The oldest Crocodylia?

A new
eusuchian from the Late Cretaceous
(Cenomanian) of Portugal

Octvio MATEUS
Universidade Nova de Lisboa FCT, Portugal;
Museu da Lourinh
American Museum of Natural History

Pedro M. CALLAPEZ
Universidade de Coimbra, Coimbra, Portugal;

Eduardo PURTOLAS-PASCUAL
Universidade Nova de Lisboa FCT, Portugal
SVP Meeting @ Calgary 2017.08.25
 Phylogeny of Crocodylomorpha Based on Holliday & Gardner
(2012), Brochu (2012), Narvez
(2015), Turner (2015) and Turner &
- Crocodylomorpha lived from the Late Triassic (late Carnian, Pritchard (2015)
~ 230 Ma) to the present.

- Great phylogenetic, ecological, morphological and trophic

variability (not living fossils).

- More than 30 families, 350 genera and 650 species.

Nowadays about 23 species, 8 genera and 3 families.
- Eusuchia true crocodiles is a clade of neosuchians which
includes all extant crocodylians and several extinct species.

- Temporal range: Early Cretaceous (Barremian) - Present.

- Global distribution.
 Phylogeny of Eusuchia
- Eusuchia is composed by several clades:

Two extinct and endemic basal clades from

Europe: Hylaeochampsidae and Allodaposuchidae.

The crown-group Crocodylia:

1) Gavialoidea.

2) Alligatoroidea.

3) Crocodyloidea.

4) Other extinct clades such as Borealosuchus,

Based on Salisbury et al. (2006), Purtolas et al. (2011), Brochu (2012), Blanco et al.
(2014), Purtolas-Pascual et al. (2014), Narvez et al. (2015) and Narvez (2015)
Planocraniidae or Aegyptosuchidae.
 Phylogeny of Eusuchia
Eusuchia is characterized by the combination of
Fully developed
three main apomorphies
bony palate, with
choana posteriorly
placed and
bordered by the
Eusuchia Non Eusuchia pterygoids

Eusuchia Non Eusuchia

Crocodylus affinis Mahajangasuchus insignis

Procoelous vertebrae

Based on Salisbury & Frey (2001), Pol et al. (2009), Salisbury et al. (2006),
Goniopholididae
Purtolas et al. (2011), Brochu (2012), Blanco et al. (2014), Purtolas-Pascual et
al. (2014), Narvez et al. (2015) and Narvez (2015) Sagitally segmented paravertebral shield
Casais dos Carecos. Tentgal Fm.
at Lusitanian Basin
Young et al., (2016), Krebs (1976)
The two oldest known crocodylomorph eggs (Russo et al. 2017)
Schwarz (2002)
Jonet (1981)
Zbyszewski (1949)
Sauvage (1898)
Antunes (1975)
 X

Casais dos Carecos

site

~95 M.a.
New specimen from Portugal (Tentgal)
 Crocodylia
New taxon diagnosis
- Presence of a small-sized and narrow slit-
shaped external mandibular fenestra (emf)
between the dentary-angular suture, with
the dentary forming its anterior and dorsal
margin and the angular forming its
posterior and ventral margin, the
surangular does not participate in the
fenestra.

Allodaposuchidae

Lohuecosuchus (Narvez et al., 2015)

Specimen from Portugal

Agaresuchus (Narvez et al., 2016) Iharkutosuchus (Hylaeochampsidae) Borealosuchus (Brochu et al., 2012)
 New taxon diagnosis
Massive postorbital bar with a very marked mediolateral
compression being twice as wide anteroposteriorly as
mediolaterally.
Specimen from Portugal

Gavialis

Posterior view
Alligator
Crocodylus

Lateral view

Tomistoma
 New taxon diagnosis
Dorsal margin of the infratemporal fenestra very
elongated, with the quadratojugal contacting the base
of the skull table posteriorly far from the postorbital
bar giving to the infratemporal fenestra a trapezoidal
contour rather than triangular.

Specimen from Portugal

Crocodylus Gavialis

Alligator Tomistoma

Lateral view
 New taxon diagnosis
In addition, this specimen differs from all other eusuchians
by the exclusive combination of the following
synapomorphies:

- Absence of a posterior process of the splenial between

the angular and coronoid.

- Maxilla broadly separates the ectopterygoid from the

maxillary tooth row.

- Braincase wall lateral to the basisphenoid rostrum

smooth, without sulcus.

Specimen from Portugal

 Phylogenetic position

Portuguese specimen

Portuguese specimen

Brevirostres

- Matrix of Narvez et al. (2016) with modifications in the codes of some first hand
studied taxa (Allodaposuchus precedens, Agaresuchus subjuniperus, Arenysuchus
gascabadiolorum, Iharkutosuchus makadii).

- Crown group Crocodylia is resolved but with little support.

- The specimen of Portugal appear within Crocodylia.

-
 Is the specimen from Portugal the oldest crocodylian?
Definition of Crocodylia

- The phylogenetic definition of Crocodylia was established by Clark (1986) and expanded by later workers (Benton & Clark
1988; Norell et al. 1994; Salisbury & Willis 1996; Brochu 1997a, 1999a, 2000, 2003).

- Crocodylia: last common ancestor of Gavialis gangeticus (Indian gharial), Alligator mississippiensis (American alligator),
and Crocodylus niloticus (Nile crocodile) and all of its descendants.

- The oldest reliable record of Crocodylia is from the Campanian - Maastrichtian with representatives of Gavialoidea in
Europe and representatives of Alligatoroidea, Crocodyloidea, Borealosuchus and Gavialoidea in North America.

- Most basal eusuchians are known from skull only.

Neosuchia according to Benton & Clark (1988) Crocodylia according to Salisbury & Willis (1996)
 Is the specimen from Portugal the oldest crocodylian?

* In bold are the synapomorphies present in Portuguese specimen.

Diagnosis of Crocodylia according to:
Clark (1986): The first caudal vertebra is biconvex; the posterior cervical vertebrae have well-developed hypapophyses; presence of a external
mandibular fenestra.
Benton & Clark (1988): scapula with nearly horizontal anterior and posterior edges.

Norell & Clark (1990): dorsal osteoderms lacking a flattened articulating surface anteriorly.

Norell (1994): descending process of prefrontal contacts palate in robust suture; pterygoids enclose choanae; choanae situated in
middle of pterygoids; retroarticular process posterodorsally curving and elongate; insertion area for M. pterygoideus posterior extends
onto lateral surface of angular (ambiguous); neural spines on posterior cervical vertebrae ateroposteriorly narrow, rodlike; cervical vertebrae
with well-developed hypapophyses; dorsal osteoderms rounded or ovate; dorsal osteoderms with straight anterior edge; dorsal osteoderms in
more than two longitudinal rows.

Salisbury & Willis (1996): All vertebrae procoelous; internal nares excluded from palatines; external mandibular fenestrae
present; enlarged 4th and 5th maxillary alveoli.

Brochu (1999): retroarticular process projects posterodorsally; anterior dentary teeth project anterodorsally; external mandibular fenestra
present; dentary teeth occlude in line with maxillary toothrow; lacrymal makes broad contact with nasal; no posterior process of maxilla;
nasals excluded, at least externally, from naris, nasals and premaxillae still in contact; mature skull table with nearly horizontal sides; significant
squamosal prongs; exoccipital with small or no boss on paroccipital process, process lateral to cranioquadrate opening long; ectopterygoid
does not extend to posterior tip of lateral pterygoid flange at maturity.
 Is the specimen from Portugal the oldest crocodylian?
Definition of Crocodylia
Salisbury et al. (2006): thoracic and lumbar vertebrae strongly procoelous; quadratojugal spine prominent at maturity; postorbital bar slender;
frontoparietal suture concavoconvex; lacrymal makes broad contact with nasal, no caudal process of the maxilla; mature skull table with nearly
horizontal sides, significant caudolateral process of the squamosal; prefrontal pillars narrow or longitudinally expanded at their dorsal part and
columnar ventrally; first caudal vertebra biconvex, remainder of caudal vertebrae procoelous, with the degree of procoely decreasing terminally;
proximal extremity of the ulna considerably wider than the distal extremity.

Purtolas-Pascual et al. (2014): palatopterygoidal suture far from caudal angle of the suborbital fenestra; ventral margin of
orbit with prominent notch; largest dentary alveolus immediately caudal to fourth is 13 or 14 and a series behind it; secondary choanae
situated towards the caudal margin of the pterygoids; antorbital fenestra absent; foramen arum set in from margin of the
retroarticular process; mature skull table with broad lateral curvature, short caudolateral process of the squamosal.

Blanco et al. (2015): anterior dentary teeth project anterodorsally; retroarticular process projects posterodorsally; frontoparietal suture concavo-
convex; mature skull table with nearly horizontal sides and long posterolateral squamosal rami along paraoccipital process; exoccipital lacks
boss on paraoccipital process; and hypapophyseal keels are present on the eleventh vertebrae behind the atlas.

Narvez et al. (2016): exoccipital with small or no boss on paroccipital process, process lateral to cranioquadrate opening long.

This study: external mandibular fenestra present with discrete concavity on angular dorsal margin; and
exoccipital with small or no boss on the paroccipital process. Ambiguous synapomorphies: Quadrate foramen
aerum on mediodorsal angle of quadrate.

* In bold are the synapomorphies present in Portuguese specimen.

 Is the specimen from Portugal the oldest crocodylian?
Definition of Crocodylia
- After the discoveries of several new basal eusuchians in recent years (e.g., Hylaeochampsidae, Allodaposuchidae...), most of the characters
proposed in previous phylogenies would not be synapomorphies of Crocodylia. These characters would be more diagnostic of Eusuchia.

-
 Is the specimen from Portugal the oldest crocodylian?
Definition of Crocodylia
- If we discard the characters that have subsequently been observed in advanced neosuchians and basal eusuchians, the most diagnostic
combination of characters for Crocodylia observed in most phylogenies correspond to:

1. Skull table with nearly horizontal sides (linear margin) and long posterior
Glen Rose Form
squamosal processes. Character also present in Allodaposuchidae?.
Hylaeochampsa

Caimans (hatchling vs adult)

Crocodylus

Hylaeochampsa
Allodaposuchidae presents the crocodylian condition

Not visible in the Portuguese specimen due to incompleteness Brochu (1997)

 Is the specimen from Portugal the oldest crocodylian?
Definition of Crocodylia
- If we discard the characters that have subsequently been observed in advanced neosuchians and basal eusuchians, the most diagnostic
combination of characters for Crocodylia observed in most phylogenies correspond to:

1. Mature skull table with nearly horizontal sides and long posterior squamosal processes.
Glen Rose Form
Hylaeochampsa
2. Exoccipital lacks boss on paroccipital process. Boss appears in most allodaposuchids and
hylaeochampsids.

Caimans (hatchling vs adult)

Crocodylus

Hylaeochampsa

Brochu (1997)
Not visible in the Portuguese specimen due to incompleteness
 Is the specimen from Portugal the oldest Crocodylian (sensu Crocodylia clade)?
Definition of Crocodylia
If we discard the characters that have subsequently been observed in advanced neosuchians and basal eusuchians, the most
diagnostic combination of characters for Crocodylia observed in most phylogenies correspond to:

1. Mature skull table with nearly horizontal sides and long posterior squamosal processes.

2. Exoccipital lacks boss on paroccipital process.

3. Presence of a mandibular fenestra. Fenestra not present in Allodaposuchidae or Hylaeochampsidae. Present in the specimen
from Portugal.

Lohuecosuchus (Allodaposuchidae
Narvez et al., 2015)
-

Specimen from Portugal

Iharkutosuchus (Hylaeochampsidae) Borealosuchus (Crocodylia, Brochu et al., 2012)

 Implications of the presence of a mandibular fenestra

- The presence of external mandibular fenestra is the only synapomorphy of Crocodylia preserved in the specimen of Portugal.

- The external mandibular fenestra is absent in most non-crocodylian neosuchians and close relatives of Eusuchia, including
Shamosuchus, Rugosuchus, Bernissartia, some goniopholidids, Theriosuchus, and basal eusuchians such as Pachycheilosuchus,
Pietraroiasuchus, Hylaeochampsidae and Allodaposuchidae.
-
- HoweverThis scenario is much more
complex (see Brochu, 2004; Brochu et al.,
2012).

- This character is present throughout

Crocodylia. The presence of this fenestra in
the basal-most gavialoid, Eothoracosaurus, is
not known because that zone is broken,
although the presence of a tiny fenestra is
plausible (Brochu, 2004).
Portuguese specimen
- Eothoracosaurus coded ?, 0, or 1 improves
resolution but not the position of our
specimen.

Matrix of Narvez et al. (2016) + the specimen from Portugal

 Conclusions
- We present a partial skull and jaw of a new eusuchian (choanae clearly enclosed
by the pterygoids) from the early upper Cenomanian (~95 M.a.) of Portugal

- Probably is a new species.

- The cladistic analysis places this specimen at the base of Crocodylia.

- - This specimen would represents the only well documented and valid eusuchian
species in the Cenomanian of Europe and probably the oldest representative of
Crocodylia worldwide (or also probably, the sister taxon of Crocodylia), pulling the
origin of the clade 15 My back.

- The discovery of this new taxon would shed light on the radiation of Eusuchia and
the origin of Crocodylia, which probably would have taken place in Europe.

- The presence of external mandibular fenestra in this new taxon may contribute
with new data on the evolution of this structure through Crocodylia and Eusuchia.

- Due to the fragmentary nature, the phylogenetic position is not very well supported.
We are facing the usual paleontological problem and potential solution: find more
specimens!
 Acknowledgments
Matilde Azenha who found the specimen. THANK YOU!
Alexandra Toms for the lab preparation.
Christopher Brochu for his remarks and advices.
OBRIGADO!
Joana Bruno for the illustrations.

Announcements
University NOVA of Lisbon (Portugal) will host:
Dinosaur Eggs and Babies symposium
4-7 October 2017

EAVP Annual Meeting

(European Association of Vertebrate Paleontologists)
27-29 June 2018

Join the Master in Paleontology at University NOVA of Lisbon, Portugal