Cretaceous Research 103 (2019) 104162

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Cretaceous Research
journal homepage: www.elsevier.com/locate/CretRes

Reassessment of cf. Plotosaurus from the upper Maastrichtian of Chile,

with comments on the South American distribution of halisaurine
mosasaurs

nez-Huidobro a, *, Rodrigo A. Otero b, Sergio Soto-Acun
Paulina Jime ~ a b,
c, d
Michael W. Caldwell
a
Division of Paleontology Institute of Geosciences, University of Bonn, Bonn, Germany
b
Red Paleontolόgica U.chile, Laboratorio de Ontogenia y Filogenia, Departamento de Biología, Universidad de Chile, Santiago, Chile
c
Department of Biological Sciences, University of Alberta, Edmonton, Alberta, T6G 2E9, Canada
d
Department of Earth and Atmospheric Sciences, University of Alberta, Edmonton, Alberta, T6G 2E9, Canada

a r t i c l e i n f o a b s t r a c t

Article history: The reassessment of a lower jaw fragment of a mosasaur from the upper Maastrichtian of central Chile
Received 26 April 2017 (Q.3105) indicates that the specimen does not belong to Plotosaurus, as previously suggested by Frey et al.
Received in revised form in 2016. The specimen does not show the characteristics of Plotosaurus, such as the long projection
26 April 2019
anterior to the anteriormost tooth socket of the dentary, the ventrally positioned Meckelian canal, nor
Accepted in revised form 19 June 2019
does it possess teeth with enamel coarsely striated at the base of the crown and a smooth apex. We
Available online 26 June 2019
continue in agreement to the previous assignment to Halisaurinae indet. proposed by Jime
nez-Huidobro
et al. in 2015. Even more, tooth morphologies such as the curvature pattern and the smooth and finely
Keywords:
Halisaurus
striated enamel, plus dentary characters such as the curvature of that bone, and the medially located
Plotosaurus Meckelian canal, support the assignment of the specimen Q.3105 to Halisaurus sp. Affinities of Q.3105
Upper Cretaceous with the halisaurine genus Eonatator are discarded, due to the shape in cross section and size of marginal
Chile teeth that decreases distally in the latter, traits not seen in the specimen Q.3105. This specimen repre-
sents the fourth halisaurine described from South America, and the second Maastrichtian halisaurine
from the southernmost part of the continent. This record contributes to the understanding of the broad
distribution that halisaurines reached during the Late Cretaceous.
© 2019 Elsevier Ltd. All rights reserved.

1. Introduction The oldest known mosasaur record from southern South

Marine reptiles from the Late Cretaceous of southern South
America include a diversity of elasmosaurid plesiosaurs (Gasparini
et al., 2001, 2003, 2015; O'Gorman et al., 2014; Otero et al., 2015),
some polycotylid plesiosaurs (Salgado et al., 2007; O'Gorman et al.,
2013; Soto-Acun ~ a et al., 2016), a pancheloniid (Gasparini and Biro

,
1986; Parham et al., 2014), and possible dermochelyid marine
turtles (Otero et al., 2012). In contrast, while mosasauroids are
present in southern South America, they are far less common than
plesiosaurs and turtles.
* Corresponding author.

nez-Huidobro), otero2112@
E-mail addresses: jimenezh@ualberta.ca (P. Jime
gmail.com (R.A. Otero), arcosaurio@gmail.com (S. Soto-Acun ~ a), mw.caldwell@
ualberta.ca (M.W. Caldwell).
America (Ameghino, 1893) is from the Cenomanian e Santonian
aged strata of the Mata Amarilla Formation in Argentina. An iso-
lated cervical vertebra of a halisaurine was described from the
Santonian(?) of Peru (Caldwell and Bell, 1995), and the fairly
complete skeleton of an endemic species, Eonatator coellensis, was
described from the Campanian of Colombia (P
aramo-Fonseca,
2013). Isolated teeth referred to indeterminate mosasauroids have
been described from the lower Maastrichtian of Chile (Otero et al.,
2012). Additionally, teeth of indeterminate tylosaurines and an
isolated tooth referred to a halisaurine from the upper Maas-
trichtian of Chile have been recorded (Jime
nez-Huidobro et al.,
2015). Articulated caudal vertebrae of an indeterminate mosa-
saurid were also reported from the upper Maastrichtian Dorotea
Formation in high latitudes of southernmost Chile (Soto-Acun ~a
et al., 2015). Several fragmentary specimens from the upper
Maastrichtian Jagüel Formation in Argentina, have been referred to

https://doi.org/10.1016/j.cretres.2019.06.008
0195-6671/© 2019 Elsevier Ltd. All rights reserved.
2 P. Jim
enez-Huidobro et al. / Cretaceous Research 103 (2019) 104162
the taxa Mosasaurinae indet., Mosasaurus aff. hoffmannii, Progna-
thodon sp., Plioplatecarpus sp. (Gasparini et al., 2001; Ferna
ndez
et al., 2008; Ferna
ndez and Gasparini, 2012). More recently,
Ferna
ndez and Talevi (2015) identified cranial and postcranial
material from the upper Maastrichtian Jagüel Formation as
belonging to the genus Halisaurus. This latter record complements
an emerging biogeographic pattern for the clade Halisaurinae,
showing a persistent presence of the group in South America dur-
ing the Santonian(?) of Peru, the upper Campanian of Colombia,
and the upper Maastrichtian of Chile and Argentina.
New records of mosasauroids are important, as they are not well
known from southern South America. Here we reassess the recent
description of a specimen assigned to cf. Plotosaurus, from the
upper Maastrichtian of the Quiriquina Formation of central Chile

nez-Huidobro et al. (2014) had
(Frey et al., 2016). Previously, Jime
referred the same specimen to cf. Halisaurinae indet. This opinion
was re-affirmed by Jime
nez-Huidobro et al. (2015), although no
generic assignment was provided (i.e., Eonatator, Halisaurus, Phos-
phorosaurus). As the recent description by Frey et al. (2016) is
different from previous identifications, noting the presence of a
plotosaur in South America, we think it is important to reassess the
anatomy of the specimen, in order to properly differentiate be-
tween the two such widely differing conclusions (i.e., mosasaurine
versus halisaurine). We provide here a detailed comparison of the
specimen with Plotosaurus bennisoni (Camp, 1942; Lindgren et al.,
Fig. 1. Map of Central Chile, the red point indicates Cocholgüe, the locality where the specimen Q.3105 was recovered. (For interpretation of the references to color in this figure
legend, the reader is referred to the Web version of this article.)
2008; Leblanc et al., 2013), as well as Halisaurus arambourgi
(Bardet et al., 2005).
2. Locality and geological settings
The material was collected in Cocholgüe, 25 km north of Con-
cepcio
n, in the Biobío Region, central Chile (Fig. 1). The specimen
was found in a transported concretion by a local collector, making it
impossible to determine its precise stratigraphic provenance. The
coastline outcrops of Cocholgüe consist mostly of sedimentary
rocks of the Quiriquina Formation (Biro
-Bago
czky, 1982). The upper
section of this unit is composed of fine sandstones-siltstones, bio-
turbated beds, and limestones with concretionary nodules. Con-
cretions often include invertebrate remains; however, different
siltstone beds do bear occasional vertebrates (Sua
rez et al., 2003;
Otero et al., 2014; Parham et al., 2014). The provenance of the
studied specimen (within a concretion) is likely from the upper
section of the Quiriquina Formation. The unit is restricted to the
upper Maastrichtian based on ammonite biostratigraphy (Salazar
et al., 2010).
3. Materials and methods
After discarding other halisaurine genera, the specimen Q.3105
was compared with Plotosaurus bennisoni and Halisaurus
 P. Jim
enez-Huidobro et al. / Cretaceous Research 103 (2019) 104162
arambourgi, in order to reassess its taxonomic assignment. Photo-
graphs were taken with a Canon EOS t3i camera and Canon macro
lens EF-S 60 mm. Images were edited with Adobe Photoshop CS6
and Adobe Illustrator CS6, using drawing tablet software and the
hardware tablet Wacom Bamboo CTH-470. Tooth cross sectional
ratios were measured as the ratio between the mesiodistal and the
labiolingual lengths of the sockets.
Institutional AbbreviationsdLACM, Los Angeles County Museum,
Los Angeles, California, USA; Q, Museo Geolo
gico Profesor Lajo

s

, Universidad de Concepcio
Biro
n, Chile; UALVP, University of
Alberta Laboratory of Vertebrate Palaeontology, Edmonton, AB,
Canada; UCMP, Museum of Paleontology University of California,
Berkeley, California, USA.
4. Systematic palaeontology
Squamata Oppel, 1811
Mosasauridae Gervais, 1852
Halisaurinae Bardet et al., 2005
Halisaurus Marsh, 1869
Halisaurus sp.
Type species. Halisaurus platyspondylus Marsh, 1869. Maastrichtian
of New Jersey, USA (Holmes and Sues, 2000).
Referred specimen. Q.3105 corresponds to a block with fragments of
left and right dentaries and a fragmented right splenial (Fig. 2).
Geographic/Stratigraphic provenance. Cocholgüe, Quiriquina For-
mation, central Chile, indeterminate levels. Upper Maastrichtian.
Fig. 2. Picture and schematic drawing of the specimen Q.3105: in A, lateral view; and B, latero-occlusal view; C, Schematic drawing of Q.3105 in lateral view; and D, schematic
drawing of Q.3105 in latero-occlusal view. Scale bars equals 5 cm. Anatomical abbreviations: 1e15, numbers position of the preserved teeth (numbers do not represent actual tooth
positions as measured from the tip of the snout, but only as preserved); alr, alveolous of replacement tooth; for, foramina; lde, left dentary; lsp, left splenial; rsp, right splenial; mc,
Meckelian canal; rde, right dentary.
3
Synonymy. Mosasauridae indet (Su
arez, 1990)
Halisaurinae indet (Jime
nez-Huidobro et al., 2015)
cf. Plotosaurus sp. (Frey et al., 2016)
5. Description and comparisons
Small-sized mosasauroid, preserved in a block that includes a
fragment of the lower jaw: left and right dentaries, and the right
splenial (Fig. 2). The left dentary is exposed in lateral view, while
the right dentary and right splenial are visible in medial view. The
right dentary, which is posteriorly incomplete, bears 15 alveoli,
whereas the left one, anteriorly and posteriorly incomplete, bears
13. In lateral view, both dentaries show straight dorsal and
ventral margins. The right dentary seems to be anteriorly almost
complete, as it gently curves down at the anterodorsal margin of
the dentary, although the tip is broken, missing the anteriormost
projection. The medial face of the bone is slightly concave. The
specimen Q.3105 (Figs. 2, 3A) and specimens of Plotosaurus ben-
nisoni (LACM2945 and UCMP 32778; Fig. 3C, D) show noticeable
and important differences. All known plotosaur dentaries (Camp,
1942; Lindgren et al., 2008; LeBlanc et al., 2013) possess a
distinctive anterior edentulous extension/process of the dentary
with a pronounced process anterior to the first tooth position that
projects downward (Fig. 3 D). The process is not figured in the
Plotosaurus drawing from Frey et al. (2016; Fig. 3). There is no
such anterior process visible on the right dentary of Q.3105,
although the anteriormost tip is absent, it is still possible to see
4 P. Jim
enez-Huidobro et al. / Cretaceous Research 103 (2019) 104162

Fig. 3. Comparison of the specimen Q.3105 with Halisaurus arambourgi and Plotosaurus bennisoni. A, specimen Q.3105, scale bar equals 5 cm; B, Halisaurus arambourgi UALVP 56123,
scale bar equal to 5 cm; C, Plotosaurus bennisoni LACM2945, scale bar equal to 10 cm; D, Plotosaurus bennisoni UCMP 32778, scale bar equal to 5 cm.

that the anterior border projects downwards, suggesting lack of
the anteriormost process. The left dentary of Q.3105 also presents
a gentle convexity marked by a horizontal line of trigeminal nerve
foramina similar to H. arambourgi, but in contrast to the straight
shape described by Plotosaurus dentaries.
In H. arambourgi the Meckelian canal is situated in the middle of
the medial face, and narrows anteriorly, becoming wider posteri-
orly. The Meckelian canal of Q.3105 is deep and situated in the
middle of the medial face of dentary, and is narrow anteriorly,
becoming wider posteriorly, as in Halisaurus arambourgi (UALVP
56123; Fig. 3B). In Plotosaurus the Meckelian canal is exposed
medially from the posterior to midpoint of the dentary, but then
descends sharply below the seventh tooth position to a ventral
position and opens ventrally, not medially as in Q.3105 (Fig. 3C, D).
The right splenial is also preserved in a Q3105, although
incomplete. The left splenial is only visible at its dorsal margin,
behind the left dentary. In lateral view, the right splenial is a long
and slender element that becomes narrower anteriorly, while it is
posteriorly wider. The posterior margin is not preserved. The
ventral margin of the bone is nearly straight, while the dorsal
margin presents some curvature.
The teeth preserved in both dentaries are sharp, with the base of
the crowns straight until about half of the crown length, and the
apex posteriorly curved. The teeth are rounded in cross-section. In
dorsal view, the alveoli in both dentaries are perfectly rounded. The
second, fourth, seventh, ninth and twelfth teeth of the right dentary
present nearly circular alveoli for replacement teeth in their
respective posterolingual faces. The replacement teeth appear
lingual with respect to the old teeth. The teeth of Q.3105 also share
the pattern of curvature of Halisaurus arambourgi (Fig. 4A, B). The
crown is nearly straight at the base, and the apex abruptly curves
posteriorly about the mid-length of the crown (Bardet et al., 2005).
The dentary of Plotosaurus possesses slender and nearly straight
mesial teeth; they become shorter and more robust distally
(Figs. 3C, D, 4C), resembling those of Eremiasaurus heterodontus
(Leblanc et al., 2012), whereas the teeth in Q.3105 show a different
pattern of curvature. The enamel of Q.3105 is ornamented with fine
striations from the apex to the base of the crown (Fig. 4A),
consistent with the teeth of H. arambourgi (Fig. 4B). On the other
hand, in Plotosaurus the enamel is coarsely striated at the base of
the crown, with a smooth apex (Fig. 4C). Another dental character
shared by Q.3105 and H. arambourgi is the rounded tooth cross-
section and associated rounded alveoli (Fig. 4D, E). The average
cross-sectional ratios are 0.924 for Q.3105, and 0.916 for
H. arambourgi.
6. Biogeography of Late Cretaceous Austral mosasaurs
AntarcticadLate Cretaceous mosasaur records from the South-
ern Hemisphere are scarce in comparison with those from the
 P. Jim
enez-Huidobro et al. / Cretaceous Research 103 (2019) 104162 5

Fig. 4. Comparison of tooth morphologies between: A, specimen Q.3105, scale bar equal to 1 cm; B, Halisaurus arambourgi UALVP 56123, scale bar equals 0.5 cm; C, Plotosaurus
bennisoni LACM2945, scale bar equal to 1 cm. Occlusal views of dentaries: D, specimen Q.3105, scale bar equal to 2 cm; and E, Halisaurus arambourgi UALVP 56123, scale bar equal to
2 cm.

Northern Hemisphere. The most diverse assemblage is known from
the upper Campanian e upper Maastrichtian of Antarctica. The
oldest Antarctic records include a well preserved skull recovered
from upper Campanian beds of the Santa Marta Formation in James
Ross Island (Olivero et al., 1986), first referred to a new genus and
species, Lakumasaurus antarcticus (Novas et al., 2002), and later
synonymized with Taniwhasaurus Hector (1874) and reassessed as
Ta. antarcticus (Martin and Fern

ndez and Martin,
andez, 2007; Ferna
2009). The same taxon has been recorded in the lower Maas-
trichtian of the Snow Hill Island Formation from Vega Island,
6 P. Jim
enez-Huidobro et al. / Cretaceous Research 103 (2019) 104162
Antarctica (Ferna
ndez y Gasparini, 2012). The upper Maastrichtian
mosasaur records from Antarctica came from the Lo
pez de Berto-
dano Formation (Rinaldi et al., 1978), exposed on Marambio
(¼Seymour) Island. Material has been referred to Plioplatecarpus
sp., Leiodon sp., Mosasaurus sp., Mosasaurus aff. hoffmannii, Mosa-
saurus cf. lemonnieri, and cf. Moanasaurus sp., as well as indeter-
minate tylosaurine remains (Martin, 2006). Reguero et al. (2015)
added the presence of Prognathodon sp., Mosasaurus sp. and Plio-
platecarpus sp. from upper levels of the Lo
pez de Bertodano For-
mation, Antarctica.
New ZealanddA well preserved assemblage, equalling the di-
versity known from Antarctica, comes from the Late Cretaceous of
New Zealand: Mosasaurus mokoroa Welles and Gregg 1971 from the
middle Campanian of the Conway Formation, Taniwhasaurus oweni
Hector 1874 from the upper Campanian of the Conway Formation,
Moanasaurus mangahouangae Wiffen 1980, and Rikisaurus tehoensis
Wiffen 1990 from the Campanian e Maastrichtian of the Maun-
gataniwha Sandstones, and finally, Prognathodon waiparaensis
Welles and Gregg 1971 from the upper Maastrichtian of the Conway
Formation.
South AmericadContrary to the rich record of mosasaurs from
Antarctica and New Zealand, Upper Cretaceous South American
records are sparse. Ameghino (1893) described isolated teeth
from Cenomanian e Santonian units of the Mata Amarilla For-
mation outcropping at Lago Argentino, Argentina, referring them
to 'Leiodon argentinus' (nomen dubium); however, the material is
currently lost. A cervical vertebra from the Santonian(?) of the
Vivia
n Formation, Peru, was referred to Halisaurus sp. by Caldwell
and Bell (1995). Gasparini et al. (2001) described vertebrae
referred to indeterminate Mosasaurinae from the upper Maas-
trichtian of the Jagüel Formation, southern Mendoza, Argentina.
From the same unit, cranial and postcranial material referred to
Mosasaurus aff. hoffmannii was described, in addition to a second
indeterminate mosasaurine that comprises fragmentary mandi-
bles, teeth and vertebrae (Ferna
ndez et al., 2008); a single iso-
lated tooth has been referred to Prognathodon (Ferna
ndez et al.,
2008; Fern
andez and Gasparini, 2012), and a tooth and cervical,
dorsal and caudal vertebrae have been referred to Plioplatecarpus
sp. (Ferna
ndez et al., 2008; Ferna
ndez and Gasparini, 2012).
Pa
ramo-Fonseca (2013) described a new halisaurine species,
Eonatator coellensis, from the Campanian of Colombia. Later,
Ferna
ndez and Talevi (2015) described a right quadrate, cervicals
and anterior dorsal vertebrae of a single specimen from the upper
Maastrichtian of the Jagüel Formation, identified as Halisaurus sp.,
making this the third record of the clade Halisaurinae in South
America.
The known mosasaur record from Chile is represented by
material from the Arauco Basin as well as the Magallanes
(¼Austral) Basin. Finds from the Arauco Basin include isolated
teeth of indeterminate mosasaurids from the lower Maastrichtian
Estratos de la Quebrada Municipalidad in north-central Chile
(Otero et al., 2012). In addition, large dorsal vertebrae of inde-
terminate mosasaurids have been recovered from two localities in
central Chile, from units of upper Maastrichtian age belonging to
the Quiriquina Formation (Jime
nez-Huidobro et al., 2015). Two
teeth referred to tylosaurines have also been recovered from the
same unit (Sua
rez et al., 2003; Jime
nez-Huidobro et al., 2014,
2015). Additionally, the specimen studied here (Q.3105), as well
as an isolated tooth also from the upper Maastrichtian of the
Quiriquina Formation, have been previously referred to indeter-
minate halisaurines (Jime
nez-Huidobro et al., 2014, 2015). Finally,
the only record from the Magallanes Basin consists of four arti-
culated caudal vertebrae, recognized as an indeterminate mosa-
saurid, recovered from upper Maastrichtian units of the Dorotea
~ a et al., 2015; Jime
Formation (Soto-Acun
nez-Huidobro et al.,
2015).
7. Discussion
Frey et al. (2016) assigned the specimen Q.3105 to cf. Plotosaurus
based on the following characters: (1) the more distally preserved
teeth are abruptly re-curved in comparison to more rostrally
crowns, character diagnostic for Plotosaurus according to Camp
(1942). The dentary of Plotosaurus bears slender and nearly
straight mesial teeth that become shorter and more robust distally,
while Q.3105 shows poorly differentiated teeth in both shape and
size, resembling those described for Halisaurus arambourgi by
Bardet et al. (2005), but with no carinae. (2) The finely striations
present in Plotosaurus and the Quiriquina specimen crowns, in
combination with absence of carinae. However, our observations
show that the enamel of Q.3105 is ornamented with fine striations
from the apex to the base of the crown, seen as fine grooves on the
lingual surface (Fig. 4A). On the other hand, the enamel of a Plo-
tosaurus tooth is finely striated only in the apex, while the base of
the crown is coarsely striated with a wrinkled texture (Fig. 4C). The
analysis of the specimen Q.3105, and comparisons to both Ploto-
saurus bennisoni and Halisaurus arambourgi, allow us to step
beyond than Jime
nez-Huidobro et al. (2015) who assigned Q.3105
to Halisaurinae indet., here assigning the specimen at generic level.
The current reassessment of Q.3105 as Halisaurus sp. represents
the third record of this genus in South America, and the fourth
occurrence of halisaurines in the continent. Even with only three
known records in South America, its biogeographic distribution is
remarkably broad across the continent, from the epicontinental sea
of Venezuela-Colombia (Putumayo Basin) during the Campanian, to
both the southwestern Atlantic as well as in the southeastern Pa-
cific during the upper Maastrichtian.
Six nominal halisaurine species are recognized for the three
genera: Halisaurus platyspondylus Marsh 1869; Phosphorosaurus
ortliebi Dollo 1889; Eonatator sternbergii (Wiman, 1920);
H. arambourgi Bardet and Pereda-Suberbiola (in Bardet et al., 2005);
E. coellensis Pa
ramo-Fonseca 2013; and P. ponpetelegans Konishi
et al. 2015 (Table 1). Their occurrences include North America,
Morocco, Japan, and South America, showing a cosmopolitan
paleodistribution of the group, at least during the Campanian e
Maastrichtian (Fig. 5).
Halisaurines are known from the Santonian (Caldwell and Bell,
1995), retaining a sub-equatorial distribution until the Campa-
nian, and reaching high latitudes during the Maastrichtian. Such
distributional patterns support possible faunal interchanges of
marine reptiles faunas along the margins of both the Pacific and the
Table 1
Comparison of temporal and geographic distribution of all described taxa from the
subfamily Halisaurinae, and the mosasaurine Plotosaurus bennisoni.
Stage Geographic distribution
Halisaurus middle e upper Eastern North America.
platyspondylus Maastrichtian
Halisaurus arambourgi Maastrichtian Morocco and Middle East.
Halisaurus sp. Santonian Peru.
upper Maastrichtian Southern Chile and
Argentina
Eonatator sternbergi upper Santonian e lower North America and
Campanian Sweden.
Eonatator coellensis upper Campanian Colombia.
Phosphorosaurus ortliebi Maastrichtian Belgium.
Phosphorosaurus lowermost Maastrichtian Northern Japan.
ponpetelegans
Plotosaurus bennisoni Maastrichtian California, North
America.
 P. Jim
enez-Huidobro et al. / Cretaceous Research 103 (2019) 104162 7

Fig. 5. Plate tectonic reconstructions during the Late Cretaceous, indicating the global occurrences of halisaurines (i.e., Eonatator and Phosphorosaurus) in black, the genus Halisaurus in
red, and the genus Plotosaurus in green. A, Santonian e Campanian; B, Campanian e middle Maastrichtian; C, late Maastrichtian. Sources of information are summarized on Table 1.
Paleogeographic reconstructions based on Blakey (2008). (For interpretation of the references to color in this figure legend, the reader is referred to the Web version of this article.)
8 P. Jim
enez-Huidobro et al. / Cretaceous Research 103 (2019) 104162
Atlantic. Therefore, the Austral marine reptile diversity not only
includes typically Weddellian representatives (e.g. mosasaurine
mosasaurs such as Moanasaurus and Rikisaurus, and elasmosaurid
plesiosaurs such as Tuarangisaurus, Vegasaurus as well as aristo-
nectines), but also, elements previously restricted to the Northern
Hemisphere in both Pacific and Atlantic oceans. Pacific elements
present in both hemispheres are so far represented by pan-
cheloniids of the genus Euclastes (Parham et al., 2014 and refer-
ences therein), cf. dermochelyids (Otero et al., 2012), and likely by
elasmosaurids with close morphological affinities (e.g., CM Zfr 115
referred to 'Mauisaurus haasti' by Hiller et al., 2005, compared to
Hydrotherosaurus alexandrae Welles, 1943). Tylosaurine mosasaurs
are also present in the Western Interior Seaway as well as in New
Zealand (Welles and Gregg, 1971), Antarctica (Novas et al., 2002;
Martin and Fern
andez, 2007) and Chile (Jime
nez-Huidobro et al.,
2014, 2015). On the other hand, an Atlantic interchange pattern
emerges with the presence of Euclastes from the Maastrichtian of
Angola (Vineyard et al., 2009), South America (Parham et al., 2014)
and Morocco (Jalil et al., 2009), and South America (Otero et al.,
2015), and now by the common presence of Halisaurus in South
America and Morocco (Bardet et al., 2005; Ferna
ndez and Talevi,
2015).
Under this scenario, the presence of Halisaurus sp. in the Quir-
iquina Formation is biogeographically plausible (not just anatomi-
cally). The genus Halisaurus seems to present a global distribution,
as seen in several mosasauroid taxa up to the generic level (Bardet
et al., 2014), contrary to the restricted occurrence of Plotosaurus so
far known only in North America, more than 10,000 km north from
the Arauco Basin, and without any geographically intermediate
record of the genus.
8. Conclusions
Closer comparisons between Q.3105 from the upper Maas-
trichtian of central Chile, and the type and referred materials of
Plotosaurus bennisoni from California, indicate that there are no
important similarities, beyond plesiomorphies, that permits
assignment of the specimen to Plotosaurus. On the other hand,
several similarities are present when compared to halisaurines,
especially with Halisaurus arambourgi. Our identification of the
higher-level affinities of Q.3105 indicates they present affinities to
halisaurines, not mosasaurines, specifically the genus Halisaurus sp.
This makes Q.3105 the third record of this genus in South America,
adding to its previous occurrence in the Santonian(?) of Peru
(Caldwell and Bell, 1995) and in the upper Maastrichtian of
Argentina (Ferna
ndez and Talevi, 2015). The present record of
Halisaurus in the upper Maastrichtian of the Quiriquina Formation
evidences a bi-oceanic distribution of the genus prior to the
extinction of mosasauroids at the K/Pg Boundary.
Acknowledgments
We would like to thank R. Blakey, Colorado Plateau Geosystems
for the paleomap used for the production of Fig. 5. We thank G.
Flores and L.A. Quinzio for access to the specimen. We are also
thankful of A. Schulp and V. Fischer for revisions and comments
that improved the manuscript. A photograph of a plotosaur tooth
was provided by A. LeBlanc. We thank the Proyecto Anillo en

n en Ciencia y Tecnología (PIA, Conicyt) ACT172099.
Investigacio
Natural Sciences and Engineering Research Council of Canada
(NSERC) Discovery Grant (238458-01), NSERC Accelerator Grant,
and Chair's Research Allowance to M.W.C.; Alexander von Hum-
boldt Postdoctoral Fellowship to P.J.H.
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Appendix A. Supplementary data
Supplementary data to this article can be found online at https://doi.org/10.
1016/j.cretres.2019.06.008.