Journal of Vertebrate Paleontology

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A new carcharodontosaurian theropod from the

Lusitanian Basin: evidence of allosauroid sympatry
in the European Late Jurassic

Elisabete Malafaia , Pedro Mocho , Fernando Escaso & Francisco Ortega

To cite this article: Elisabete Malafaia , Pedro Mocho , Fernando Escaso & Francisco Ortega
(2020) A new carcharodontosaurian theropod from the Lusitanian Basin: evidence of allosauroid
sympatry in the European Late Jurassic, Journal of Vertebrate Paleontology, 40:1, e1768106, DOI:
10.1080/02724634.2020.1768106

To link to this article: https://doi.org/10.1080/02724634.2020.1768106

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Journal of Vertebrate Paleontology e1768106 (8 pages)
© 2020 by the Society of Vertebrate Paleontology
DOI: 10.1080/02724634.2020.1768106

ARTICLE

A NEW CARCHARODONTOSAURIAN THEROPOD FROM THE LUSITANIAN BASIN:

EVIDENCE OF ALLOSAUROID SYMPATRY IN THE EUROPEAN LATE JURASSIC

ELISABETE MALAFAIA, *,1,2,3 PEDRO MOCHO, 1,2,3,4 FERNANDO ESCASO, 2,3 and FRANCISCO ORTEGA 2,3
1
Instituto Dom Luiz, Universidade de Lisboa, Edifício C6, Campo Grande, 1749-016 Lisboa, Portugal, efmalafaia@fc.ul.pt;
p.mochopaleo@gmail.com;
2
Grupo de Biología Evolutiva, Facultad de Ciencias, Universidad Nacional de Educación a Distancia, Paseo de la Senda del Rey 9,
28040 Madrid, Spain, fescaso@ccia.uned.es; fortega@ccia.uned.es;
3
Laboratório de Paleontologia e Paleoecologia, Sociedade de História Natural, Polígono Industrial do Alto do Ameal, Pav. H02 e H06,
2565-641, Torres Vedras, Portugal;
4
The Dinosaur Institute, Natural History Museum of Los Angeles County, 900 Exposition Blvd., Los Angeles, California 90007, U.S.A.

ABSTRACT—Carcharodontosaurian allosauroids were temporally restricted to the Cretaceous, being known from all land
masses with the exception of Antarctica. In addition to Veterupristisaurus from Tanzania, exceptions to this distribution have
been reported recently, consisting on fragmentary materials from Upper Jurassic strata of China, Germany, and Portugal.
Here, we propose a new Late Jurassic carcharodontosaurian taxon, Lusovenator santosi, gen. et sp. nov. based on the
reevaluation of previously described specimens from the Lusitanian Basin, Portugal. The performed phylogenetic analysis
recovered Lusovenator santosi as an early branching carcharodontosaurian allosauroid diagnosed by an exclusive combination
of characters, including three autapomorphic features: (1) large recesses in neural arch of anterior dorsal vertebrae; (2) well-
developed and continuous longitudinal laminae extending from the tip of the prezygapophyses to the distal end of the
postzygapophyses in mid-caudal vertebrae; and (3) supraacetabular crest of ilium forming a prominent ventrolaterally
projecting shelf. Lusovenator santosi is the oldest carcharodontosaurian allosauroid yet discovered from Laurasia and supports
unequivocally the hypothesis of a pre-Cretaceous scenario for the radiation of the clade. The identification of this taxon
highlights the high diversity of medium- to large-bodied theropods in the later part of the Late Jurassic of the Iberian
Peninsula. Carcharodontosauria is not yet known in correlative levels of the North American Morrison Formation, and the
existence of contacts after the late Tithonian between these landmasses could explain the distribution of this clade and other
dinosaur groups present in the Iberian Jurassic and in the North American Lower Cretaceous.

http://zoobank.org/urn:lsid:zoobank.org:pub:0EEC830B-26EC-484B-860E-7777C1A1D6FA

SUPPLEMENTAL DATA—Supplemental materials are available for this article for free at www.tandfonline.com/UJVP

Citation for this article: Malafaia, E., P. Mocho, F. Escaso, and F. Ortega. 2020. A new carcharodontosaurian theropod from the
Lusitanian Basin: evidence of allosauroid sympatry in the European Late Jurassic. Journal of Vertebrate Paleontology. DOI:
10.1080/02724634.2020.1768106.

INTRODUCTION Ceratosaurus (e.g., Malafaia et al., 2015), Torvosaurus (e.g., Hen-

The Portuguese Upper Jurassic record of theropod dinosaurs
comprises members of deeply nested clades that are abundant
and diverse in Cretaceous faunas. It includes Carcharodonto-
sauria (Malafaia et al., 2019), Tyrannosauroidea (Rauhut,
2003a), and a great diversity of maniraptorans (Zinke and
Rauhut, 1994; Zinke, 1998; Hendrickx and Mateus, 2014a; Mala-
faia et al., 2017b). The Portuguese specimens represent some of
the earliest evidence of these clades in Laurasia and have signifi-
cant implications for understanding the paleobiogeographic
context and dynamics of Late Jurassic theropod fauna in the
peri-Atlantic realm. The Upper Jurassic theropod record in the
Lusitanian Basin is mostly composed of medium- to large-sized
taxa with a trans-Atlantic distribution, such as the genera
*Corresponding author.
This article was originally published with errors, which have now been
corrected in the online version. Please see Correction (https://doi.org/10.
1080/02724634.2020.1810451).
Color versions of one or more of the figures in the article can be found
online at www.tandfonline.com/ujvp.
drickx and Mateus, 2014b), and Allosaurus (e.g., Pérez-Moreno
et al., 1999; Mateus et al., 2006). However, this record also
includes some exclusive taxa interpreted as related to Jurassic
forms from both Laurasian (e.g., Lourinhanosaurus antunesi;
Mateus, 1998) and Gondwanan (e.g., Carcharodontosauria; Mala-
faia et al., 2019) landmasses. The combination of shared and
endemic forms is compatible with a model of incipient vicariance
pattern for the Late Jurassic dinosaur faunas of North American
and Iberian domains (e.g., Ortega et al., 2017). This vicariant
pattern was possibly related to the establishment of geographic
barriers (e.g., opening of the North Atlantic Ocean) that would
enforce an increase in continent-scale endemism during the
Late Jurassic and the Early Cretaceous (e.g., Benson et al., 2013).
It has been assumed that Carcharodontosauria has a ghost
lineage extending back to the Late Jurassic (Brusatte et al.,
2008). More recently, some isolated elements from Upper Jurassic
strata in different localities have been identified to this clade,
including a specimen from the Upper Jurassic of the Lusitanian
Basin (Malafaia et al., 2019), some isolated teeth, caudal ver-
tebrae, and an ilium from the Tendaguru Formation (Rauhut,
2011; Carrano et al., 2012), some teeth from the Middle–Upper

Published online 10 Jul 2020

 Malafaia et al.—New carcharodontosaurian dinosaur from Portugal (e1768106-2)
Jurassic Shishugou Formation of China (Han et al., 2011), and
possibly some teeth from the Lower Saxony Basin of Germany
(Gerke and Wings, 2016). Here, a new taxon, Lusovenator
santosi, gen. et sp. nov., related to Carcharodontosauria, is
defined, and the phylogenetic position of the type and referred
specimens is presented. The new taxon supports the previously
referred evidence for the presence of carcharodontosaurian ther-
opods in Portugal (Malafaia et al., 2019). It is the first taxon of the
group defined in the Jurassic of Laurasia and extends the fossil
record of this clade already represented in the Lower Cretaceous
of Europe (e.g., Brusatte et al., 2008; Ortega et al., 2010).
Institutional Abbreviations—MB.R, Museum für Naturkunde,
Berlin, Germany; ML, Museum of Lourinhã, Lourinhã, Portugal;
NCSM, North Carolina Museum of Natural Sciences, Raleigh,
U.S.A.; SHN, Sociedade de História Natural, Torres Vedras,
Portugal.
MATERIALS AND METHODS
A phylogenetic analysis of the specimens here assigned to the
new species Lusovenator santosi was performed using the data
matrix of Carrano et al. (2012), including modifications proposed
by Coria and Currie (2016) and Zanno and Makovicky (2013), as
well as the addition of seven new morphological characters and
four taxa. A modification of the description for character 278, con-
sisting of elimination of the second state (because most Neothero-
poda taxa have ilia with postacetabular length that is doubled the
ischial peduncle length) proposed by Rauhut et al. (2016), was
also included. Some characters were rescored for Lourinhano-
saurus antunesi Mateus, 1988, based on review of the holotype
(see Appendix S1 in Supplemental Data). A left femur (ML 555)
previously assigned to Lourinhanosaurus by Antunes and Mateus
(2003) was removed from the scoring of this taxon because it was
collected from a different site and does not share any diagnostic
characters with the holotype; ML 555 has a femoral head that pro-
jects dorsomedially, which is a feature that has been interpreted as a
synapomorphy of Carcharodontosauria (Brusatte et al., 2008;
Benson et al., 2010). Considering this hypothesis, the fossil record
of carcharodontosaurs in the Lusitanian Basin extends from the
Kimmeridgian to the upper Tithonian (see Fig. S1).
The data matrix, with 358 characters and 65 taxa, was analyzed
through TNT 1.1 (Goloboff et al., 2008). The 358 characters were
scored using Mesquite 2.72 (Maddison and Maddison, 2009) and
were considered to be unordered and equally weighted in the per-
formed analyses. Herrerasaurus Reig, 1963, was selected as the out-
group taxon. Given the large size of the matrix and following the
procedures executed in the analyses of Carrano et al. (2012), a heur-
istic search using the ‘New Technology Search’ options was per-
formed. These included noncollapsing rules and the default
settings for sectorial, ratchet, tree drift, and tree fusion, using a
driven search that stabilized consensus twice with a factor of 25. Sub-
sequently, the resulting most parsimonious trees (MPTs) were sub-
jected to tree bisection and reconnection (TBR) branch swapping.
The strict consensus using all trees was then calculated.
For the phylogenetic analysis, the holotype and the referred
material were coded separately. When the two specimens are
included, the resulting consensus tree places them within Allosauria,
but the resolution of the clade is low, with most taxa forming a large
polytomy. When the referred specimen (SHN.019) is excluded, there
is a great improvement in the resolution within Allosauria. An analy-
sis with both specimens codified together was also performed in order
to verify the differences in the results between the two analyses, but
the typology of the consensus tree is the same.
SYSTEMATIC PALEONTOLOGY
THEROPODA Marsh, 1881
TETANURAE Gauthier, 1986
AVETHEROPODA Paul, 1988
ALLOSAUROIDEA Marsh, 1878
CARCHARODONTOSAURIA Benson, Carrano, and
Brusatte, 2010
LUSOVENATOR SANTOSI, gen. et sp. nov.
(Figs. 1, 2A, C, D)
Holotype—SHN.036, a partial postcranial skeleton preserving
odontoid, atlantal intercentrum, a cervical vertebra (previously mis-
interpreted as the articulated axial intercentrum and axis; Malafaia
et al., 2017a), isolated cervical neural spines, dorsal vertebrae, frag-
ments of sacral vertebrae, caudal vertebrae, chevrons, fragments of
cervical and dorsal ribs, right ilium, both pubes, and ischia (Fig. 1).
See Malafaia et al. (2017a) for a full description of the specimen.
SHN.036 is interpreted as a juvenile individual because it has open
neurocentral sutures on all preserved posterior dorsal, sacral, and
anterior caudal vertebrae, which is a criterion traditionally used to
assess ontogeny in archosaurs (e.g., Brochu, 1996; Irmis, 2007).
Etymology—The generic name is composed from the Latin
words Luso, referring to Lusitania, the province in Roman Hispa-
nia related to the current Portugal, and venator, from the Latin
word for hunter; the specific name santosi after José Joaquim
dos Santos, who found and collected the holotype.
Locality and Horizon—Praia de Valmitão, in the locality of
Ribamar and municipality of Lourinhã (central-west region of
the Lusitanian Basin, Portugal). Praia da Amoreira-Porto Novo
Formation (upper Kimmeridgian) (see Fig. S1).
Diagnosis—A carcharodontosaurian theropod diagnosed by the
following autapomorphies (the diagnosis is only based on the type
specimen, SHN.036): large recesses in neural arch of anterior
dorsal vertebrae (Fig. 1C); well-developed and continuous longi-
tudinal laminae extending from the tip of the prezygapophyses
to the distal end of the postzygapophyses in mid-caudal vertebrae
(Fig. 1H); and supraacetabular crest of ilium forming a prominent
ventrolaterally projecting shelf (Fig. 1I). Additionally, Lusovenator
differs from other carcharodontosaurs in the following unique
combination of features: absence of ventral keel on anterior
dorsal vertebrae (Fig. 1D); presence of hyposphene-hypantrum
articulation in at least the anterior-most caudal vertebrae (Fig.
1G); presence of a lateral lamina projecting from posterior articu-
lar facet to base of caudal rib in mid-caudal vertebrae (Fig. 1H);
mid-caudal vertebrae with spinoprezygapophyseal lamina project-
ing from the medial surface of the prezygapophysis (Fig. 1H); and
almost flat ventral surface of the pubic boot (Fig. 1J).
Referred Material—SHN.019, a partial skeleton represented by a
series of articulated caudal vertebrae and an almost complete right
pes collected from another locality and horizon. See Malafaia et al.
(2019) for a full description of the specimen. Although SHN.019 was
found in a different locality and formation (around 8 Ma younger
than SHN.036), this specimen is tentatively referred to Lusovenator
santosi based on the presence of a shared feature proposed as diag-
nostic of this taxon: the presence of longitudinal laminae extending
from the tip of the prezypapophyses to the distal end of the postzy-
gapophyses in mid-caudal vertebrae. Besides, both specimens also
share the presence of a distinct lateral lamina in mid-caudal ver-
tebrae, which is a feature unknown in other theropods from the
Late Jurassic of the Lusitanian Basin.
SHN.019 was collected at Praia de Cambelas, in the locality of
São Pedro da Cadeira and municipality of Torres Vedras (central-
west region of the Lusitanian Basin, Portugal) and comes from
sediments belonging to the Freixial Formation (upper Tithonian).
RESULTS
The first search of the performed cladistics analysis, based on
the ‘New Technology Search’ option, retained 77 MPTs with
length of 1,084, and the following ‘Traditional Search’ produced
 Malafaia et al.—New carcharodontosaurian dinosaur from Portugal (e1768106-3)

FIGURE 1. Lusovenator santosi, gen. et sp. nov., SHN.036, holotype, selected skeletal remains. A, odontoid and atlantal intercentrum in anterior view.
B, cervical vertebra in right lateral view. C, D, anterior dorsal vertebra in C, right lateral and D, ventral views. E, sacral neural arch in posterior view. F,
neural arch of anterior caudal vertebra in left lateral view. G, H, middle caudal vertebra in G, dorsal and H, left lateral views. I, right ilium in lateral view;
J, K, pubes in J, left lateral and K, ventral views. L, silhouette showing preserved elements of SHN.036. A plus sign (+) indicates a diagnostic character
and an asterisk (*) indicates an autapomorphy of Lusovenator santosi, gen. et sp. nov. Abbreviations: acdl, anterior centrodiapophyseal lamina; act,
acetabulum; at in, atlantal intercentrum; bf, brevis fossa; cpf, cupedicus fossa; isp, ischial peduncle; mec, medial crest; ms, medial symphysis; ncs, neu-
rocentral suture; ns, neural spine; od, odontoid; pcdl, posterior centrodiapophyseal lamina; posdf, postzygapophyseal spinodiapophyseal fossa; poz,
postzygapophysis; pp, parapophysis; prz, prezygapophysis; pup, pubic peduncle; spof, spinopostzygapophyseal fossa; tp, transverse process. Scale
bars equal 25 mm (A), 50 mm (B–H), 100 mm (I–K), and 1 m (L).

5,832 MPTs with length of 1,084 steps. The consensus tree
recovers Lusovenator santosi as an early branching carcharo-
dontosaur within Allosauroidea (Fig. S1). The consensus tree
has a consistency index (CI) of 0.397 and a retention index
(RI) of 0.684 and places Lusovenator santosi in an unresolved
carcharodontosaurian polytomy with Siats meekerorum and
the clade including neovenatorid and carcharodontosaurid
carcharodontosaurs. The consensus tree topology is similar to
those obtained by previous works (Carrano et al., 2012; Coria
and Currie, 2016). Only the early branching position of Siats
differs from that obtained by Zanno and Makovicky (2013),
which places this taxon within the neovenatorid carcharodonto-
saurian group. Synapomorphies that clearly nested Lusovenator
within carcharodontosaurian allosauroids include the presence
of a ventral ridge on the ventral surface of anterior caudal ver-
tebrae (character 202), a strongly developed ridge on the
medial surface of the ilium adjacent to the preacetabular notch
(character 273) (Fig. 2A, B), and the peg-and-socket mor-
phology of the ischium articulation with the ilium (character
294) (Fig. 2C–F). In addition, SHN.036 shares with other carch-
arodontosaurian theropods the absence of ventral keel on
anterior dorsal vertebrae (shared with Neovenator; Brusatte
et al., 2008); the presence of spinoprezygapophyseal laminae
in mid-caudal vertebrae extending from the medial surface of
the base of the prezygapophysis (shared with Veterupristisaurus:
Rauhut, 2011) (Fig. 2I, J); and the presence of well-defined
anterior centrodiapophyseal lamina with associated centropre-
zygapophyseal fossa in anterior mid-caudal vertebrae (shared
with some carcharodontosaurian allosauroids, including Neove-
nator [Brusatte et al., 2008], Veterupristisaurus [Rauhut, 2011],
and Siats [Zanno and Makovicky, 2013]). Also, the presence of
well-developed lateral lamina at the base of the neural arch of
mid- and posterior caudal centra is a feature shared with other
carcharodontosaurs, such as Veterupristisaurus (Rauhut, 2011)
and Concavenator (Cuesta et al., 2019). However, in Lusovena-
tor, there is an additional smaller lamina adjacent to the pos-
terior articular facet that is not present in Veterupristisaurus or
Concavenator (Fig. 2G, H).
 Malafaia et al.—New carcharodontosaurian dinosaur from Portugal (e1768106-4)

FIGURE 2. Selected elements of Lusovenator santosi, gen. et sp. nov., and other carcharodontosaurs showing some of the features that clearly nest the
new taxon within Carcharodontosauria. A, L. santosi, SHN.036, right ilium in medial view. B, MB.R 3628, possible carcharodontosaur from Tendaguru,
right ilium in medial view. C, D, L. santosi, SHN.036, right ischium in C, proximal and D, lateral views. E, F, Acrocanthosaurus atokensis, NCSM 14345,
left ischium in E, proximal and F, lateral views. G, I, L. santosi, SHN.036, middle caudal vertebrae in G, lateral and I, dorsal views. H, J, MB.R 1938,
Veterupristisaurus milneri, middle caudal vertebrae in H, lateral and J, dorsal views. Abbreviations: acdl, anterior centrodiapophyseal lamina; alr,
anterior lateral ridge; ilc, iliac peduncle; pamr, preacetabular medial ridge; polr, posterior lateral ridge; sprl, spinoprezygapophyseal lamina. Scale
bars equal 100 mm (A–F) and 50 mm (G–J).

DISCUSSION of the prezygapophyses to the distal end of the postzygapophyses

in mid- and posterior caudal vertebrae. The morphology of the
SHN.036 was previously interpreted as belonging to an indeter-
ilium, oval with an extremely convex dorsal margin and relatively
minate allosauroid with a combination of features distinct from
short anteroposteriorly, is distinct from the almost straight and
other taxa known for the Upper Jurassic of the Iberian Peninsula,
elongated ilium of Allosaurus (Gilmore, 1920; Madsen, 1976;
Allosaurus and Lourinhanosaurus antunesi (Malafaia et al.,
Chure, 2000) and Lourinhanosaurus antunesi (Mateus, 1998).
2017a). The type specimen of Lusovenator santosi (SHN.036)
Another unusual feature of the ilium of Lusovenator santosi is
differs from Allosaurus in several features, including the
the supraacetabular crest that forms a prominent ventrolaterally
absence of ventral keel on anterior dorsal vertebrae, the triangu-
projecting shelf, which is somewhat similar to the hypertrophied
lar morphology of the hypantrum, mid-caudal vertebrae with
supraacetabular crest present in some basal theropods, such as
well-developed lateral crests projecting from the dorsal margin
Monolophosaurus jiangi, Ceratosaurus nasicornis, and Dilopho-
of the centra adjacent to the anterior articular facet, and presence
saurus wetherilli (Zhao et al., 2010; Carrano et al., 2012). Some
of well-developed and continuous laminae projecting from the tip
 Malafaia et al.—New carcharodontosaurian dinosaur from Portugal (e1768106-5)

FIGURE 3. Time-calibrated evolutionary tree for Allosauroidea. Agreement subtree produced in TNT, with additional taxa incorporated (Veterupris-
tisaurus milneri) marked with dashed line (see Fig. S1 for the full version of this tree). The pie charts represent relative frequencies of the clades in
different paleogeographic areas calculated based on the number of taxa known in each area. Silhouette of dinosaur modified from the drawing of
Scott Hartman. Global paleogeographic reconstructions modified from maps in Blakey (2019).
 Malafaia et al.—New carcharodontosaurian dinosaur from Portugal (e1768106-6)
small Allosaurus specimens have a more oval, shorter, and higher
morphology of the ilium, suggesting that this feature is somewhat
variable. However, even in those small-sized specimens, the
supraacetabular crest does not project ventrolaterally and the
overall shape of the ilium is more elongate than in SHN.036.
Lusovenator santosi is the oldest carcharodontosaurian allosaur-
oid yet discovered from Laurasia and supports unequivocally
the hypothesis of a pre-Cretaceous radiation of the clade
(Fig. 3). Until recently (Rauhut, 2011; Malafaia et al., 2019), the
fossil record of carcharodontosaurs was restricted to the Cretac-
eous and they were a significantly diverse group of large-bodied
carnivorous dinosaurs during the Late Cretaceous and, especially,
present in southern landmasses. The first exception to this tem-
poral distribution was Veterupristisaurus milneri from the
Middle Dinosaur Member of Tendaguru Formation in southeast-
ern Tanzania (Rauhut, 2011). In the case of the contemporaneous
L. santosi, its presence in uppermost Kimmeridgian to Tithonian
strata represents a second exception. Thus, the temporal distri-
bution of Lusovenator and the carcharodontosaurian affinity of
V. milneri clearly indicate that the origin of this allosauroid
clade of predators should move to at least the earliest Kimmerid-
gian (∼157 Ma). Despite sharing some features, L. santosi can be
distinguished from V. milneri by the morphology of the caudal
vertebrae, namely, by the absence of an additional lateral
lamina adjacent to the posterior articular facet in V. milneri or
the much more developed anterior lateral lamina in the taxon
from Tendaguru. These differences indicate that despite being
closely related, L. santosi and V. milneri are sufficiently distinct
not to be considered synonyms. The discovery of L. santosi
demonstrates that the oldest representatives of Carcharodonto-
sauria, similarly to Tyrannosauroidea (Rauhut, 2003a), were
already present in the Iberian Peninsula before reaching the
apices of the trophic pyramids during the Cretaceous. Similarly,
the dinosaurian fossil record of the Iberian Peninsula and Tanza-
nia also includes the possible first occurrences of somphospondy-
lan sauropods, a group that also became dominant during the
Cretaceous (Mannion et al., 2019; Mocho et al., 2019).
The identification of carcharodontosaurian theropods in the
Upper Jurassic of the Lusitanian Basin highlights an interesting eco-
logical feature, the sympatric coexistence of, at least, two medium- to
large-sized allosauroids: Allosaurus europaeus and L. santosi, poss-
ibly in addition to Lo. antunesi. The last taxon has been recovered in
some recent phylogenetic analyses as belonging to Coelurosauria
(e.g., Carrano et al., 2012). The analysis performed here also recov-
ered this taxon as a coelurosaurian theropod based on a single
feature related to the morphology of anterior and middle chevrons
with unexpanded distal ends. In fact, this morphology is shared with
some coelurosaurs (e.g., Compsognathus longipes; Peyer, 2006), but
also with ceratosaurs (e.g., Ceratosaurus nasicornis; Madsen and
Welles, 2000) and Concavenator corcovatus (Ortega et al., 2010;
Cuesta et al., 2019). Moreover, Lo. antunesi shares with allosauroid
tetanurans the presence of a distinct kink in the anterior margin of
mid-caudal neural spines, which has been interpreted as a synapo-
morphy of Allosauroidea (Rauhut, 2003b), and the presence of an
anterior spur in the neural spine of mid-caudal vertebrae (also
shared with some non-allosauroid tetanurans; e.g., Rauhut et al.,
2016). The phylogenetic position of this taxon is unstable, but its
relationship with early branching allosauroids, closely related to
metriacanthosaurids, currently seems to be the consensus hypothesis
(Mateus, 1998; Benson, 2010; Benson et al., 2010).
The identification of L. santosi as a member of Carcharodonto-
sauria has some relevant paleobiogeographic implications.
Despite the similarity of Late Jurassic faunas from the Morrison
Formation and the Lusitanian Basin, carcharodontosaurian allo-
sauroids are absent in the former area. The development of a ter-
restrial dispersal route between Europe and North America after
the late Tithonian could explain the absence of this clade in the
Morrison Formation and, additionally, their presence during the
Early and Late Cretaceous of North America (e.g., Harris,
1998; Zanno and Makovicky, 2013). A land reconnection
between Europe and North America due to a drop in sea level
during the mid-Valanginian (ca. 137 Ma) was previously proposed
explaining the paleobiogeographic context of other dinosaur
groups such as the turiasaurian sauropods (Royo-Torres et al.,
2017). Within this paleobiogeographic context, the fossil record
indicates that the Iberian Peninsula had a key role in the dis-
persion of dinosaurs to North American landmasses during the
Jurassic–Cretaceous transition.
CONCLUSION
A new carcharodontosaurian theropod, Lusovenator santosi, gen.
et sp. nov., is defined based on a partial skeleton of a juvenile indi-
vidual, SHN.036, collected in upper Kimmeridgian levels of the Lusi-
tanian Basin. A second specimen, SHN.019, corresponding to some
axial and appendicular elements of a large-sized individual, from
upper Tithonian strata in the same geographic area, is tentatively
proposed as referred material. Also, an isolated femur previously
tentatively assigned to Lourinhanosaurus antunesi is interpreted
as belonging to a carcharodontosaur. Thus, the fossil record of
these allosauroids in the Lusitanian Basin spans from the Kimmer-
idgian to the upper Tithonian. Lusovenator santosi represents the
oldest member of Carcharodontosauria defined in the Upper Juras-
sic of Laurasia and extends the record of this clade, which was
already represented in the Lower Cretaceous of Europe. Despite
the great similarity in the theropod faunal composition, the recog-
nition of a carcharodontosaurian allosauroid taxon provides a com-
ponent of the Upper Jurassic Lusitanian Basin distinct from its
contemporaneous Morrison Formation in North America. The
development of a terrestrial dispersal route connecting these land-
masses after the late Tithonian could explain the absence of Carch-
arodontosauria in the diverse theropod faunas of the Morrison
Formation and, additionally, their presence during the Early and
Late Cretaceous of North America. The new carcharodontosaurian
theropod provides the first evidence for sympatry among allosaur-
oid theropods in the Late Jurassic of Europe. Additionally, the dis-
covery of L. santosi sheds light on a poorly known period of
carcharodontosaurian history and adds important data to the knowl-
edge of the early evolution of these allosauroids.
ACKNOWLEDGMENTS
We thank J. J. dos Santos for collecting the specimen. J. J. dos
Santos donated to the Sociedade de História Natural (SHN) a
large collection of vertebrate fossils that he collected during
more than 20 years, mostly from the Upper Jurassic of the Lusita-
nian Basin, and that has allowed the description of several new
taxa, including the pleurosternid turtle Selenemys lusitanica, the
ornithopod Eousdryosaurus nanohallucis, and the sauropod
Oceanotitan dantasi. We also thank the editors and reviewers
for comments and suggestions, G. Ramalheiro for preparation
of some elements of SHN.019, E. Cuesta for photos of carcharo-
dontosaur specimens, C. Moniz and M. Cachão for comments on
the paper, and the following for access to specimens: B. C. Silva
(Sociedade de História Natural), R. Castanhinha and C. Tomás
(Museu da Lourinhã), V. dos Santos (Museu Nacional de História
Natural e da Ciência), S. Langreo (Museo Paleontológico de Cas-
tilla-La Mancha), R. Allain (Muséum national d’Histoire natur-
elle), L. Chiappe (Dinosaur Institute of the Natural History
Museum of Los Angeles County), K. Carpenter (Denver
Museum of Nature and Science), R. Scheetz and B. Britt
(Brigham Young University), M. Getty, M. Loewen, and
R. Irmis (Natural History Museum of Utah), D. Chure (Dinosaur
National Monument), S. Chapman (Natural History Museum),
P. Jeffery (Oxford University Museum of Natural History), and
 Malafaia et al.—New carcharodontosaurian dinosaur from Portugal (e1768106-7)
T. Schossleitner and D. Schwarz (Museum für Naturkunde). Fun-
dação para a Ciência e a Tecnologia grant SFRH/BD/84746/2012
to E.M. and a protocol between Câmara Municipal de Torrres
Vedras (CMTV) SHN supported this research. P.M.’s research
is funded by Fundação para a Ciência e a Tecnologia/Ministério
da Ciência, Tecnologia e Ensino Superior (FCT/MCTES) for
one Concurso Estímulo ao Emprego Científico (CEEC) individ-
ual contract (CEECIND/00726/2017).
ORCID
Elisabete Malafaia http://orcid.org/0000-0003-4894-4257
Pedro Mocho http://orcid.org/0000-0002-3348-5572
Fernando Escaso http://orcid.org/0000-0001-7642-1555
Francisco Ortega http://orcid.org/0000-0002-7431-354X
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Submitted May 13, 2019; revisions received March 30, 2020;
accepted April 9, 2020.
Handling editor: Lindsay Zanno.