of 1981 and recovered over 400 additional verte-
brate specimens for the USNM collections (Hulbert and
Whitmore 2006). In December of 1981, Whitmore was joined
on this expedition by personnel from the Red Mountain
Museum (RMM) in Birmingham, AL, USA, who collected a
small number of vertebrate remains from the site. In 1994, the
RMM transferred these latter specimens to the scientific collec-
tions at McWane Science Center (MSC) in Birmingham, AL,
USA, where they reside today. In October of 1988, James Dobie
of the Auburn University of Paleontology (AUMP), Auburn,
AL, USA, revisited the site and collected a small number of
vertebrate remains for the AUMP collections, including those
belonging to a horse, Protohippus gidleyi. This specimen was
later described and figured by MacFadden and Dobie (1998).
Savage and Russell (1983) included the Mauvilla fossil site in
their review of North American paleofaunas (but under the
name ‘Chickasaw Creek’), and Hulbert and Whitmore (2006)

CONTACT Jun A. Ebersole jebersole@mcwane.org McWane Science Center, Birmingham, AL, USA
© 2018 Informa UK Limited, trading as Taylor & Francis Group

Published online 09 Oct 2018

2 J. A. EBERSOLE AND S. J. JACQUEMIN

Figure 1. General location of the Mauvilla fossil locality, Mobile County, Alabama, USA, and aerial extent of Miocene undifferentiated deposits in Alabama. Scale
bars = 20 km.

later provided systematic descriptions of the mammalian speci- Geologic setting

mens recovered from this locality that were housed at the
USNM. This latter study described 15 distinct mammalian
taxa from the site, 12 of which either represented new species
or were new to the state. These authors also, for the first time,
referred to these mammals as belonging to the ‘Mauvilla local
fauna’. MacFadden and Dobie (1998) also mentioned the pre-
sence of fishes and turtles at the Mauvilla fossil site, the latter of
which were to be described by Dobie. Dobie, unfortunately,
passed away before completing this work. One of these speci-
mens, however, a trionychid turtle (referred to Apalone sp. cf. A.
spinifera), was later described and figured by Jasinski (2013).
Aside from the brief remarks by Isphording and Lamb
(1971) and Hulbert and Whitmore (2006) on the presence
of freshwater fishes at the Mauvilla locality, as of yet, none of
these specimens have been formally documented. Here we
rectify this absence by describing the late Miocene freshwater
fishes collected from the Mauvilla fossil site, in turn adding to
the list of confirmed vertebrate taxa that comprise the
Mauvilla local fauna.
The Mauvilla fossil locality is located along Chickasaw Creek
near the town of Mauvilla in northern Mobile County in
Alabama, USA (Figure 1). The vertebrate remains recovered
from this locality were derived from a 60 cm thick clay and
sand bed exposed at the site. This vertebrate lens was origin-
ally interpreted to be the basal component of the Citronelle
Formation with an age as late as the middle Pliocene (see
Isphording and Lamb 1971; Tedford and Hunter 1984). Later
investigations, however, showed that this fossil rich layer
disconformably underlies the Pliocene Citronelle Formation
and is part of an expansive unit in south Alabama that has
been mapped locally as ‘Miocene Undifferentiated’ (see
Raymond et al. 1988). Isphording (1977) divided the
Miocene surface exposures in south Alabama into two dis-
tinct, but informal, units, the marine and transitional marine,
middle Miocene, ‘Mobile Clay’, and the fluvial, late Miocene,
‘Ecor Rouge Sand’ (Figure 2).
The Ecor Rouge Sand is a dark gray, carbonaceous silty
clay and clayey sand layer that is composed of alternating

Figure 2. Neogene surface stratigraphy in Alabama. Abbreviations: (E) early; (Fm.) formation; (L) late; (M) middle); (mya) millions of years ago. North American Land
Mammal Ages after Barnosky et al. (2014).

beds of clay and sand (Isphording and Lamb 1971). At the
Mauvilla fossil locality, all the described vertebrate remains
were derived from the sand layers within this informal unit
(Hulbert and Whitmore 2006). The depositional setting for
the Ecor Rouge Sand is interpreted as being a deltaic flood-
plain, with the vertebrate material being deposited during
flooding events (Isphording and Lamb 1971; Isphording
1977; Hulbert and Whitmore 2006). The deltaic nature of
the Ecor Rouge Sand is supported by the presence of terres-
trial mammals, freshwater turtles (such as Apalone), croco-
diles, and fish remains (see Isphording and Lamb 1971;
MacFadden and Dobie 1998; Hulbert and Whitmore 2006;
Jasinski 2013). The late Miocene age for the Mauvilla local
fauna is supported by a mammal assemblage that best resem-
bles those from late early Hemphillian North American Land
Mammal Age (NAMLA) sites in central Florida (Hulbert and
Whitmore 2006).
Materials and methods
A total of 62 lots comprising 324 freshwater fish elements
were examined as part of this study, all of which were col-
lected from the Mauvilla fossil site. These specimens were not
collected by the present authors. Rather, they were historically
collected and directly examined by one or both of the authors
within the scientific collections at the following institutions:
AUMP: Auburn Museum of Paleontology, Auburn
University, Auburn, Alabama, USA.
MSC: McWane Science Center, Birmingham,
Alabama, USA.
USNM: United States National Museum, Washington, D.
C., USA.
Direct examination of the recovered material at MSC and
USNM showed that each institution cataloged the fish remains
into lots, with each lot containing one or more elements.
Although the elements that comprised a lot were cataloged
under the same number, it could not be determined whether
the elements belonged to an associated specimen or represented
isolated remains of one or more individuals that were cataloged
together. In many instances, this latter explanation appeared
most plausible as remains belonging to different taxa were fre-
quently cataloged together within the same lot. In these
instances, the elements were separated, and each identified
taxon was provided with a unique catalog number.
The specimens examined herein were collected from the
Mauvilla fossil locality at different times by different institu-
tions and individuals. The USNM specimens were collected
during the two Whitmore excavations in 1967 and 1981.
These specimens were recovered from a large excavation of
the creek bank, but also by screen washing sediments in the
creek (Hulbert and Whitmore 2006; Figure 2). Although the
date of collection was not recorded for several of the USNM
specimens examined, they were most certainly collected dur-
ing one of these two expeditions to the site. The MSC speci-
mens were all collected in December of 1981 by personnel
from the RMM. While several of the specimens were surface
collected from Whitmore’s main excavation, unpublished
field notes on file at MSC indicate that many of the smaller
specimens were derived from a bulk field sample that was
HISTORICAL BIOLOGY 3
collected from the same excavation site, then later processed
at the RMM. The AUMP specimens were collected in October
of 1988 by James Dobie, however his methods of collection
were not documented.
All figured specimens were photographed with a Nikon
D80 camera with Tamron macro lens, and all photographs
were rendered in Adobe Photoshop CC 2017 as part of
the production of the presented figures. The taxonomic
identifications presented herein were determined through
the direct comparison of elements, or photographs of the
elements, to extant osteological specimens housed at MSC
and the University of Tennessee Zooarchaeology
Collection (UT) in Knoxville, Tennessee, USA. The tax-
onomy utilised follows that of Nelson et al. (2016) and
anatomical terms follow that of Rojo (1991) and other
references cited herein. The North American Land
Mammal Ages (NAMLA) presented in Figure 2 follows
that of Barnosky et al. (2014).
Systematic paleontology
Class Osteichthyes Huxley 1880
Subclass Actinopterygii Klein 1885
Unranked Neopterygii Regan 1923
Infraclass Holostei Müller 1845
Division Ginglymodi Cope 1872
Order Lepisosteiformes Hay 1929
Family Lepisosteidae Cuvier 1825
Gen. et. sp. indet.
(Figure 3)
Material
USNM 171044 (vertebra).
Description
The lone Lepisosteidae vertebra in our sample preserves the
centrum only. This centrum measures 2.6 cm in anteropos-
terior length and has a greatest anterior diameter of 2.2 cm.
The vertebra is distinctly opisthocoelous – being convex ante-
riorly and concave posteriorly. The lateral edges are antero-
posteriorly concave and a series of alternating ossified ridges
and depressions are present. Both the anterior and posterior
faces are smooth and lack apparent growth rings. Although
the hemal and neural spines and transverse processes are not
preserved, the articulation points for these spines and pro-
cesses are visible.
Remarks
Among North American freshwater fishes, the opisthocoelous
nature of the vertebra in our sample is unique to the members
of the Lepisosteidae. This family currently consists of two
extant genera, Atractosteus and Lepisosteus (Nelson et al.
2016), and, according to Boschung and Mayden (2004), four
extant species are considered native to Alabama, Atractosteus
spatula (Lacepède 1803), Lepisosteus oculatus Winchell
(1864), Lepisosteus osseus (Linnaeus 1758), and Lepisosteus
platostomus Rafinesque (1820). While specimen USNM
177044 likely belongs to one of these aforementioned species,
4 J. A. EBERSOLE AND S. J. JACQUEMIN

Figure 3. Recent and late Miocene Lepisosteidae remains. (a–c) MSC 20959, Recent Lepisosteus oculatus Winchell (1864) vertebra in (a) anterior; (b) lateral; and (c)
posterior views. (d–f) USNM 171044, late Miocene Lepisosteidae vertebra in (d) anterior; (e) lateral; and (f) posterior views. (g–k) MSC 39043, Recent Atractosteus
spatula scales in lateral view; (g) Type 2 scale; (h) Type 3 scale; (i-j) Type 1 scales; (k) unserrated scale. (l–u) late Miocene Atractosteus sp. cf. A. spatula scales; (l) MSC
2238.7, Type 3 scale; (m) MSC 2238.4, Type 2 scale; (n) USNM 641879.1, Type 1 scale; (o) USNM 641879.2, Type 1 scale; (p) MSC 2238.1, unserrated scale; (q) USNM
331030, Type 2 scale; (r) USNM 629339.1, Type 3 scale (reversed for comparison); (s) USNM 641879.1, Type 1 scale; (t) USNM 629339.2, Type 1 scale (reversed for
comparison); (u) MSC 2238.18, unserrated scale (reversed for comparison). Scale bars = 1 cm.

the lack of differential characteristics on gar vertebrae prohi-
bits a more refined specific or generic placement. Although
Atractosteus individuals can achieve much larger body sizes
than those of Lepisosteus, their size ranges can overlap when
presented with younger (i.e. smaller) Atractosteus specimens
(see Boschung and Mayden 2004), rendering size as an insuf-
ficient characteristic to separate the vertebrae of these two
genera.
In Alabama, fossil members of the Lepisosteidae have been
reported from the Upper Cretaceous (Ikejiri et al. 2013),
middle Eocene (Maisch et al. 2016), and late Pleistocene
(Jacquemin et al. 2016). Elsewhere, the earliest reported fossil
member of the Lepisosteidae, Atractosteus, has been docu-
mented from deposits over 100 million years in age (Grande
2010; Wright et al. 2012). Molecular divergence estimates
suggest a deeper fossil history for the Lepisosteidae, with an
estimated familial divergence of approximately 260 mya
(Betancur-R et al. 2013). All members of this family inhabit
primarily low gradient lentic and lotic freshwater habitats and
are typically classified as generalist ambush style predators
that occupy some of the top trophic niches in aquatic habitats
(Page and Burr 2011). Although Miocene members of the

Lepisosteidae have been reported elsewhere in eastern North
America (see Smith 1981), the specimens reported here and
below represent the first Miocene occurrences of the family in
the Gulf Coastal Plain.
Genus Atractosteus Rafinesque 1820
Atractosteus sp. cf. A. spatula (Lacepède 1803)
(Figure 3)
Material
AUMP 3463 (scale), MSC 2238 (26 scales), USNM 173637
(scale), USNM 331017 (scale), USNM 331023 (3 scales),
USNM 331030 (5 scales), USNM 340707 (scale), USNM
629339 (10 scales), USNM 641873 (2 scales), USNM 641876
(5 scales), USNM 641879 (3 scales), USNM 641881 (4 scales).
Description
Our sample consists of 62 ganoid scales. These scales range in
size and shape, with the largest scales reaching almost 3 cm in
greatest dimension. The scales have a bony interior and an
outer layer of ganoine. The medial face of the scales is gen-
erally convex and smooth, but also appears pitted on some
specimens. The lateral face of the scales has varying degrees of
ganoine ornamentation which ranges from rhomboidal to
more globular, and some scales have finger-like lateral pro-
jections. Some scales display a punctate ganoine ornamenta-
tion, while others are smooth. The ganoine ornamentation
extends to the edges of the lateral faces on some scales and is
more centralised on others. The scales in our sample fall into
several distinct morphological groups, referred to as types I,
II, and III by Yang et al. (2013). Type I scales are the most
common and covered a majority of the body of the fish. These
scales are sub-rectangular in shape, have an elongated trian-
gular projection, and a single serrated edge. Type II scales
occupy the regions just anterior to the dorsal and anal fins
and are sub-circular in shape and lack serrated edges. Type III
scales reside just anterior to the caudal fin and are similar in
morphology to Type 1 scales by being sub-rectangular in
outline and by having an elongated projection. These latter
scales, however, have two distinctly serrated edges. A fourth
morphology is diamond-shaped in outline and lacks serrated
edges.
Remarks
Traditionally, scales belonging to Atractosteus and Lepisosteus
have been separated in the fossil record by the presence or
absence of ganoine ornamentation on the lateral face of the
scales (with unornamented scales being assigned to
Lepisosteus; see Wiley 1976; Maisch et al. 2016). Grande
(2010), however, discussed how the presence of ganoine
ornamentation on scales can be related to ontogeny (with
those on older individuals being more ornamented), and
can vary by the location of the scale on the body (i.e. more
prevalent in the trunk region, and often absent on scales in
other areas). Although Grande (2010) showed that unorna-
mented scales can belong to either genera, the scales recov-
ered in our sample all have distinct ganoine ornamentation,
HISTORICAL BIOLOGY 5
and therefore are all assigned to Atractosteus. Grande (2010)
and Nelson et al. (2016) recognised three extant species of
Atractosteus, with only one, A. spatula, having a modern
range in North America, including Alabama (Boschung and
Mayden 2004). The two other extant species, Atractosteus
tristoechus (Bloch and Schneider 1801) and Atractosteus tro-
picus Gill (1863), have current distributions in Cuba and from
Mexico to Costa Rica, respectively (Page and Burr 2011).
Given these modern distributions, it is likely the scales in
our sample belong to A. spatula. However, because the pre-
historic ranges of the other two Atractosteus is unknown, and
because scales cannot be used to differentiate these three taxa,
the specimens in our sample are here only tentatively
assigned Atractosteus sp. cf. A. spatula.
Division Teleosteomorpha Arratia et al. 2004
Subdivision Teleostei Müller 1846
Gen. et. sp. indet.
(Figure 4)
Material
MSC 2300 (vertebra), USNM 340995 (vertebra), USNM
341003 (vertebra), USNM 341489 (dorsal spine), USNM
341581 (vertebra), USNM 341688 (spine), USNM 347889
(vertebra), USNM 347899 (vertebra), USNM 641877 (dorsal
spine), USNM 697246 (vertebra), USNM 697247 (dorsal
spine), USNM 697248 (dorsal spine), USNM 697252
(vertebra).
Description
Both dorsal spines and vertebrae are present in our sample.
The vertebrae range in size from 2 to 5 cm in greatest anterior
diameter. In lateral view, the centra range in anteroposterior
length from 1 to 5 cm. The neural and hemal spines are not
preserved on these specimens, but their points of articulation
are visible. Articulation points for the transverse processes are
present on some specimens, but absent on others, indicating
both dorsal and caudal vertebrae are present in our sample.
The anterior and dorsal faces of the centra are strongly con-
cave, and in profile view, the lateral edges are slightly concave.
Growth rings are visible on the anterior and posterior faces of
the centrum on some specimens, and the shape of the cen-
trum ranges from circular to ovate in outline. Two or more
ossified ridges are present on the lateral edges of the centra.
The spines consist of both complete and incompletely pre-
served specimens. When proximal ends are preserved, all exhi-
bit two lateral condyles and a median foramen, indicating they
are dorsal spines. On all specimens, a shallow medioposterior
groove is present that extends nearly the length of the spine.
The spines taper to a distal tip and all lack lateral ornamenta-
tion. The anterior face on some specimens have an anterome-
dial groove, while others are either uniformly convex or have a
medial ridge that extends the length of the spine.
Remarks
Unfortunately, the isolated vertebrae with a bi-convex cen-
trum and the unornamented spines in our sample could not
6 J. A. EBERSOLE AND S. J. JACQUEMIN

Figure 4. Late Miocene Teleostei remains. (a–b) USNM 340995, vertebra in (a) medial; and (b) lateral views. (c–d) USNM 341581, vertebra in (c) medial; and (d) lateral
views. (e–f) USNM 347889, vertebra in (e) medial; and (f) lateral views. (g–i) USNM 697247, dorsal spine in (g) anterior; (h) posterior; and (i) lateral views. (j–l) USNM
341489, proximal end of a dorsal spine in (j) anterior; (k) posterior; and (l) lateral views. (m–o) USNM 341688, spine in (m) anterior; (n) posterior; and (o) lateral views.
(p–r) USNM 641877, proximal end of a dorsal spine in (p) anterior; (q) posterior; and (r) lateral views. (s–u) USNM 697248, proximal end of a dorsal spine in (s)
anterior; (t) posterior; and (u) lateral views. (v–w) USNM 347899, vertebra in (v) medial; and (w) lateral views. (x–y) USNM 347899, vertebra in (x) medial; and (y)
lateral views. (z–aa) USNM 347899, vertebra in (z) medial; and (aa) lateral views. Scale bars = 1 cm.

be identified beyond the Teleostei. Despite the lack of differ-
ential characteristics on these elements, the size and morphol-
ogy of the dorsal spines suggests a degree of taxonomic
diversity. Of those with preserved proximal ends, the distance
between the lateral extent of the lateral condyles in anterior or
posterior views ranges from 4 mm to 15 mm. This, in con-
junction with the presence or absence of an anteromedial
groove, suggests they belong to at least two separate taxa.
Although the ability to identify these remains beyond the
Teleostei could potentially reveal additional taxa at this site,
it is also likely that a majority of these elements belong to one
or more of the taxa identified herein.
Subseries Siluriphysi Fink and Fink 1996
Order Siluriformes Cuvier 1817
Suborder Siluroidei Frizzell and Dante 1965
Superfamily Ictaluroidea Gill 1861
Family Ictaluridae Gill 1861
Genus Ictalurus Rafinesque 1820
Ictalurus spp.
(Figures 5–6)
Material
MSC 2225 (3 spines), MSC 2299 (spine), MSC 2303 (spine),
MSC 38695 (spine), MSC 38,698 (spine), MSC 38,699 (spine),
USNM 173,625 (spine), USNM 340,670 (spine), USNM
340,708 (spine), USNM 340,740 (neurocranium), USNM
340,747 (spine), USNM 340,942 (pectoral spine), USNM
340,981 (spine), USNM 341,015 (spine), USNM 341,507 (pec-
toral spine), USNM 341,540 (spine), USNM 341,552 (dorsal
spine), USNM 341,657 (pectoral spine), USNM 341,675 (pec-
toral spine), USNM 341,691 (pectoral spine), USNM 347,963
(spine), USNM 347,984 (spine), USNM 641,874 (spine),
USNM 641,875 (spine), USNM 697,250 (pectoral spine),
USNM 697,251 (spine).
Description
Dorsal and pectoral spines and a fragmentary neurocranium
were identified in our sample. The pectoral spines have a
robust proximal end and elongated shaft that tapers distally
to a sharp point. The spines have a slight posterior bend in
dorsal and ventral views and have a slight ventral bend in
anterior and posterior views. The length of the spine is
ornamented with a series of alternating parallel ridges and
grooves. In basal view, the anterior articular process is ovate
and extends slightly dorsal to the dorsal anterior process. The
anterior and dorsal articular processes are separated by a
shallow groove. The dorsal articular process is three times
the size of the anterior articular process and is ovate in out-
line. In dorsal view, the dorsal articular process projects both
proximally and dorsally while the anterior articular process
projects proximally. A posterior medial groove is present that
extends from a deep basal recess to almost the distal tip of the
spine. The base of the medial groove is lined with a series of
posterior serrae. The tips of the posterior serrae are not
preserved on most specimens, but when present, consists of
 HISTORICAL BIOLOGY 7

Figure 5. Recent and late Miocene Ictalurus spp. spines. (a–c) UT 3460, Recent Ictalurus furcatus dorsal spine in (a) anterior; (b) posterior; and (c) lateral views. (d–f)
UT 3460, Recent Ictalurus furcatus pectoral spine in (d) anterior; (e) posterior; and (f) dorsal views. (g–i) UT 8474, Recent Ictalurus punctatus pectoral spine in (g)
anterior; (h) posterior; and (i) dorsal views. (j–l) UT 8417, Recent Ictalurus punctatus pectoral spine in (j) anterior; (k) posterior; and (l) dorsal views. (m–o) USNM
173625, late Miocene Ictalurus sp. dorsal spine fragment in (m) anterior; (n) posterior; and (o) lateral views. (p–r) USNM 341552, late Miocene Ictalurus sp. dorsal
spine fragment in (p) anterior; (q) posterior; and (r) lateral views. (s–u) USNM 697250, Late Miocene Ictalurus sp. pectoral spine in (s) anterior; (t) posterior; and (u)
dorsal views. (v–x) USNM 340942, late Miocene Ictalurus sp. pectoral spine in (v) anterior; (w) posterior; and (x) dorsal views. (y–aa) USNM 341675, late Miocene
Ictalurus sp. pectoral spine in (y) anterior; (z) posterior; and (aa) dorsal views. (bb–dd) USNM 340708, late Miocene Ictalurus sp. spine in (bb) anterior; (cc) posterior;
and (dd) dorsal views. (ee–gg) USNM 340747, late Miocene Ictalurus sp. spine in (ee) anterior; (ff) posterior; and (gg) dorsal views. (hh–jj) USNM 347984, late
Miocene Ictalurus sp. spine in (hh) anterior; (ii) posterior; and (jj) dorsal views. (kk–mm) USNM 341540, late Miocene Ictalurus sp. spine in (kk) anterior; (ll) posterior;
and (mm) dorsal views. Scale bars = 1 cm.

proximally inclined triangular barbs that decrease in size
distally. The anterior edge of the spine lacks serrae, but
instead has a shallow anteromedial groove that is visible
along the entire length of the shaft. This groove is shallower
on larger specimens, and, at times, a medial ridge is present
within the groove. Only the proximal ends of the dorsal
spines are present in our sample. Although these spines are
similar to those we assigned to the Teleostei, they were
differentiated by having lateral ornamentation consisting of
alternating parallel ridges and grooves.
The neurocranium in our sample (Figure 6(a–b)) is
incomplete and preserves only the basioccipital. This robust
element measures 67 mm in greatest anteroposterior length,
and 33 mm in mediolateral diameter across the articular
8 J. A. EBERSOLE AND S. J. JACQUEMIN

Figure 6. Recent and late Miocene Ictalurus neurocrania. (a–b) USNM 340740, late Miocene cf. Ictalurus sp. basioccipital in (a) ventral; and (b) posterior views. (c–e)
UT 1137, Recent Ictalurus punctatus neurocranium; (c) ventral view; (d) posterior view); and (e) close-up of basioccipital in posterior view. Scale bars = 2 cm.

surface of the basioccipital in posterior view. In ventral view,
the deep foramen magnum is visible on the basioccipital and
is located just anterior to the articular surface. In posterior
view, only the ventral half of the basioccipital is preserved.
The preserved portion has a slightly concave articular face
that is covered with a series of growth lines that form con-
centric circles around a shallow medial canal. In posterior
view, the ventral edge of basioccipital has a distinct medial
depression.
Remarks
Given the large number of fossil spines recovered and rela-
tively good preservation of many, careful consideration was
given to comparisons with other genera and species that
could possibly be encountered from the region. The fossil
spines in our sample were compared to representatives from
each of the native genera within the Ictaluridae (i.e. Ameiurus,
Ictalurus, Noturus, and Pylodictis). Furthermore, because we
could not rule out the possible presence of sea catfishes at the
Mauvilla fossil locality, we also compared the fossil specimens
to two members of the Arridae that have modern ranges in
the northern Gulf of Mexico, the Hardhead Catfish Arius felis
Linneaus 1766 and Gafftopsail Catfish Bagre marinus Mitchell
1815 (Hoese and Moore 1998). In particular, spines were
compared using size and shape of the anterior and posterior
serrae, overall spine curvature, and the presence or absence of
striations or fluting along the spine shafts.

The ornamentation on the spines in our sample was
remarkably consistent and is represented by a series of alter-
nating parallel ridges and grooves. These longitudinal stria-
tions delineate these fossil spines from the native members of
Noturus that have either unornamented spines (i.e. N. gilberti,
N. lachneri, N. leptacanthus) or spines that are fluted, rather
than striated (i.e. N. elegans, N. flavater, N. eleutherus, N.
flavippinis, N. flavus, N. gyrinus, N. insignis, N. miurus, N.
munitus, and N. stigmosus). Furthermore, the fossil spines
lack the extremely large and recurved posterior serrae present
on N. elegans, N. eleutherus, N. flavater, N. flavippinis, N.
miurus, N. munitus, and N. stigmosus. Although similar lat-
eral ornamentation can be seen on the spines of the native
members of Ameiurus and Pylodictis, the fossil spines in our
sample are consistent in lacking anterior serrae or notches,
separating them from A. melas, A. natalis, A. nebulosus, and
P. olivaris. Of the sea catfish, the spines of Ariopsis felis and
Bagre marinus differ from those in our sample by a combina-
tion of being fluting and by having both anterior and poster-
ior serrae.
The lateral ornamentation, spine angle, and posterior ser-
rae patterning on the spines in our samples are all diagnostic
for Ictalurus. There are currently two recognized native spe-
cies of Ictalurus in Alabama, I. furcatus (Valenciennes in
Cuvier and Valenciennes 1840) and I. punctatus (Rafinesque
1818) (Boschung and Mayden 2004), and at least one extinct
taxon, I. countermani (Lundberg and Luckenbill 2012), has
been documented from Miocene deposits in Maryland, USA.
Our specimens differ from I countermani by lacking the small
anterior serrae present on the pectoral spine of this taxon.
Unfortunately, close comparison of the spines in our sample
to extant osteological specimens, as well to others in the
literature, did not facilitate further delineation beyond
Ictalurus. Although I. furcatus spines can grow much longer
than those of I. punctatus, there appears to be a paucity of
additional consistent characteristics that can be used to dif-
ferentiate the pectoral spines of these two species (see Duvall
2007). Due to the similarity of extant I. furcatus and I.
punctatus spines and the size overlap of the of the fossil
spines in our sample, it is possible that that one or both of
these Ictalurus species is present at the Mauvilla Fossil Site.
It should also be noted that some of the spines in our
sample lack distinguishing features such an articular head or
preserved serrae, precluding identification as pectoral or dor-
sal spines. These specimens, consisting mostly of a laterally
ornamented shaft, with or without serrae, could not deli-
neated further than ‘spine’. The partial neurocranium in our
sample compared very favorably to that from large extant
Ictalurus furcatus and Ictalurus punctatus specimens in
respect to its size, the position of the foramen magnum, and
the distinct medial depression on the ventral edge of the
basioccipital. Unfortunately, due to the incomplete preserva-
tion of the fossil specimen, it could only tentatively be
assigned to this genus.
The Ictaluridae consists of seven extant genera, Ictalurus,
Ameiurus, Noturus, Pylodictus, Satan, Trogloglanis, and
Prietella (Nelson et al. 2016), with 19+ species representing
the former four genera considered native to Alabama
(Boschung and Mayden 2004). The members of this family
HISTORICAL BIOLOGY 9
inhabit a range of lentic and lotic ecosystems and are typically
classified as benthic omnivores known to consume a wide
variety of prey including insects, fish, mollusks, crayfish, and
annelids (Page and Burr 2011). The earliest known members
of the Ictaluridae date to the late Eocene, with remains of
Ictalurus being documented from 30+ million-year-old
deposits (Prothero 1995). Molecular evidence pushes the ori-
gin of this family back even further, suggesting a familial
divergence estimate of approximately 56 mya (Mariangeles
et al. 2017). Although Miocene members of the Ictaluridae
have been reported from both eastern and western North
America (Smith 1981), the specimens in our sample represent
the first Miocene occurrences in Alabama. Fossil Ictalurus
specimens have been documented from late Pleistocene
deposits in both the northern and central regions of the
state (Jacquemin et al. 2016).
Cohort Eutelostei Rosen 1985
Superorder Acanthopterygii Greenwood et al. 1966
Series Percomorpha Cope 1871
Subseries Ovalentaria Smith and Near 2012 in
Wainwright et al. 2012
Order Acanthuriformes Jordan 1923
Suborder Sciaenoidei Cuvier and Valenciennes 1828
Family Sciaenidae Linnaeus 1758
Genus Aplodinotus Rafinesque 1819
Aplodinotus grunniens Rafinesque 1819
(Figure 7)
Material
AUMP 3456 (pharyngeal tooth), AUMP 3457 (pharyngeal
tooth), MSC 2297 (188 pharyngeal teeth), MSC 38696 (phar-
yngeal tooth), MSC 38700 (23 pharyngeal teeth), USNM
340746 (pharyngeal tooth), USNM 341700 (pterygiophore),
USNM 347964 (upper left pharyngeal), USNM 641872 (2
pharyngeal teeth), USNM 641880 (pharyngeal tooth).
Description
This taxon is represented in our sample by a partial pharyn-
geal, pharyngeal teeth, and a pterygiophore. The lone phar-
yngeal in our sample (Figure 4(p–r)) is incompletely
preserved, but a majority of the occlusal surface and symphy-
seal margin is intact. The occlusal surface is covered with
tightly packed cylindrical aveoli that decrease in diameter
towards the lateral edge. The symphyseal margin is smooth
and uniform in thickness. The element is slightly concave
dorsally and has a slightly convex oral surface. The dorsal
face of the pharyngeal is smooth and preserves a ridge that
extends nearly the entire length of the symphyseal margin,
terminating at a dorsally projected muscle attachment point
near the posterior margin of the plate. The lateral, anterior,
and posterior portions of the tooth plate are not preserved.
This specimen is identified as an upper left pharyngeal.
The pharyngeal teeth in our sample (Figure 4(d–l)) range
in size from 2.0 to 4.0 mm in greatest occlusal diameter. In
profile view, the lateral edges of the teeth are biconvex, and
many of the teeth are slightly tapered basally. The teeth vary
10 J. A. EBERSOLE AND S. J. JACQUEMIN

Figure 7. Recent and late Miocene Aplodinotus grunniens remains. (a–c) UT 9375, Recent tooth in (a) occlusal; (b) lateral; and (c) basal views. (d–f) MSC 2297, late
Miocene tooth in (d) occlusal; (e) lateral; and (f) basal views. (g–i) USNM 641872.1, late Miocene tooth in (g) occlusal; (h) lateral; and (i) basal views. (j–l) USNM
641872.2, late Miocene tooth in (j) occlusal; (k) lateral; and (l) basal views. (m–o) MSC 7945, Recent upper left pharyngeal in (m) dorsal; (n) occlusal; and (o)
symphyseal views. (p–r) USNM 347964, late Miocene upper left pharyngeal in (p) dorsal; (q) occlusal; and (r) symphyseal views. (s–u) USNM 341700, late Miocene
pterygiophore in (s) anterior; (t) posterior; and (u) proximal views. (v–x) MSC 7945, Recent pterygiophore in (v) proximal; (w) anterior; and (x) posterior views. Scale
bars = 1 cm.

in occlusal outline, being rounded, sub-rounded, and some-
times angular, and the oral surface of the teeth ranges from
flat to slightly concave. The apical one-half to two-thirds of
the teeth are covered by a smooth enamel coating. A shallow,
rounded, medial pulp cavity is present on the tooth base. The
teeth vary in diameter depending on their position within the
mouth, with smaller teeth generally occupying the lateral
positions on the tooth plate.
The pterygiophore (Figure 7(s–u)) measures nearly 8 cm
as preserved. This spine is sub-triangular in cross-section
and has two prominent medial ridges along the length of the
anterior face that are bisected by a shallow groove. The
posterior face is smooth and slightly convex. A slight medial
ridge is present, but it does not extend to the tip of the
spine. Two distinct depressions flank the medial ridge near
the proximal margin of the spine. In lateral view, the spine
gently tapers distally, and an anteroposterior concavity
extends nearly the length of the element. In proximal view
(Figure 7(u)), the pterygiophore has seven distinct, blunt,
laterally extending, projections. The anterior pointed projec-
tion is formed by the anterior medial ridge. This projection
is flanked by two anterior articular facets that extent

anterolaterally. The two largest projections extend laterally
and are formed by two proximal articular facets. The two
posterior projections are formed by the bifurcation of the
posterior median ridge. The anterior and posterior articular
facets are slightly concave on the proximal face. The dorsal
tip of the pterygiophore is not preserved.
Remarks
The fossil sciaenid elements in our sample resemble those of
three extant members within the family, the Freshwater
Drum (Aplodinotus grunniens Rafinesque 1819), the saltwater
Black Drum (Pogonias cromis (Linnaeus 1766)), and the salt-
water Red Drum (Sciaenops ocellatus (Linnaeus 1766)). Due
to the deltaic nature of the sediments exposed at the Mauvilla
fossil locality, we could not rule out the possibility of estuar-
ine fishes being present within our sample. As a result, the
fossil specimens were compared to extant remains belonging
to each of the aforementioned sciaenid taxa. Although the
fossil teeth and pterygiophore were nearly indistinguishable
from those belonging to each of the extant species, slight
differences were observed between the upper pharyngeals of
the three taxa.
On the fossil pharyngeal, a prominent ridge is preserved
on the dorsal surface that extends nearly the entire length of
the symphyseal margin and terminates at a dorsally projected
muscle attachment point towards the posterior margin of the
tooth plate. This ridge is prominent on the upper pharnygeals
of A. grunniens and P. cromis, and a comparison between the
two showed this characteristic to be very similar in overall
morphology on both taxa. This ridge, however, is absent on
the upper pharyngeals of S. ocellatus, indicating that the fossil
tooth plate does not belong to a Red Drum. Although the
fossil pharyngeal in our sample is incompletely preserved, it
can be differentiated from that of the similar P. cromis by the
angle of the symphyseal face, which, as observed on extant
specimens, is more vertical on A. grunniens and more medi-
ally flared on P. cromis. Furthermore, our observations of
adult and juvenile extant specimens showed that sub-adult
A. grunniens upper pharyngeals have a similar oval-shaped
tooth patch to that on P. cromis. The adult A. grunniens
specimens, however, show a degree of ontogenetic variation
as the adult upper pharyngeals become more laterally
extended as the fish increases in size. This degree of ontoge-
netic change was not observed on the upper pharyngeals of P.
cromis as the tooth patch on both adult and sub-adult speci-
mens appear to retain their oval outline. Because the fossil
pharyngeal compared more favorably to a smaller, rather than
larger, Freshwater Drum, it can be assumed that the fossil
specimen belonged to a sub-adult individual. Despite the lack
of differential characteristics on the remaining sciaenid ele-
ments in our sample, all are here assigned to A. grunniens
based on the presence of the identified pharanygeal. Although
saltwater specimens may be present at the Mauvilla Fossil
Site, additional, and more diagnostic, material is needed to
confirm the presence of these other taxa.
A. grunniens is the only currently recognised native fresh-
water species within the Sciaenidae that resides in Alabama
and is known to inhabitant a range of lentic and lotic habitat
types ranging from mid-to-large sized rivers and streams to
HISTORICAL BIOLOGY 11
lakes (Boschung and Mayden 2004). Members of the
Sciaenidae are typically classified as benthic foragers and are
known to consume a wide variety of prey (mollusks, insects,
fish, etc.) in their respective habitats (Page and Burr 2011;
Jacquemin et al. 2014).
Although assessing the complete prehistoric range of the
Sciaenidae is not possible at this time, based on fossil records
of A. grunniens, it is clear this taxon maintained a large
geographic range in the past, likely similar to the extent of
the modern drum across North America. Molecular analyses
have suggested A. grunniens diverged from a euryhaline
ancestor in North America approximately 19.5 mya (Lo
et al. 2015). Based on the examination of otoliths, Nolf
(2003) recognised several lower Oligocene and Miocene sciae-
nid species within marine deposits along the Gulf Coast of
North America. This led Nolf (2003) to surmise that fresh-
water sciaenid taxa from North and South America were
likely derived from a marine ancestor, likely during the
Miocene. Given that A. grunniens is the only freshwater
sciaenid in North America, the specimens in our sample
provide an important contribution to the fossil record as
they represent some of the earliest definitively freshwater
evidence for a member of the Sciaenidae. A. grunniens speci-
mens have also been reported from late Pleistocene deposits
in Alabama (Jacquemin et al. 2016).
Discussion
In this study we examined 324 elements recovered from the
late Miocene (Hemphillian) Mauvilla fossil site, and identified
three unequivocal taxa, including Alligator Gar (Atractosteus
sp. cf. A. spatula), Channel or Blue Catfish (Ictalurus spp.),
and Freshwater Drum (Aplodinotus grunninens). Although
these remains date to the latest Miocene, we support the
utilisation of extant genus and species names as these ele-
ments are nearly identical morphologically to those of extant
representatives. Furthermore, previous molecular divergence
estimates suggest the presence of these taxa in deposits pre-
dating those here, lending further support to their presence in
the Miocene. The combination of these lines of evidence
strongly suggests that these remains belong to early fossil
representatives of the modern taxa rather than different gen-
era or species. In addition, while molecular divergence times
indicate that these taxa existed at least 10+ million years prior
to the late Miocene, these specimens help provide tangible
fossil evidence that helps to fill the gaps in the ghost lineages
that exist between phylogenetic divergence estimates and the
modern taxa.
Not supported in this study is the reported occurrence of
‘Scombroidei, mackerels and related fish’ at the Mauvilla site
as listed by Isphording and Lamb (1971, p. 776). This report
is likely the result of the misidentification of the recovered
remains, as no such taxa were identified in our sample (which
includes all fish remains recovered from the site), and extant
members of the Scombroidei are largely marine taxa (see
Nelson et al. 2016).
While the presence of only three unequivocal taxa does not
lend itself to a complete faunal reconstruction, the identified
taxa do provide some inferences towards their habitat and
12 J. A. EBERSOLE AND S. J. JACQUEMIN
larger assemblage. Based on modern assemblage-habitat rela-
tionships, the co-occurrence of these three taxa indicates that
the habitat was likely a medium-to-large river with a diversity
of microhabitats (Jacquemin and Pyron 2011). It is also likely
that this medium-to-large river habitat contained a variety of
other freshwater fish typical of these habitats today. However,
conspicuously absent from our Mauvilla sample are any
number of typical taxa one would expect to co-occur in a
typical large river fish assemblage in this area (e.g.
Catostomidae such as Moxostoma, Cyprinidae such as
Nocomis, Centrarchidae such as Micropterus, and more;
Jacquemin et al. 2016). This lack of other native species
typical of an Alabama large river assemblage likely indicates
a taphonomic bias. This bias towards the preservation of
robust elements from large taxa extends beyond fishes, as
prior studies describing the mammal assemblage from this
site also reported the absence of small taxa (see Isphording
and Lamb 1971; MacFadden and Dobie 1998; Hulbert and
Whitmore 2006). The Mauvilla site was interpreted by
Isphording (1977), Isphording and Lamb (1971), and
Hulbert and Whitmore (2006) as representing a deltaic flood-
plain, with the vertebrate material likely deposited during
flooding events. Hulbert and Whitmore (2006) suggested
that this depositional energy was likely responsible for this
bias towards larger taxa as smaller individuals were likely
swept away by the currents.
The only watershed in Alabama that currently sustains
population of Atractosteus spatula, Ictalurus spp., and
Aplodinotus grunninens is the Mobile Basin (Boschung and
Mayden 2004). This expansive alluvial plain is the second
largest delta in the United States and serves as the drainage
for the Apalachee, Blakely, Middle, Mobile, Raft, Spanish, and
Tensaw rivers. Although a geomorphological study by Smith
(1988) suggested that the Mobile Basin did not take its cur-
rent form until after the late Pleistocene, it was also suggested
that one or more of the constituting rivers was likely present
in pre-Pleistocene times. Of the rivers that make up this delta,
the Mobile River is the closest the Mauvilla fossil locality.
Located only 15 kilometers to the east, this river may be the
source of the specimens at the site.
Despite of the taphonomic bias present in this sample, the
identification of these three taxa not only adds to the diversity
of the Mauvilla local fauna, but each also represents the first
Miocene freshwater fishes described from the Gulf Coastal
Plain in North America. Equally important, this study exem-
plifies the paucity of faunal remains for this epoch in this
region. Hopefully future investigations within the Miocene
strata in Alabama will yield additional remains and further
add to the diversity of fossil fishes known from in this region
as well as lend insights into the origins of many of the
modern taxa.
Conclusions
Three unequivocal taxa, Atractosteus sp. cf. spatula,
Ictalurus spp., and Aplodinotus grunninens were identified
from 324 elements historically collected from the late
Miocene Mauvilla fossil locality in Alabama, USA. The
taxonomic assignment to these extant taxa is supported by
their morphological similarity to modern representatives as
well as molecular divergence estimates that suggests these
taxa were present in deposits that predate the Miocene.
From a stratigraphic perspective, these specimens help fill
the gaps in the ghost lineages that exist between phyloge-
netic divergence estimates and the modern taxa and repre-
sent some of the earliest fossil occurrences of these extant
taxa. From a biogeographical standpoint, these identified
taxa represent the first Miocene occurrences for each in
Alabama, as well as within the Gulf Coastal Plain of North
America. The co-occurrence of these three taxa indicate
they were likely part of a medium-to-large river fish assem-
blage, and the absence of typical taxa such as Micropterus,
Moxostoma, and Nocomis, suggests a strong taphonomic
bias is present within our sample that appears to favor
larger taxa. Extant representatives of the identified taxa
have modern ranges in the nearby Mobile Basin, strongly
suggesting that one or more of the rivers that make up the
modern delta were present in the late Miocene. Overall, this
study lends to our understanding of Miocene fish distribu-
tions and provides some indication as to biodiversity as well
as habitat conditions in the region at this time.
Acknowledgments
We extend our gratitude to Amanda Millhouse for granting access to the
paleobiology collections at the USNM and for her assistance in assigning
new catalog numbers to many of the USNM specimens. We also thank
Connie Randall, Meagan Dennison, Jennifer Green, and Alex Bentley of
the Department of Anthropology at University of Tennessee, Knoxville
for their assistance with accessing comparative specimens in the UT
osteology collections. Ray Wilhite is thanked for allowing access to the
AUMP collections, and Walter Klippel (UT, retired) and John Lundberg
(Drexel University, Philadelphia, Pennsylvania) are thanked for their
insights concerning several of the Mauvilla fossil remains. Sandy
Ebersole (Geological Survey of Alabama, Tuscaloosa) is thanked for
providing the GIS shapefiles necessary to produce Figure 1. Finally, we
thank Jon Armbruster (Auburn University, Alabama), Gary L. Stringer
(University of Louisiana at Monroe), and Gareth J. Dyke (Historical
Biology) for their insightful reviews of this work.
Disclosure statement
No potential conflict of interest was reported by the authors.
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