First record of the ichthyodectiform fish Xiphactinus (Teleostei) from Patagonia, Argentina

Abstract

We report the first record of the genus Xiphactinus from southern South America. The recovered fossil material consists of an associated maxilla and abdominal vertebra, probably derived from latest Maastrichtian marine deposits of the Salamanca Formation in Chubut Province, Argentina. Xiphactinus has been widely reported from Late Cretaceous strata throughout the Northern Hemisphere, although to date, equivalent Southern Hemisphere occurrences include only a single specimen from Venezuela. Our new discovery thus indicates that Xiphactinus had a much more cosmopolitan distribution, encompassing the southern paleolatitudinal extremities of South America during the terminal Cretaceous.

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First record of the ichthyodectiform fish
Xiphactinus (Teleostei) from Patagonia, Argentina
Julieta J. De Pasqua, Federico L. Agnolin & Sergio Bogan
To cite this article: Julieta J. De Pasqua, Federico L. Agnolin & Sergio Bogan (2020): First record
of the ichthyodectiform fish Xiphactinus (Teleostei) from Patagonia, Argentina, Alcheringa: An
Australasian Journal of Palaeontology, DOI: 10.1080/03115518.2019.1702221
To link to this article: https://doi.org/10.1080/03115518.2019.1702221
Published online: 02 Mar 2020.
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First record of the ichthyodectiform fish Xiphactinus (Teleostei) from
Patagonia, Argentina
Julieta J. De Pasqua, Federico L. Agnolin, and Sergio Bogan
ABSTRACT
We report the first record of the genus Xiphactinus from southern South America. The recovered
fossil material consists of an associated maxilla and abdominal vertebra, probably derived from lat-
est Maastrichtian marine deposits of the Salamanca Formation in Chubut Province, Argentina.
Xiphactinus has been widely reported from Late Cretaceous strata throughout the Northern
Hemisphere, although to date, equivalent Southern Hemisphere occurrences include only a single
specimen from Venezuela. Our new discovery thus indicates that Xiphactinus had a much more
cosmopolitan distribution, encompassing the southern paleolatitudinal extremities of South
America during the terminal Cretaceous.
ARTICLE HISTORY
Received 21 May 2019
Revised 15 November 2019
Accepted 2 December 2019
KEYWORDS
Ichthyodectiformes; South
America; terminal
Cretaceous; Gondwana
Julieta J. De Pasqua [julietadepasqua@gmail.com], Laboratorio de Anatom

ıa Comparada y Evoluci
on de los Vertebrados, Museo Argentino de
Ciencias Naturales ‘Bernardino Rivadavia’, Av.

Angel Gallardo, 470, C1405DJR, Buenos Aires, Argentina; Federico L. Agnolin
[fedeagnolin@yahoo.com.ar], Laboratorio de Anatom

ıa Comparada y Evoluci
on de los Vertebrados, Museo Argentino de Ciencias Naturales
‘Bernardino Rivadavia’, Av.

Angel Gallardo, 470, C1405DJR, Buenos Aires, Argentina. CONICET; Fundaci
on de Historia Natural ‘F
elix de Azara’,
Departamento de Ciencias Naturales y Antropolog

ıa, Universidad Maim
onides; Hidalgo 775 piso 7, C1405BDB, Buenos Aires, Argentina; Sergio
Bogan [sergiobogan@yahoo.com.ar], Fundaci
on de Historia Natural ‘F
elix de Azara’, Departamento de Ciencias Naturales y Antropolog

ıa,
Universidad Maim
onides; Hidalgo 775 piso 7, C1405BDB, Buenos Aires, Argentina.
ICHTHYODECTIFORMES, most commonly known as
‘bulldog fishes’, were a geographically widely distributed clade
of stem teleosts, whose fossils have been recovered from mar-
ine deposits throughout North and South America, Antarctica,
Africa, Australia, Asia and Europe, and with a chronostrati-
graphical range from the from the Middle Jurassic to Late
Cretaceous (Chang 1963, Bardack 1965, Patterson & Rosen
1977, Schaeffer & Patterson 1984,Tanimoto&Kinkyo2001,
Forey et al. 2003, Arratia et al. 2004,Kimet al. 2014,
Alvarado Ortega and Porras M
uzquiz 2015,Yabumotoet al.
2018). Their general body-plan, preserved gut contents, and
jaw structure suggest an ecological role as higher trophic-level
pelagic macropredators (Cavin et al. 2013,2015).
The oldest recorded ichthyodectiform taxon, Saurocephalus,
was described in the early nineteenth century, and thought to
be a reptile (Harlan 1824). However, over 100 years later,
Bardack & Sprinkle (1969) established that Saurocephalus and
its relatives were indeed fishes, and could be classified as the
Order Ichthyodectiformes. Today, Ichthyodectiformes com-
prises more than 25 nominal genera, including Xiphactinus
Leidy, 1870 (Taverne 2010, Carillo-Brice~
no et al. 2012,Cavin
& Berrell 2019,King&Super2019), which is recognized as
one of the largest-bodied stem teleost taxa known from the
Late Cretaceous (Bardack 1965, Schwimmer et al. 1997,
Shimada et al. 2006, Everhart et al. 2010).
Often spectacular skeletons of Xiphactinus have been
found in the Niobrara Formation of Kansas, USA, and
reveal individuals of up to 5 m in maximum snout–tail
length (Cope 1872, Bardack 1965). Nevertheless, remains of
Xiphactinus have also been recovered in South America,
with fossils previously reported from the Late Cretaceous La
Luna Formation of Venezuela (Carrillo–Brice~
no et al. 2012),
and here for the first time, from the Late Cretaceous–earliest
Paleogene successions exposed in the Chubut Province of
Argentina. The aim of this paper, therefore, is to document
the new Argentinian occurrence of Xiphactinus and assess
its paleobiogeographical implications.
Geological setting
The new Argentinian Xiphactinus specimens, MACN Pv
15979 and MACN Pv 15980, were excavated in April 1946
within the Colhu
e Huapi area of southeastern Chubut
Province in Patagonia (Fig. 1). The Colhue Huapi area con-
tains Late Cretaceous to Paleogene strata including the
Coniacian–Maastrichtian Lago Colhu
e Huapi Formation,
which represents a fluvial channel sequence (Casal et al.
2015, Vallati et al. 2016), and the overlying marine
Salamanca Formation, which is Maastrichtian–lower
Thanetian in age (Clyde et al. 2014); this unit also incorpo-
rates the Cretaceous–Paleogene boundary at its base
(Legarreta & Uliana 1994, Gelfo et al. 2009, Clyde et al.
2014). The Salamanca Formation reflects the transgressive
onset of the Atlantic margin into the San Jorge Basin of cen-
tral Patagonia (Camacho 1967), and has yielded a rich fossil
record of marine micro- (Frenguelli 1936, Camacho 1954,
1967,M

endez 1966, Masiuk 1967, Bertels 1975, Matheos
et al. 2005) and macro-invertebrates (Ihering 1903, Feruglio
1949, Camacho 1967, Chebli & Serraiotto 1974, Andreis
1977, Parma & Casad
ıo2005), together with shark teeth
(Arratia & Cione 1996). Although uncertain, we suspect that
ß2020 Geological Society of Australia Inc., Australasian Palaeontologists
ALCHERINGA: AN AUSTRALASIAN JOURNAL OF PALAEONTOLOGY
https://doi.org/10.1080/03115518.2019.1702221
Published online 02 Mar 2020

MACN Pv 15979 and MACN Pv 15980 probably also
derived from this succession based on the associated assem-
blage and depositional setting.
Systematic paleontology
Order ICHTHYODECTIFORMES Bardack & Sprinkle, 1969
Suborder ICHTHYODECTOIDEI Romer, 1966
Family ICHTHYODECTIDAE sensu Patterson &
Rosen, 1977
Genus Xiphactinus Leidy, 1870
Xiphactinus sp.
Referred material. MACN Pv 15979, an incomplete maxilla
(Fig. 2A–C), and MACN Pv 15980, an abdominal vertebra (Fig.
3A–C); these specimens likely represent a single individual.
Description. MACN Pv 15979 is 18 mm in maximum trans-
verse width, 76 mm in maximum height, and 146mm in
maximum length (Fig. 2). The dorsal margin is incompletely
preserved, but can be identified as a maxilla because of its
length, transverse flattening, and straight profile; this differs
from the typically sinuous profile of the dentary in
Xiphactinus (see Bardack 1965). MACN Pv 15979 is triangu-
lar in lateral view and tapering posteriorly. The lateral sur-
face bears faint longitudinal striae; the anterolateral surface
is shallowly concave. Eleven thecodont alveoli are visible;
these steadily increase in diameter toward the mid-length of
the row, and then gradually decrease posteriorly. The pre-
served teeth are sub-circular in basal outline and of variable
size, with minimum/maximum diameters of 4/11 mm.
Their crowns have a narrow band of dentin surrounding
the pulp cavity; no carinae were observed.
MACN Pv 15980 is poorly preserved but was clearly
dorsoventrally compressed and flattened ventrally, with
articular facets for the autogenous neural and hemal arches
(Fig. 3). The lateral surface of the centrum bears a promin-
ent longitudinal ridge bordered by deeply incised fossae.
The articular surfaces are sub-circular in outline with con-
centric growth rings. A fragment of the articulated neural
arch has a sub-oval longitudinal fossa on its lateral side.
Discussion
Taxonomic assignment
MACN Pv 15979 and MACN Pv 15980 can be referred to
Ichthyodectiformes based on the presence of an elongate
maxilla with a single row of teeth, and autogenous neural
arches that are not fused to the centrum (Bardack 1965,
Patterson & Rosen 1977, Mkhitaryan & Averianov 2011).
They are also consistent with the genus Xiphactinus because
of the irregular maxillary tooth size, thecodont tooth
implantation, and longitudinal lateral ridge on the vertebral
centrum that is delimited by deeply incised fossae (Bardack
1965, Nelson 1973). MACN Pv 15979 differs from basal ich-
thyodectiforms, such as Thrissops, which has a curved maxil-
lary profile with small conical teeth of regular size and
uniform spacing; the anterior extremity of the jaw is also
edentulous (Taverne 2008). In contrast, derived ichthyodec-
tiforms, such as Xiphactinus and saurodontids possess
shorter maxillae with reduced edentulous extremities and
raised dental rows (Taverne 2008), as in MACN Pv 15979.
Like MACN Pv 15979, ichthyodectids including Gillicus,
Cooyoo and Ichthyodectes, exhibit sub-circular tooth crowns
with thecodont implantation, but differ in their more uni-
form tooth size in comparison to Xiphactinus (Newton
1877, Bardack 1965, Lees & Bartholomai 1987, Taverne
2008). Gillicus also has minute teeth and a curved maxillary
shape (Lees & Bartholomai 1987, Taverne 2008).
Furthermore, Aidachar can be distinguished from both
MACN Pv 15979 and Xiphactinus by its anteriorly in-curved
(by about 20–25) maxilla (evident elsewhere in Gillicus and
Chirocentrites: Taverne 2008) that has a sinusoidal alveolar
margin (Mkhitaryan & Averianov 2011). Lastly, MACN Pv
Figure 1. Map showing the recovery locality of the Argentinean
Xiphactinus fossils.
2 JULIETA J. DE PASQUA ET AL. XIPHACTINUS IN SOUTH AMERICA

15979 lacks the conspicuous tooth foramina (¼replacement
alveoli) of saurodontines (Cavin et al. 2013).
Within Xiphactinus, MACN Pv 15979 most closely resem-
bles Xiphactinus audax in its sub-circular tooth cross-sections,
and lack of distinct carinae; the teeth are otherwise elliptical
in Xiphactinus vetus, whose dentition also displays prominent
carinae (Schwimmer et al. 1997,Vavreket al. 2016).
Paleobiogeographical implications
Historically, the remains of Xiphactinus have been widely
reported from Cenomanian–Maastrichtian strata across the
Northern Hemisphere (Bardack 1965, Russell 1988,1993,
Schwimmer et al. 1997, Cumbaa et al. 2006, DeMar &
Breithaupt 2006, Vavrek et al. 2016), although recently
Figure 2. A–C,Xiphactinus sp. (MACN Pv 15979) incomplete maxilla in A, occlusal and B, lateral views. C, Reconstruction of Xiphactinus cranium indicating the
anatomical position of MACN Pv 15979. Abbreviations: te: teeth, al: alveoli. Scale bar ¼50 mm in A, B; C, not to scale and modified from Bardack (1965).
ALCHERINGA: AN AUSTRALASIAN JOURNAL OF PALAEONTOLOGY 3

recovered diagnostic fossils from the Cenomanian of
Venezuela show that the genus was also distributed in the
emerging Atlantic region of northern South America during
the mid-Cretaceous (Carrillo-Brice~
no et al. 2012). Smith
Woodward (1894) also described remains of ‘Portheus’aus-
tralis from the Early Cretaceous (late Albian) of Australia.
Hay (1898) designated Portheus as a junior synonym of
Xiphactinus, although Lees & Bartholomai (1987) later estab-
lished that the Australian specimens constituted a separate
genus, Cooyoo. The new Patagonian Xiphactinus occurrence
that we report here is therefore significant, because it not
only extends the stratigraphic range of the taxon to the ter-
minal-Cretaceous in South America, but also reveals a
southern Atlantic higher mid-latitude dispersal that concurs
with the cosmopolitan distribution of ichthyodectiforms and
their inferred pelagic lifestyle (Bardack 1965, Patterson &
Rosen 1977, Schaeffer & Patterson 1984, Tanimoto &
Kinkyo 2001, Forey et al. 2003, Arratia et al. 2004).
Acknowledgements
We thank M. Ezcurra (MACN) for access to collections. Our study was
supported by A. Giacchino (Universidad Maimonides), and Fundaci
on
de Historia Natural ‘Felix de Azara’. Editorial revisions by the
Alcheringa Editorial Group, together with reviews from Lionel Cavin
and Lance Grande greatly improved our manuscript.
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