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An overview of non-avian theropod discoveries and classification

January 2015
PalArch's Journal of Vertebrate Palaeontology 12:1-73

January 2015
12:1-73

Authors:

Christophe Hendrickx

National Scientific and Technical Research Council

Scott Hartman

University of Wisconsin–Madison

Octávio Mateus

University NOVA of Lisbon

Theropods form a taxonomically and morphologically diverse group of dinosaurs that include extant birds. Inferred relationships between theropod clades are complex and have changed dramatically over the past thirty years with the emergence of cladistic techniques. Here, we present a brief historical perspective of theropod discoveries and classification, as well as an overview on the current systematics of non-avian theropods. The first scientifically recorded theropod remains dating back to the 17th and 18th centuries come from the Middle Jurassic of Oxfordshire and most likely belong to the megalosaurid Megalosaurus. The latter was the first theropod genus to be named in 1824, and subsequent theropod material found before 1850 can all be referred to megalosauroids. In the fifty years from 1856 to 1906, theropod remains were reported from all continents but Antarctica. The clade Theropoda was erected by Othniel Charles Marsh in 1881, and in its current usage corresponds to an intricate ladder-like organization of textquoteleftfamilytextquoteright to textquoteleftsuperfamilytextquoteright level clades. The earliest definitive theropods come from the Carnian of Argentina, and coelophysoids form the first significant theropod radiation from the Late Triassic to their extinction in the Early Jurassic. Most subsequent theropod clades such as ceratosaurs, allosauroids, tyrannosauroids, ornithomimosaurs, therizinosaurs, oviraptorosaurs, dromaeosaurids, and troodontids persisted until the end of the Cretaceous, though the megalosauroid clade did not extend into the Maastrichtian. Current debates are focused on the monophyly of deinonychosaurs, the position of dilophosaurids within coelophysoids, and megaraptorans among neovenatorids. Some recent analyses have suggested a placement of dilophosaurids outside Coelophysoidea, Megaraptora within Tyrannosauroidea, and a paraphyletic Deinonychosauria with troodontids placed more closely to avialans than dromaeosaurids.

Phylogeny and stratigraphic distribution of theropod clades. The phylogenetic classification of theropods follows the results of the cladistic analyses obtained by Sues et al. (2011) for non-neotheropod Theropoda, Smith et al. (2007) and Ezcurra & Brusatte (2011) for non-averostran Neotheropoda, Pol & Rauhut (2012) and Tortosa et al. (2014) for Ceratosauria, Carrano et al. (2012) for non-coelurosaur Tetanurae, Loewen et al. (2013), Lü et al. (2014) and Porfiri et al. (2014) for Tyrannosauroidea, Lee et al. (2014) for Ornithomimosauria, Lamanna et al. (2014) for Oviraptorosauria, and Turner et al. (2012), Godefroit et al. (2013a) and Choiniere et al. (2014b) for non-tyrannosauroid Coelurosauria. Silhouettes by Michael Bech Hussein (Coelophysoidea, Dilophosauridae, Therizinosauria, and Alvarezsauroidea), Jaime Headden (Caenagnathidae), Michael Keesey (Deinocheiridae), William Parker (Daemonosaurus), Travis Tischler (Megaraptora), and Scott Hartman (all others).

…

Earliest historical records of theropod remains in the world. A-B) Distal part of a left femur of Megalosaurus from Cornwell, U.K., in posterior view, and first reported by Plot (1677); A, illustrations by Plot (1677, table 8, fig.4); and B, Brookes (1763, p. 312, figure 317) showing the label 'Scrotum Humanum'; C) Isolated theropod tooth (likely Megalosaurus) from the Stonesfield, U.K., illustrated by Lhuyd (1699, plate 16, figure 1328); D) Right femur of Megalosaurus from Stonesfield, U.K., in anterior view, illustrated by Platt (1758, table 19); E) Right dentary of Megalosaurus bucklandii from Stonesfield, U.K., in medial and posterior views, illustrated by Buckland (1824, plate 40).

…

Earliest historical records of theropod remains in A-B, North America; C, Asia; D, Africa; E-F, South America; G, Oceania; and H, Antarctica. Isolated teeth of: A) Troodon formosus; and B, Deinodon horridus (= Albertosaurus sarcophagus) from the Upper Cretaceous Judith River of Colorado and first reported by Leidy (1856; modified from Leidy 1860, plate 9); C) Isolated theropod tooth of 'Massospondylus rawesi', an abelisaurid from the Upper Cretaceous of India (localities of Takli and Maleri) first reported by Hislop (1861, 1864, illustration by Lydekker 1890, fig. 1); D) Casts of theropod tracks from the Upper Cretaceous of Jebel Bou-Khaïl, Algeria, and first reported by Le Mesle and Peron (1880; illustration by Mesle & Peron 1880, figures 65 and 66); E-F) Isolated theropod tooth from the Upper Cretaceous of Par-Aïk, Argentina, referred to Loncosaurus argentines and first reported by Ameghino (1899); E, illustration by Ameghino (1900, p. 160) and Ameghino (1906, Figure 8); and F, Huene (1929a, plate 41); G) Pedal ungual of an indeterminate theropod from the Upper Cretaceous of Cape Patterson, Australia, and first reported by Woodward (1906); H) Distal part of a tibia of a megalosauroid? theropod from the Upper Cretaceous of Col Crame, Antarctica, discovered in 1988 (modified from Molnar et al. 1996).

…

Cladogram of basal Theropoda showing the relationships of tyrannosauroids and non-coelurosaur theropods. The phylogenetic classification follows the results of the cladistic analyses obtained by Sues et al. (2011) for non-neotheropod Theropoda, Smith et al. (2007) and Ezcurra & Brusatte (2011) for non-averostran Neotheropoda, Pol & Rauhut (2012) and Tortosa et al. (2014) for Ceratosauria, Carrano et al. (2012) for non-coelurosaur Tetanurae, Loewen et al. (2013), Lü et al. (2014) and Porfiri et al. (2014) for Tyrannosauroidea, and Choiniere et al. (2014b) for basalmost Coelurosauria.

…

Cladogram of non-tyrannosauroid Coelurosauria showing the relationships of compsognathids and maniraptoriforms. The phylogenetic classification follows the results of the cladistic analyses obtained by Choiniere et al. (2014b) for basalmost Coelurosauria and Compsognathidae, Longrich & Currie (2009a) and Choiniere et al. (2010b) for Alvarezsauroidea, Lee et al. (2014) for Ornithomimosauria, Senter et al. (2012a) and Pu et al. (2013) for Therizinosauria, Lamanna et al. (2014) for Oviraptorosauria, Turner et al. (2012) for Paraves, and Foth et al. (2014) for Avialae.

…

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Hendrickx et al., Non-Avian Theropods PalArch’s Journal of Vertebrate Palaeontology, 12, 1 (2015)

PalArch Foundation 1

AN OVERVIEW OF NON-AVIAN THEROPOD

DISCOVERIES AND CLASSIFICATION

Christophe Hendrickx*, Scott A. Hartman# & Octávio Mateus*

* Universidade Nova de Lisboa, CICEGe, Departamento de Ciências da Terra, Faculdade

de Ciências e Tecnologia, Quinta da Torre, 2829-516, Caparica, Portugal & Museu da

Lourinhã, 9 Rua João Luis de Moura, 2530-158, Lourinhã, Portugal

# University of Wisconsin-Madison, Department of Geosciences, Madison, WI, 53716

USA

Corresponding author: christophe.hendrickx@hotmail.com

Christophe Hendrickx, Scott A. Hartman & Octávio Mateus. 2015. An Overview of Non-

Avian Theropod Discoveries and Classiﬁcation. - PalArch’s Journal of Vertebrate Palaeon-

tology 12, 1 (2015), 1-73. ISSN 1567-2158. 73 pages + 15 ﬁgures, 1 table.

Keywords: Dinosauria, Theropoda, Discovery, Systematics

ABSTRACT

Theropods form a taxonomically and morphologically diverse group of dinosaurs that

include extant birds. Inferred relationships between theropod clades are complex and

have changed dramatically over the past thirty years with the emergence of cladistic

techniques. Here, we present a brief historical perspective of theropod discoveries and

classiﬁcation, as well as an overview on the current systematics of non-avian thero-

pods. The ﬁrst scientiﬁcally recorded theropod remains dating back to the 17th and

18th centuries come from the Middle Jurassic of Oxfordshire and most likely belong to

the megalosaurid Megalosaurus. The latter was the ﬁrst theropod genus to be named

in 1824, and subsequent theropod material found before 1850 can all be referred to

megalosauroids. In the ﬁfty years from 1856 to 1906, theropod remains were reported

from all continents but Antarctica. The clade Theropoda was erected by Othniel Charles

Marsh in 1881, and in its current usage corresponds to an intricate ladder-like organi-

zation of ‘family’ to ‘superfamily’ level clades. The earliest deﬁnitive theropods come

from the Carnian of Argentina, and coelophysoids form the ﬁrst signiﬁcant theropod

radiation from the Late Triassic to their extinction in the Early Jurassic. Most subsequent

theropod clades such as ceratosaurs, allosauroids, tyrannosauroids, ornithomimosaurs,

therizinosaurs, oviraptorosaurs, dromaeosaurids, and troodontids persisted until the

end of the Cretaceous, though the megalosauroid clade did not extend into the Maas-

trichtian. Current debates are focused on the monophyly of deinonychosaurs, the posi-

tion of dilophosaurids within coelophysoids, and megaraptorans among neovenatorids.

Some recent analyses have suggested a placement of dilophosaurids outside Coelo-

physoidea, Megaraptora within Tyrannosauroidea, and a paraphyletic Deinonychosauria

with troodontids placed more closely to avialans than dromaeosaurids.

Hendrickx et al., Non-Avian Theropods PalArch’s Journal of Vertebrate Palaeontology, 12, 1 (2015)

PalArch Foundation 2

Introduction

Theropods form a clade of bipedal tetrapods

among which birds and all strictly carnivorous

dinosaurs are found (e.g., Gauthier 1986; Sere-

no 1997; Holtz & Osmólska 2004; Holtz 2012;

Naish 2012). Along with sauropodomorph and

ornithischian clades, they appeared in the Late

Triassic (Figure1) and rapidly acquired a world-

wide distribution, being present on every con-

tinent by the Lower Jurassic (Tykoski & Rowe

2004). In the Jurassic (possibly as early as the

Middle Jurassic, based on ghost ranges; e.g., Hu

et al. 2009; Godefroit et al. 2013a, b), small the-

ropods gave rise to birds, the only dinosaurs to

survive the Cretaceous-Paleocene (K-Pg) mass

extinction event 66 million years ago (Figure1).

After surviving the K-Pg extinction event, birds

radiated into ecological niches left by non-avian

dinosaurs (Padian & Chiappe 1998; Chiappe &

Witmer 2002; Naish 2012). As a result, thero-

pods are one of the most successful groups

of tetrapods, and the most morphologically

and taxonomically diverse clade of dinosaurs

(Rauhut 2003a; Holtz 2012; Foth & Rauhut

2013).

Non-avian theropods (i.e., Theropoda exclud-

ing Avialae) were the dominant terrestrial pred-

ators in Jurassic and Cretaceous ecosystems

worldwide (Rauhut 2003a; D’Amore 2009).

Though their diversity and disparity remained

high through the end of the Cretaceous, they

became extinct at the end of the Cretaceous

concurrent with all other clades of non-avian

dinosaurs (Rauhut 2003a; Holtz et al. 2004; Up-

church et al. 2011; Brusatte et al. 2012b, 2015).

While non-avian theropods include the major-

ity (if not all) of meat-eating dinosaurs, many

theropod clades became secondarily adapted to

herbivorous diets (Barrett 2005; Xu et al. 2009b;

Zanno et al. 2009; Zanno & Makovicky 2011),

and several taxa have been described as omni-

vores (Holtz et al. 1998; Lee et al. 2014), insec-

tivores (Senter 2005) or ﬁlter feeders (Norell et

al. 2001). The non-avian theropod body plan un-

derwent relatively little modiﬁcation during the

evolution of the clade, being almost exclusively

bipedal and exhibiting, for the large majority

of them, elongated necks and a long, horizon-

tally projecting tail (n.b., some theropods such

as tyrannosaurids and caudipterids had a short

neck and short tail, respectively). Variation in

the postcranium mostly occurs in the forelimb,

manual and pelvic morphology, hind limbs pro-

portion as well as the vertebral counts, ossiﬁca-

tion, and elongation of the neural spine. Some

theropods like abelisaurids had short stubby

arms bearing four short ﬁngers (e.g., Ruiz et

al. 2011; Burch & Carrano 2012) whereas oth-

ers like therizinosaurids possess elongated fore-

limbs with three slender ﬁngers bearing large

claws (Clark et al. 2004; Zanno 2010a). Like-

wise, although a large majority of theropods

exhibit short neural spines, some spinosaurids,

allosauroids and deinocheirids have developed

hypertrophied spines forming a hump or a sail

on the back of these animals (Bailey 1997; Lee

et al. 2014). Unlike the postcranial skeleton,

there is a tremendous diversity of skull mor-

phology in non-avian theropods, from the elon-

gated skull of spinosaurids showing a terminal

spatulate rosette (Charig & Milner 1997; Dal

Sasso et al. 2005) to the short parrot-like skull

and edentulous jaws of oviraptorids (Xu & Han

2010). Recent discoveries of non-avian thero-

pods such as the rodent-like Incisivosaurus (Xu

et al. 2002a), the beaked Limusaurus (Xu et al.

2009b), the crested Guanlong (Xu et al. 2006),

the long snouted Buitreraptor (Makovicky et al.

2005) and the duck-billed Deinocheirus (Lee et

al. 2014) indicate a particularly high variety of

skull morphologies among theropod dinosaurs

(Brusatte et al. 2012c; Foth & Rauhut 2013).

Given such morphological and taxonomic

diversity, it is not surprising that theropod clas-

siﬁcation is particularly complex, with Thero-

poda currently comprising more than 20 clades

at the ‘family’ and ‘super-family’ level. With the

emergence of cladistic approaches and the dis-

covery of a large number of new theropod taxa,

higher-level theropod relationships have also

changed dramatically over the past thirty years.

This paper aims to present an overview of the

current state of knowledge on the systematics

of non-avian theropods and a general descrip-

tion of each subclade. A historical perspective

of initial discoveries and the evolution of thero-

pod classiﬁcation are also provided.

Hendrickx et al., Non-Avian Theropods PalArch’s Journal of Vertebrate Palaeontology, 12, 1 (2015)

PalArch Foundation 3

Figure 1. Phylogeny and stratigraphic distribution of theropod clades. The phylogenetic classiﬁcation of theropods follows

the results of the cladistic analyses obtained by Sues et al. (2011) for non-neotheropod Theropoda, Smith et al. (2007) and

Ezcurra & Brusatte (2011) for non-averostran Neotheropoda, Pol & Rauhut (2012) and Tortosa et al. (2014) for Ceratosauria,

Carrano et al. (2012) for non-coelurosaur Tetanurae, Loewen et al. (2013), Lü et al. (2014) and Porﬁri et al. (2014) for

Tyrannosauroidea, Lee et al. (2014) for Ornithomimosauria, Lamanna et al. (2014) for Oviraptorosauria, and Turner et al.

(2012), Godefroit et al. (2013a) and Choiniere et al. (2014b) for non-tyrannosauroid Coelurosauria. Silhouettes by Michael

Bech Hussein (Coelophysoidea, Dilophosauridae, Therizinosauria, and Alvarezsauroidea), Jaime Headden (Caenagnathidae),

Michael Keesey (Deinocheiridae), William Parker (Daemonosaurus), Travis Tischler (Megaraptora), and Scott Hartman (all

others).

Hendrickx et al., Non-Avian Theropods PalArch’s Journal of Vertebrate Palaeontology, 12, 1 (2015)

PalArch Foundation 4

Historical Background

First Discoveries

The description of the ﬁrst theropod remains

and the ﬁrst dinosaur material go hand in hand,

as the ﬁrst dinosaur bones and teeth reported

in the literature belong to theropods (Lebrun

2004). All theropod material reported in the

17th, 18th, and the ﬁrst half of the 19th cen-

tury came from England and France, and has

been referred to megalosauroid theropods, with

most remains being assigned to Megalosauri-

dae. This coincidence can be explained by two

independent factors: 1) the emergence of verte-

brate paleontology in the Early modern period

and early 19th century in Western Europe, with

scientists like Georges Cuvier, Gideon Mantell,

and Richard Owen; and 2) the excavation, at

that time, of vertebrate remains from Middle

Jurassic limestone quarries of Stonesﬁeld (Ox-

fordshire) and Caen (Normandy), a period of

time when megalosauroids were the dominant

theropods in Europe.

Theropod fossils were almost certainly

found by prescientiﬁc societies prior to the 17th

century, but the discovery of these unusual re-

mains were interpreted in ways that gave rise

to myths and legends (Buffetaut 1994; Lebrun

2004; Spalding & Sarjeant 2012). Theropod

tracks from the Lower Cretaceous sandstones

of Paraíba in north-eastern Brazil were, for in-

stance, considered by Amerindians to pertain to

giant running birds (Leonardi 1984; Mayor &

Sarjeant 2001). Likewise, a set of theropod

tracks visible on Cenomanian limestone in the

south of Algeria was believed by Arabs to be-

long to a giant ostrich, property of a venerated

man buried nearby (Taquet 2010).

The ﬁrst published record of a theropod

bone is of an incomplete left femur described

and ﬁgured by Robert Plot in his 1677 ‘Natural

History of Oxfordshire’ (Figure 2A). The fossil

was dug up from a quarry in the Parish of Corn-

well, Oxfordshire, and probably pertains to the

megalosaurid Megalosaurus (Delair & Sarjeant

1975, 2002). Plot (1677) correctly identiﬁed the

bone as a distal femoral condyle (capita femoris

inferiora), and wondered whether this partial

femur belonged to an elephant brought to Brit-

ain by the Romans. Plot (1677), however, noted

many differences with the femur of elephants

and instead referred the bone to a human giant

also brought by the Romans (Evans 2010). This

portion of femur was reillustrated by English

naturalist Richard Brookes (1763) who labeled

the ﬁgure ‘Scrotum Humanum’, given the super-

ﬁcial similarity of the distal condyle to human

testicles (Figure 2B). Although this binomial

term was clearly used as a descriptive appel-

lation by Brookes (Spalding & Sarjeant 2012),

some have proposed its use as a valid scientiﬁc

name. That would make ‘Scrotum Humanum’ the

ﬁrst formal binomial name given to a dinosaur

and a senior synonym of Megalosaurus buck-

landii (Halstead 1970; Delair & Sarjeant 1975),

a proposition which was rejected by the Inter-

national Zoological Commission (Halstead &

Sarjeant 1993; Delair & Sarjeant 2002).

Isolated theropod teeth were ﬁrst described

and ﬁgured in 1699 by Welsh naturalist Edward

Lhuyd in his catalogue of fossils and minerals

Lithophylacii Britannici Ichnographica (Lhuyd

1699). The specimen number 1328 (Lhuyd

1699, plate 16), originally ascribed to a ﬁsh by

Lhuyd (1699), corresponds to an isolated tooth

from the Middle Jurassic Great Oolite of Stones-

ﬁeld (Figure 2C). This shed tooth greatly resem-

bles Megalosaurus and most likely belongs to

that taxon (Delair & Sarjeant 2002). Additional

theropod ﬁndings reported in the 18th century

include a limb bone from Stonesﬁeld labeled

specimen a.1 by John Woodward (1729) in his

catalogue of British fossils from his personal

collection. This section of limb bone is current-

ly preserved in the Sedgwick Museum of Cam-

bridge (specimen D.30.1) and, once again, likely

pertains to Megalosaurus (Delair & Sarjeant

1975, 2002). It may, therefore, be the earliest-

discovered bone that can still be identiﬁed as

belonging to a theropod with conﬁdence (De-

lair & Sarjeant 1975, 2002). Later, an incomplete

femur described and illustrated by Platt (1758)

was identiﬁed as belonging to a hippopotamus,

a rhinoceros, or an unknown animal of large

size (Figure 2D). This large femur, which is also

from Stonesﬁeld, was recently referred to Meg-

alosaurus bucklandii (Evans 2010).

The ﬁrst theropod taxon to be recognized

as reptilian and formally described in the lit-

erature is, in fact, Megalosaurus, coined by Wil-

liam Buckland in 1824 (although the generic

name was already announced by James Par-

kinson in 1822). Material originally ascribed to

Megalosaurus included a right dentary with a

well-preserved erupted tooth (Figure 2E), ribs,

hind-limb elements, pelvic bones, and sacral

Hendrickx et al., Non-Avian Theropods PalArch’s Journal of Vertebrate Palaeontology, 12, 1 (2015)

PalArch Foundation 5

Figure 2. Earliest historical records of theropod remains in the world. A-B) Distal part of a left femur of Megalosaurus from

Cornwell, U.K., in posterior view, and ﬁrst reported by Plot (1677); A, illustrations by Plot (1677, table 8, ﬁg.4); and B, Brookes

(1763, p. 312, ﬁgure 317) showing the label ‘Scrotum Humanum’; C) Isolated theropod tooth (likely Megalosaurus) from the

Stonesﬁeld, U.K., illustrated by Lhuyd (1699, plate 16, ﬁgure 1328); D) Right femur of Megalosaurus from Stonesﬁeld, U.K.,

in anterior view, illustrated by Platt (1758, table 19); E) Right dentary of Megalosaurus bucklandii from Stonesﬁeld, U.K., in

medial and posterior views, illustrated by Buckland (1824, plate 40).

Hendrickx et al., Non-Avian Theropods PalArch’s Journal of Vertebrate Palaeontology, 12, 1 (2015)

PalArch Foundation 6

and caudal vertebrae, all collected in the Tayn-

ton Limestone Formation (middle Bathonian)

of Stonesﬁeld, Oxfordshire. As Buckland (1824)

did not provide a species name for Megalosau-

rus, the type species Megalosaurus conybeari

was proposed by Ferdinand von Ritgen in 1826

(von Ritgen 1826). This author failed to provide

a description and diagnosis for the species, al-

lowing Mantell (1827) to be the ﬁrst scientist

to name and diagnose a theropod species, i.e.,

Megalosaurus bucklandii, which is the name

currently accepted by the scientiﬁc community.

Streptospondylus altdorfensis (Meyer 1832)

and Poekilopleuron bucklandii (Eudes-Deslong-

champs 1837) from France were the ﬁrst non-

avian theropods to be described in the literature

outside England, and the second and third Me-

sozoic theropods to be formally named. These

two megalosauroids, considered valid species

(Carrano et al. 2012), are only known from post-

cranial remains. The material of Streptospondy-

lus, discovered in the Callovian Vaches Noires

cliffs around 1770, was mixed with crocodilian

remains, and interpreted as a crocodile by Cu-

vier (1808, 1812, 1824). The remains of Poeki-

lopleuron from the Calcaire de Caen Formation

(middle Bathonian) in Caen, Normandy, were

correctly identiﬁed as belonging to a large rep-

tile closely related to Megalosaurus. Unfortu-

nately, the material was lost during World War II

and, besides the original illustrations provided

by Eudes-Deslongchamps (1837), only casts of

some bones remain (Allain & Chure 2002).

Although Buckland (1824) and Mantell

(1827) were the ﬁrst to give a relatively good de-

scription of the dentition of Megalosaurus, Rich-

ard Owen was the ﬁrst scientist to exhaustively

investigate the tooth anatomy of theropods and

many other vertebrates. In his treatise on verte-

brate teeth, ‘Odontography’ (Owen 1840-1845),

and his richly illustrated four volume ‘A History

of British Fossil Reptiles’ (Owen 1849-1884),

Owen provided a comprehensive description

and illustration of the crowns, denticles, and

internal structure of the teeth of Megalosaurus

bucklandii and Suchosaurus cultridens. The lat-

ter was erected by Owen (1840-1845) based on

isolated teeth from the Wealden of Tilgate For-

est, near Cuckﬁeld (Sussex). Interestingly, the

teeth of Suchosaurus were discovered by Man-

tell, and were ﬁrst described and illustrated by

Mantell (1822) and Cuvier (1824), respectively

(Buffetaut 2010). Cuvier (1824), Mantell (1827,

1833), and Owen (1840-1845; 1849-1884) all re-

ferred these isolated teeth to crocodilians, yet

they closely resemble those of the spinosaurid

Baryonyx walkeri discovered much later. Sucho-

saurus teeth are now considered as belonging

to either Baryonyx or an unnamed member of

Baryonychinae (Milner 2003; Buffetaut 2007;

Mateus et al. 2011).

The ﬁrst non-megalosauroid theropod to be

formally described is Nuthetes destructor from

the Purbeck Formation (Berriasian, Early Cre-

taceous) of Durlston Bay, Dorset. This tentative

dromaeosaurid was erected by Owen (1854)

based on an incomplete dentary and some

isolated teeth originally assigned to a lizard

or a varanid (Milner 2002). A few years later,

Compsognathus longipes (Wagner 1861), from

the Solnhofen Limestone of Germany, was the

ﬁrst non-avian theropod preserving a nearly

complete and slightly disarticulated skull and

skeleton to be reported in the literature. This

theropod was discovered in Germany around

1859 (Wellnhofer 2008) and was reported by

Wagner (1859) the same year. It remained one

of the most completely known theropods for

more than a century (Ostrom 1978).

After Europe, North America became the

second continent to yield theropod remains de-

scribed by paleontologists. The ﬁrst theropod

fossils reported were isolated teeth discovered

in 1855 by eminent American scientist Ferdi-

nand Vandiveer Hayden from the Upper Cre-

taceous of Montana, at the conﬂuence of the

Missouri and Judith rivers (Breithaupt 1999).

The dental material was brieﬂy described one

year later by Leidy (1856) who erected two

new species, Deinodon horridus based on sev-

eral fragment of teeth (Figure 3A) and Troodon

formosus based on a single shed tooth (Figure

3B). Troodon and Deinodon were originally

thought to belong to a ‘lacertian’ (a large Moni-

tor according to Leidy 1860) and a relative of

Megalosaurus, respectively (Leidy 1856, 1860).

Troodon is now considered to be a valid species

of troodontid (Currie 1987), whereas Deinodon

has been recognized as belonging to an uniden-

tiﬁed tyrannosaurid, probably Albertosaurus

known from the same deposits (Breithaupt

1999; Breithaupt & Elizabeth 2008).

Shortly after Leidy’s description of theropod

teeth from North America, the Reverend Ste-

phen Hislop (1861, 1864) reported the discov-

ery of isolated theropod teeth from the Upper

Hendrickx et al., Non-Avian Theropods PalArch’s Journal of Vertebrate Palaeontology, 12, 1 (2015)

PalArch Foundation 7

Cretaceous of India. One of them was discov-

ered by Mr. Rawes in the locality of Takli, in the

Nagpur area of Maharashtra, and represents the

earliest historical record of theropod dinosaurs

in Asia (Carrano et al. 2010). The shed tooth was

sent to the Geological Society’s Museum of Lon-

don (which is now part of the Natural History

Museum) and studied and illustrated by English

naturalist Richard Lydekker (1879, 1885, 1890;

Figure 3C). Although the latter recognized the

theropod afﬁnity of the tooth, he assigned it to

a new species of basal sauropodomorph, Mas-

sospondylus rawesi (Lydekker 1890). The tooth

was later referred to Megalosaurus (Vianey-

Liaud et al. 1988) and is currently assigned to

an indeterminate theropod, almost certainly an

abelisaurid (Carrano et al. 2010, 2012; C.H. pers.

obs.).

In the span of a decade, between the latest

part of the 19th century and the ﬁrst part of

the 20th century, theropod skeletal material

was reported on three continents of the South-

ern Hemisphere, Africa, South America and

Australia. The French were the ﬁrst to collect

and describe material belonging to Gondwanan

theropods. The ﬁrst deﬁnitive theropod skeletal

remains to be reported in the Southern Hemi-

sphere, in fact, belong to the well-known abelis-

aurid Majungasaurus crenatissimus unearthed

in the Maevarano Formation (Maastrichtian) of

Madagascar. The species was erected as Megalo-

saurus crenatissimus by French paleontologist

Charles Depéret in 1896, based on fossils col-

lected by Mr. Landillon in the Mahajanga Ba-

sin one year before (Depéret 1896a, b; Krause

et al. 2007). Nevertheless, theropod tracks dis-

covered in Cenomanian limestone in the Jebel

Bou-Khaïl (near the city of Laghouat), Algeria,

by French geologist G. Le Mesle were already

reported by Le Mesle & Peron (1880; Figure 3D)

sixteen years before, and account for the ﬁrst

described theropod material from Africa (Ta-

quet 2010; Chabou et al. 2015). Almost twenty

years later, a theropod vertebra and isolated

teeth assigned to Spinosauridae were discov-

ered in the Djoua country (near Timassânine),

Algeria, during a mission led by French ofﬁcer

François Lami and explorer Fernand Foureau in

1898 (Buffetaut 2010; Taquet 2010). Based on

the material collected by the latter, French pa-

leontologist Emile Haug (1904, 1905) reported

on and illustrated the ﬁrst skeletal material of

a theropod (and a dinosaur) from the Sahara,

though the teeth were interpreted as belonging

to an ichthyodectid ﬁsh (Buffetaut 2005, 2010).

The ﬁrst theropod material to be reported in

South America was an isolated tooth described

by famous Argentinian paleontologist Floren-

tino Ameghino in 1899 (Ameghino 1899; Co-

ria & Salgado 1996; Figure 3E-F). Based on this

fragmentary tooth and a partial femur found in

the Upper Cretaceous of Par-Aïk, Shehuen Riv-

er, Santa Cruz Province of Argentina, Ameghi-

no erected the taxon Loncosaurus argentines

which was initially classiﬁed as a megalosaurid

(Ameghino 1899). Although the partial tooth

most likely belongs to a theropod, the femur of

Loncosaurus is that of an ornithopod (Coria &

Salgado 1996). Genyodectes serus, named and

described by Woodward only two years later,

is the ﬁrst valid theropod (and dinosaur) to

be reported from Argentina (Woodward 1901;

Rauhut 2004b). Until the 1970s, this cerato-

saurid remained one of the most complete the-

ropods known from that continent (Rauhut

2004b).

Theropod material is scarcer in Oceania,

yet the ﬁrst representative fossil was reported

in Australia by the beginning of the 20th cen-

tury (Agnolín et al. 2010). Woodward (1906)

described a theropod ungual unearthed from

Cape Patterson, on the south coast of Victoria

(Figure 3G). This claw, the ﬁrst dinosaur materi-

al reported from Australia, was collected by Mr.

W. H. Ferguson in the Wonthaggi Formation

(early Aptian; Agnolín et al. 2010). The pedal

ungual was initially thought to be from a tax-

on closely related to Megalosaurus, and some-

times as Megalosaurus itself (Woodward 1906;

Huene 1926a). It is now considered an indeter-

minate theropod (Agnolín et al. 2010; Carrano

et al. 2012). Four years later, Woodward (1910)

brieﬂy reported the discovery of a tooth and a

posterior caudal vertebra of what he assumed

to be a small megalosaurian theropod. The ma-

terial was found by T.C. Wollaston and comes

from the Griman Creek Formation (Albian)

of Lightning Ridge near Walgett, New South

Wales. Huene (1932) described and referred

the vertebra to the new taxon Walgettosuchus

woodwardi, an indeterminate theropod current-

ly considered a nomen dubium (Agnolín et al.

2010).

Antarctica is the last continent to have yield-

ed non-avian theropod material. The ﬁrst dis-

covery of theropod remains occurred in 1988

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Figure 3. Earliest historical records of theropod remains in A-B, North America; C, Asia; D, Africa; E-F, South America; G,

Oceania; and H, Antarctica. Isolated teeth of: A) Troodon formosus; and B, Deinodon horridus (= Albertosaurus sarcophagus)

from the Upper Cretaceous Judith River of Colorado and ﬁrst reported by Leidy (1856; modiﬁed from Leidy 1860, plate 9);

C) Isolated theropod tooth of ‘Massospondylus rawesi’, an abelisaurid from the Upper Cretaceous of India (localities of Takli

and Maleri) ﬁrst reported by Hislop (1861, 1864, illustration by Lydekker 1890, ﬁg. 1); D) Casts of theropod tracks from

the Upper Cretaceous of Jebel Bou-Khaïl, Algeria, and ﬁrst reported by Le Mesle and Peron (1880; illustration by Mesle &

Peron 1880, ﬁgures 65 and 66); E-F) Isolated theropod tooth from the Upper Cretaceous of Par-Aïk, Argentina, referred to

Loncosaurus argentines and ﬁrst reported by Ameghino (1899); E, illustration by Ameghino (1900, p. 160) and Ameghino

(1906, Figure 8); and F, Huene (1929a, plate 41); G) Pedal ungual of an indeterminate theropod from the Upper Cretaceous

of Cape Patterson, Australia, and ﬁrst reported by Woodward (1906); H) Distal part of a tibia of a megalosauroid? theropod

from the Upper Cretaceous of Col Crame, Antarctica, discovered in 1988 (modiﬁed from Molnar et al. 1996).

Hendrickx et al., Non-Avian Theropods PalArch’s Journal of Vertebrate Palaeontology, 12, 1 (2015)

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when Agelandro Lopez Angriman found the

distal part of a tibia in the Coniacian-Santonian

Hidden Lake Formation of Antarctica (Molnar

et al. 1996; Figure 3H). The bone, known as the

Hidden Lake specimen (Carrano et al. 2012),

comes from the north of Col Crame in the Cape

Lachman region, north-western James Ross Is-

land. This tibia was assigned to an indetermi-

nate tetanuran by Molnar et al. (1996), and to a

megalosauroid by Carrano et al. (2012), which

makes it the latest surviving member of this

clade found to date. Although the Hidden Lake

specimen was the ﬁrst theropod material to be

found in Antarctica, this partial tibia was only

described in 1996, and the ﬁrst theropod to be

reported in the literature is, in fact, Cryolopho-

saurus ellioti, described by Hammer and Hick-

erson (1994) two years earlier. Cryolophosaurus

material was collected during the 1990-1991

and 2003-2004 ﬁeld seasons, and this taxon is

the most complete theropod from Antarctica,

one of the largest from the Early Jurassic, and

possibly one of the earliest tetanurans hitherto

discovered (Smith et al. 2007; Carrano et al.

2012).

History of Classiﬁcation

The clade Dinosauria was erected as a tribe (or

a sub-order) by Richard Owen in 1842 to con-

tain three taxa of large reptiles, Megalosaurus,

Iguanodon, and Hylaeosaurus. Owen (1842) did

not include the already named theropods Poeki-

lopleuron, Streptospondylus, and Suchosaurus,

all considered to be crocodilian taxa at the time.

‘Goniopoda’ was the ﬁrst clade of dinosaurs to

gather two valid theropod dinosaurs. This order

was erected by Edward Drinker Cope in 1866 to

encompass Laelaps (now known as Dryptosau-

rus; Brusatte et al. 2011) and ‘probably’ Mega-

losaurus. ‘Goniopoda’ was, by then, opposed to

the ‘Orthopoda’ consisting of Scelidosaurus, Hy-

laeosaurus, Iguanodon, and Hadrosaurus (Cope

1866).

Although the taxa ‘Goniopoda’ and ‘Orthop-

oda’ were used in Matthew & Brown’s (1922)

classiﬁcation of theropods in the 20th century,

these two groups were abandoned in favor of

clades coined by Othniel Charles Marsh by

the end of the 19th century. Marsh (1881) ﬁrst

erected the taxon Theropoda to contain the

family Allosauridae, initially represented by the

North American genera Allosaurus, Creosaurus,

and Labrosaurus. The term ‘Theropoda’ derived

from the old Greek words θηρίον, thérion mean-

ing ‘wild beast, animal’, and ποδος, pous, podos

meaning ‘foot’. Theropods, with ‘beast feet’ were,

at that time, separated from ornithopods, mean-

ing ‘bird feet’, and sauropods, meaning ‘reptile

feet’, which were coined by Marsh in 1871 and

1878, respectively. A year after naming the tax-

on Theropoda, Marsh (1882) already included

six ‘families’ in this clade, namely Megalosau-

ridae, Zanclodontidae, Amphisauridae, Labro-

sauridae, Coeluridae, and Compsognathidae.

A few years later, Seeley (1887) used the orien-

tation and morphology of the pubis to divide

the clade of Dinosauria into two major groups,

the Saurischia and the Ornithischia. Theropods

and sauropodomorphs were grouped among

saurischian dinosaurs with reptile-like pelves,

whereas ornithischians with bird-like pelves in-

cluded Stegosauria and Ornithopoda. Ironically,

saurischian theropods with beast-like feet and a

reptile-like pelvis ultimately give rise to birds,

instead of the ornithischians with a bird-like

pelvis, and the ornithopods with bird-like feet.

By the end of the 19th century, four currently

valid theropod clades (Ceratosauridae, Mega-

losauridae, Compsognathidae, Omithomimi-

dae), two sauropodomorph (Plateosauridae,

Anchisauridae) and four unrecognized archo-

saur clades (i.e., Labrosauridae, Dryptosauridae,

Coeluridae, and Hallopidae) were gathered into

Theropoda by Marsh (1895, 1896).

The classiﬁcation of theropods was marked-

ly affected by the work of German paleontolo-

gist Friedrich von Huene (1909, 1914a, b, 1923,

1926a, b, 1929b, 1932) in the ﬁrst half of the

20th century. Up until 1932, Huene ignored the

name Theropoda and erected two new clades

to encompass all saurischian dinosaurs, Coe-

lurosauria and ‘Pachypodosauria’. In Huene’s

earlier classiﬁcations, coelurosaurs comprised

theropods such as Coelophysis, Ceratosaurus,

Compsognathus, Proceratosaurus, Tyranno-

saurus, and Ornithomimus, whereas pachypo-

dosaurs included the Carnosauria, consisting

of Megalosaurus, Spinosaurus, and Allosaurus

(formerly known as Antrodemus), as well as the

Prosauropoda and the Sauropoda, two clades

currently classiﬁed as sauropodomorphs. In

the 1930s, Huene (1932) modiﬁed his view on

theropod systematics and abandoned the taxon

‘Pachypodosauria’. At that time, saurischian di-

nosaurs included Coelurosauria, Carnosauria,

Prosauropoda, and Sauropoda, and the separa-

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tion between coelurosaurs and carnosaurs was

mostly based on size (Rauhut 2003a). Among

carnivorous saurischians, coelurosaurs were as-

signed to relatively small, slenderly built, pre-

daceous theropod clades such as Coelophysidae

(formerly known as ‘Podokesauridae’), Comp-

sognathidae, and Ornithomimidae, whereas

carnosaurs encompassed large, heavily built

predators with massive skulls such as Megalo-

sauridae, Spinosauridae, Tyrannosauridae (for-

merly known as ‘Dinodontidae’), and Allosauri-

dae (Huene 1932).

In the beginning of the second half of the

20th century, Alfred Sherwood Romer, in his

authoritative book ‘Osteology of the Reptiles’,

(1956) proposed a slightly modiﬁed version of

the saurischian classiﬁcation. Romer separated

saurischian dinosaurs into Theropoda and Sau-

ropoda, and included all bipedal saurischians

within theropods, including Prosauropoda, Coe-

lurosauria, and Carnosauria. Romer adopted the

size criteria followed by Huene (1932) and re-

stricted carnosaurs to Teratosauridae (now con-

sidered to be a clade of rauisuchian archosaurs;

e.g., Benton 1986), Megalosauridae (represented

at that time by theropods such as Ceratosaurus,

Megalosaurus, Spinosaurus, Allosaurus, Carcha-

rodontosaurus, and Proceratosaurus), and Ty-

rannosauridae. From the 1960s to the beginning

of the 1980s, authors working on theropods, in-

cluding Walker (1964), Colbert (1964), Colbert

& Russell (1969), Ostrom (1976a), and Russell

(1984) did not deviate signiﬁcantly from the

classiﬁcation scheme of Romer (1956). Most of

them, however, did acknowledge that coeluro-

saurs and carnosaurs were likely to be grades

rather than clades (Thulborn 1984). A few

authors like Ostrom (1972), Barsbold (1977),

Welles (1984) and Carroll (1988) abandoned

the size-based dichotomy between coelurosaurs

and carnosaurs, and Barsbold (1977) included

newly erected clades such as Oviraptorosauria

(with Oviraptoridae), Deinonychosauria (with

Dromaeosauridae and Troodontidae, formerly

known as ‘Saurornithoididae’), and Therizino-

sauria (formerly known as ‘Deinocheirosauria’

by Barsbold, 1977, then ‘Segnosauria’ by Bars-

bold and Perle, 1980) among theropods.

The adoption in the early 1980s of phyloge-

netic methodology developed by German ento-

mologist Willi Hennig (1950) in the beginning

of the second half of the 20th century, was a

major step in the history of theropod systemat-

ics, and the results of those cladistic analyses

radically changed prevailing views on theropod

phylogeny. Thulborn (1984) was the ﬁrst to in-

vestigate theropod interrelationships through

a cladistic approach by addressing the system-

atics of Archaeopteryx and other stem-group

birds. Gauthier’s (1986) work on saurischian

interrelationships was the ﬁrst to outline the

current phylogenetic classiﬁcation of non-avian

theropods. Based on a cladistic analysis per-

formed on a data matrix of 84 characters, the

American paleontologist conﬁrmed the mono-

phyly of dinosaurs and corroborated Seeley’s

idea that Sauropodomorpha and Theropoda

were sister-groups within Saurischia. Gauthier

(1986) recovered Theropoda as a well-supported

clade divided into Ceratosauria and Tetanurae,

and provided the modern phylogenetic deﬁni-

tion of theropods as birds and all saurischians

closer to birds than to sauropodomorphs. He

recognized a dichotomy between Carnosauria

and Coelurosauria within tetanuran theropods,

and erected the clade Maniraptora to encom-

pass coelurosaurs more derived than Ornitho-

mimidae. At that time, Ceratosauria contained

Coelophysis, Dilophosaurus, and Ceratosaurus,

carnosaurs included Allosaurus, Acrocanthosau-

rus and tyrannosaurids, and non-avian coeluro-

saurs comprised Compsognathus, Ornitholestes,

and the Ornithomimidae, Caenagnathidae, and

Deinonychosauria (Gauthier 1986).

Since the pioneering work of Gauthier

(1986), the availability of parsimony-based phy-

logenetic software has enabled a large number

of authors to investigate theropod interrelation-

ships via cladistic analysis, resulting in major

revisions to theropod systematics. Novas (1992)

was the ﬁrst to include abelisaurids and tyran-

nosaurids among ceratosaurs and coelurosaurs,

respectively (Rauhut 2003a), and Holtz (1994)

was the ﬁrst major phylogenetic analysis that

recovered the clade Avetheropoda (erected by

Paul 1988 and also known as ‘Neotetanurae’;

Table 1) to include Allosauridae and Coeluro-

sauria (Figure 4). The same year, Sereno et al.

(1994) found that Megalosauroidea (formerly

known as ‘Torvosauroidea’ and ‘Spinosauroi-

dea’) formed the sister group of Avetheropoda

and was divided into Megalosauridae (formerly

known as ‘Torvosauridae’) and Spinosauridae

(Figure 4). Two years later, Sereno et al. (1996)

found the new clade Allosauroidea (also termed

‘Carnosauria’ sensu Padian et al. 1999), which

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PalArch Foundation 11

Figure 4. Cladogram of basal Theropoda showing the relationships of tyrannosauroids and non-coelurosaur theropods. The

phylogenetic classiﬁcation follows the results of the cladistic analyses obtained by Sues et al. (2011) for non-neotheropod

Theropoda, Smith et al. (2007) and Ezcurra & Brusatte (2011) for non-averostran Neotheropoda, Pol & Rauhut (2012) and

Tortosa et al. (2014) for Ceratosauria, Carrano et al. (2012) for non-coelurosaur Tetanurae, Loewen et al. (2013), Lü et al.

(2014) and Porﬁri et al. (2014) for Tyrannosauroidea, and Choiniere et al. (2014b) for basalmost Coelurosauria.

Hendrickx et al., Non-Avian Theropods PalArch’s Journal of Vertebrate Palaeontology, 12, 1 (2015)

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gathered Allosaurus, Sinraptoridae, and Carcha-

rodontosauridae, to be the sister-group of Coelu-

rosauria. Following these preliminary analyses,

Sereno’s (1997, 1998, 1999) major phylogenetic

analyses of dinosaurs proceeded to deﬁne ma-

jor theropod clades such as Neotheropoda, Coe-

lophysoidea, Megalosauroidea, Allosauroidea,

Tyrannosauroidea, Ornithomimosauria (‘Orni-

thomimoidea’ sensu Sereno 1998), Therizino-

sauroidea, Paraves, and Deinonychosauria (Fig-

ure 5).

Subsequent studies on theropod systematics,

whose results are summarized by Holtz (1998),

Rauhut (2003a), Senter (2007), Carrano &

Sampson (2008) and Carrano et al. (2012), bet-

ter resolved the relationships of non-avian

theropods and deﬁned additional clades such

as Noasauridae (Coria & Salgado 1998), Piat-

nitzkysauridae (Carrano et al. 2012), Megarap-

tora (Benson et al. 2010), and Proceratosauridae

(Rauhut et al. 2010; Figures 4-5). In 2015, the

current consensus on non-avian theropod clas-

siﬁcation is based on the results of the most re-

cent large scaled phylogenetic analyses obtained

by Sues et al. (2011) for non-neotheropod The-

ropoda, Smith et al. (2007) and Ezcurra & Bru-

satte (2011) for non-averostran Neotheropoda,

Pol & Rauhut (2012) and Tortosa et al. (2014)

for Ceratosauria, Carrano et al. (2012) for non-

coelurosaur Tetanurae, Loewen et al. (2013), Lü

et al. (2014; which is based on Brusatte et al.,

2010d) and Porﬁri et al. (2014) for Tyrannosau-

roidea, and Godefroit et al. (2013a), Choiniere

et al. (2014b) and Brusatte et al. (2014; the most

updated version of the Theropod Working

Group (TWIG) dataset) for non-tyrannosauroid

Coelurosauria (Figures 4-5).

As noted by Turner et al. (2012), Theropoda

is now comprised of numerous well-supported

‘family’ or ‘super-family’-level subclades that

form a pectinate, ladder-like organization, with

each rung corresponding to a node-based clade

that has not always received a name. Although

the relationships between most theropod clades

are currently well understood, several aspects

of theropod systematics remain controversial.

Current debate occurs over the phylogenetic

placement of Eoraptor and/or herrerasaurids

within non-theropod saurischians (e.g., Langer

& Benton 2006; Alcober & Martinez 2010; Ez-

curra 2010; Martinez et al. 2011; Sereno et al.

2013) or at the base of Theropoda (Nesbitt et

al. 2009; Ezcurra & Brusatte 2011; Nesbitt 2011;

Sues et al. 2011; Langer & Ferigolo 2013; Fig-

ure 4), and over the monophyly or paraphyly of

Coelophysoidea (i.e., Coelophysidae + Dilopho-

sauridae; e.g., Tykoski 2005; Yates 2005; Ezcurra

& Cuny 2007; Ezcurra & Novas 2007; Smith et

al. 2007; Nesbitt et al. 2009; Ezcurra & Brusatte

2011; Xing 2012), and Deinonychosauria (i.e.,

Dromaeosauridae + Troodontidae; e.g., Senter

2011; Turner et al. 2012; Godefroit et al. 2013a,

b; Brusatte et al. 2014; Choiniere et al. 2014b;

Foth et al. 2014; Tsuihiji et al. 2014). Recent de-

bate also focuses on the position of megarap-

torans within neovenatorid allosauroids (Ben-

son et al. 2010; Carrano et al. 2012) or among

tyrannosauroid coelurosaurs (Novas et al. 2013;

Porﬁri et al. 2014; Figure 4).

Current Classiﬁcation

First Theropods

Theropoda can be deﬁned as the most inclusive

clade containing the house sparrow Passer do-

mesticus (Linnaeus 1758) but not the titanosau-

rid sauropod Saltasaurus loricatus Bonaparte

& Powell 1980 (Sereno 2005; Table 1 [see ap-

pendix]). Regardless of the status of inclusion

of Eoraptor and herrerasaurids within Thero-

poda, the oldest deﬁnitive theropod remains

come from the mid-Carnian (early Late Trias-

sic; ~231 Ma) of Argentina (Figure 1). Similar

in age to Eoraptor lunensis (Sereno et al. 2013;

Figure 6A) and the herrerasaurids Herrerasau-

rus ischigualastensis (Sereno & Novas 1994; Fig-

ure 6B) and Sanjuansaurus gordilloi (Alcober &

Martinez 2010), the oldest unquestioned the-

ropod taxon Eodromaeus murphi (Martinez et

al. 2011) is from the Ischigualasto Formation of

San Juan Province. Eodromaeus, Eoraptor and

herrerasaurids were small to large sized (1-6m

long; Sereno & Novas 1992) bipedal sauris-

chians with relatively elongated skulls. These

primitive saurischians retained the ancestral

dinosauromorph habit of obligate bipedality

and the ziphodont dentition present in more

primitive archosauriforms (Holtz et al. 1998;

Barrett et al. 2010; Holtz 2012), leading to their

consideration as carnivorous dinosaurs. Never-

theless, Eoraptor, recently interpreted to be a

basal sauropodomorph (Martinez et al. 2011;

Sereno et al. 2013), exhibits constricted crowns

and pointed denticles that suggest that this

primitive saurischian, as well as the ﬁrst dino-

saurs, might have been omnivorous (Barrett et

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Figure 5. Cladogram of non-tyrannosauroid Coelurosauria showing the relationships of compsognathids and

maniraptoriforms. The phylogenetic classiﬁcation follows the results of the cladistic analyses obtained by Choiniere et

al. (2014b) for basalmost Coelurosauria and Compsognathidae, Longrich & Currie (2009a) and Choiniere et al. (2010b) for

Alvarezsauroidea, Lee et al. (2014) for Ornithomimosauria, Senter et al. (2012a) and Pu et al. (2013) for Therizinosauria,

Lamanna et al. (2014) for Oviraptorosauria, Turner et al. (2012) for Paraves, and Foth et al. (2014) for Avialae.

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Figure 6. Skeletal reconstructions of three non-neotheropod saurischians (and possibly three basalmost theropods). A) The

possible primitive sauropodomorph Eoraptor lunensis; B) The herrerasaurid Herrerasaurus ischigualastensis; C) The very

basal theropod Tawa hallae. Reconstructions by Scott Hartman.

al. 2010; Langer et al. 2010; Sereno et al. 2013).

Tawa hallae (Nesbitt et al. 2009; Figure 6C) and

Daemonosaurus chauliodus (Sues et al. 2011)

from the Norian and possibly Rhaetian of New

Mexico, respectively, are currently recovered be-

tween Eodromaeus and neotheropods (Figure 4).

Unlike Eoraptor, these two recently reported

taxa possess the short subnarial gap present in

basal neotheropods and an antorbital fossa re-

stricted to the vicinity of the antorbital fenestra,

as seen in Herrerasaurus (Nesbitt et al. 2009;

Sues et al. 2011; Langer 2014). This condition

contrasts with the expanded antorbital fossa

of Eoraptor and Eodromaeus. Daemonosaurus

is unique in having a short and tall skull ﬁlled

with procumbent premaxillary and dentary

teeth (Sues et al. 2011). Tawa is closer to coelo-

physoids than Daemonosaurus and other primi-

tive theropods in having an elongated snout

and a more gracile body. Tawa shares with Dae-

monosaurus greatly enlarged maxillary teeth as

well as pneumatic fossae (pleurocoels) in the

cervical vertebrae (Nesbitt et al. 2009; Sues et

al. 2011).

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Coelophysoidea and Dilophosauridae

Neotheropoda (Bakker 1986), the least inclusive

clade containing Coelophysis bauri (Cope 1889)

and Passer domesticus (Linnaeus 1758) (Sereno

2005), currently comprises theropods more de-

rived than Tawa (Nesbitt et al. 2009; Nesbitt

2011; Sues et al. 2011). Among their derived

features, neotheropods are characterized by

an intramandibular joint and a hinge between

the dentary and the postdentary bones (Holtz

2012). Current consensus on basal theropod

phylogeny suggests that neotheropods encom-

pass a basal clade that can be referred to Coe-

lophysoidea and a slightly more derived clade

named Dilophosauridae (sensu Charig & Milner

1990; Figure 4; the clade Dilophosauridae is

here deﬁned phylogenetically for the ﬁrst time,

see Table 1). Dilophosaurus is thought to belong

to Coelophysoidea by some authors (e.g., Car-

rano et al. 2005; Tykoski 2005; Ezcurra & Cuny

2007; Ezcurra & Novas 2007; Xing 2012). How-

ever, results of more recent and/or larger scale

analyses recover Dilophosauridae as a more

derived clade of neotheropods, and the sister-

group of Averostra (e.g., Smith et al. 2007; Nes-

bitt et al. 2009; Ezcurra & Brusatte 2011; Sues et

al. 2011; Ezcurra 2012). Consequently, though

the phylogenetic relationships of dilophosau-

rids remains unresolved, these basal theropods

seem to be more derived than coelophysoids.

Coelophysoidea (sensu Sereno 2005; Table 1)

encompasses small to medium sized theropods

(2-6m long) with slender skulls, and lightly

built, gracile, and elongated bodies character-

ized by elongated cervical centra (Tykoski &

Rowe 2004; Brusatte et al. 2010c; Holtz 2012).

The ﬁrst coelophysoids are already present in

the Norian of Europe (Procompsognathus trias-

sicus, Camposaurus arizonensis; Sereno & Wild

1992; Rauhut & Hungerbühler 1998; Ezcurra &

Brusatte 2011) and North America (Coelophysis

bauri; Colbert 1989; Figure 7A). Although coe-

lophysoids form the ﬁrst radiation of neothero-

pods, they were not apex terrestrial predators in

the Late Triassic, as pseudosuchian carnivores

such as rauisuchians and phytosaurs were larg-

er and more abundant at that time (Brusatte et

al. 2010c; Holtz 2012). Unlike most large pseu-

dosuchian archosaurs, coelophysoids survived

the Triassic/Jurassic boundary, and Jurassic

coelophysoids are known from the Hettangian-

Pliensbachian of China (Panguraptor lufengen-

sis; You et al. 2014), South Africa (Coelophysis

rhodesiensis; Raath 1969, 1977; Bristowe &

Raath 2004), and North America (Coelophysis

kayentakatae; Rowe 1989; n.b., this taxon was

originally coined ‘Syntarsus’ by Rowe, 1989; it

is also referred to as Megapnosaurus by some

authors as the genus name ‘Syntarsus’ was pre-

occupied by a beetle, and the entomologists

Ivie et al. 2001 replaced it with Megapnosaurus;

‘Syntarsus’ is thought to be a junior synonym

of Coelophysis by many authors such as Downs

2000, Bristowe & Raath 2004 and Carrano et al.

2012). Zupaysaurus rougieri (Arcucci & Coria

2003; Ezcurra 2007) and Liliensternus lilienster-

ni (Huene 1934) from the Norian of Argentina

and Germany, respectively, are either classiﬁed

as coelophysoids (e.g., You et al. 2014) or recov-

ered as more derived neotheropods positioned

between Coelophysoidea and Dilophosauridae

(Nesbitt et al. 2009; Ezcurra & Brusatte 2011;

Sues et al. 2011; Ezcurra 2012; Figure 4).

Dilophosauridae is a poorly supported clade

that may contain medium to large sized (4-7m

long) theropods, such as Dilophosaurus wether-

illi (Welles 19841; Figure 7B) and Dracovenator

regenti (Yates 2005) from the Early Jurassic of

North America and South Africa, respectively.

Similar to coelophysoids, these two taxa pos-

sess a subnarial gap and anteriormost maxillary

teeth facing anteroventrally, yet they share with

averostrans a promaxillary fenestra and a re-

duced number of maxillary teeth (Holtz 2012).

The clade has been recovered by some authors

(Yates 2005; Smith et al. 2007; Xu et al. 2009b);

however, an over-atomization of cranial crest

characters may have been leading phylogenetic

analyses to artiﬁcially ﬁnd such a dilophosaurid

clade (Brusatte et al. 2010a). In fact, ‘Dilopho-

saurus’ sinensis (Hu 1993), considered to be a

synonym junior of Sinosaurus triassicus (Dong

2003; Xing et al. 2013a, 2014), and Cryolopho-

saurus ellioti (Smith et al. 2007) from the Early

Jurassic of China and Antarctica, respectively,

were formerly interpreted as dilophosaurid taxa

and are now classiﬁed among basal tetanurans

(Benson 2010a; Brusatte et al. 2010c; Carrano et

al. 2012; Xing 2012). The cranial crest of Dilo-

phosaurus, Cryolophosaurus, and ‘Dilophosau-

rus’ sinensis was convergently acquired in these

taxa and evolved independently in dilophosau-

rids and basal tetanurans (Brusatte et al. 2010a;

Xing 2012), or was a derived feature present in

the common ancestor of dilophosaurids and

basal averostrans. Although relatively common

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PalArch Foundation 16

Figure 7. Skeletal reconstructions of two non-averostran neotheropods and one basal ceratosaur. A) The coelophysoid

Coelophysis bauri; B) The dilophosaurid Dilophosaurus wetherilli; C) The ‘elaphrosaur’ Limusaurus inextricabilis.

Reconstructions by Gregory Paul for Coelophysis and Dilophosaurus (modiﬁed), and Ville Sinkkonen for Limusaurus

(modiﬁed).

and diverse entering the Jurassic, coelophysoids

and dilophosaurids became extinct at or near

the end of Early Jurassic (Carrano & Sampson

2004; Ezcurra & Novas 2007; Langer et al. 2010).

Ceratosauria

Averostra (Paul 2002), the least inclusive clade

containing Ceratosaurus nasicornis Marsh

1884a and Passer domesticus (Linnaeus 1758)

(Allain et al. 2012; Table 1), radiated into two

main clades, Ceratosauria and Tetanurae (Fig-

ure 4). Basal averostrans are characterized by

the oreinirostral condition of their head, de-

ﬁned as a transversally narrow and dorsoven-

trally high skull (Holtz 2012). According to Car-

rano et al. (2012), the derived features shared by

averostrans include a reduced prefrontal which

remains unfused to the postorbital in adults,

the moderate size of the acromion process of

the scapula, a ridge-like medial epicondyle on

the femur, an interpubic fenestra, the subtri-

angular morphology of the distal end of the is-

chium, and a centrally positioned ﬁbular fossa

on the medial surface of the ﬁbula. The ﬁrst

averostrans are known from the Early Juras-

sic and are distributed widely across the globe

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PalArch Foundation 17

with remains found in China (‘Dilophosaurus’

sinensis; Hu 1993), Antarctica (Cryolophosaurus

ellioti; Smith et al. 2007), Africa (Berberosau-

rus liassicus; Allain et al. 2007), South America

(Tachiraptor admirabilis; Langer et al. 2014), and

possibly Europe (‘Saltriosaurus’; Dal Sasso 2003;

Benson 2010b). Ceratosaurs currently include a

basal clade informally referred to as ‘elaphro-

saurs’, a more derived family named Ceratosau-

ridae, and a major clade known as the Abelisau-

roidea (Wilson et al. 2003; Sereno et al. 2004;

Carrano & Sampson 2008; Pol & Rauhut 2012;

Tortosa et al. 2014). ‘Elaphrosaurs’ are a poorly

known group of primitive ceratosaurs including

Elaphrosaurus bambergi from the Kimmeridg-

ian-Tithonian of Tendaguru (Carrano & Samp-

son 2008), Limusaurus inextricabilis from the

Oxfordian of China (Xu et al. 2009b; Figure 7C),

and Spinostropheus gauthieri from the Middle

Jurassic of Niger (Carrano & Sampson 2008;

Rauhut and López-Arbarello 2009; Remes et al.

2009). The ‘elaphrosaur’ clade was retrieved in

all recent cladistic analyses on ceratosaurs (Xu

et al. 2009b; Pol & Rauhut 2012; Farke & Sertich

2013; Tortosa et al. 2014) and always gathers

the taxa Elaphrosaurus and Limusaurus. The lat-

ter is the only ‘elaphrosaur’ known from cranial

material and the only non-maniraptoriform

theropod to possess an edentulous skull conver-

gent with that of ornithomimids (Figure 7C).

Although recovered as ‘elaphrosaurs’ in all re-

cent large scaled cladistic analyses on cerato-

saurs (Pol & Rauhut 2012; Tortosa et al. 2014),

Elaphrosaurus and Limusaurus have also been

suggested to belong to Noasauridae (Canale et

al. 2009; Stiegler et al. 2014).

Ceratosauridae only contains two taxa, the

eponymous Ceratosaurus from the Kimmerid-

gian-Tithonian of North America (C. nasicornis;

Gilmore 1920; Madsen & Welles 2000; Carrano

& Sampson 2008; Figure 8A) and Europe (Cera-

tosaurus sp.; Mateus & Antunes 2000; Malafaia

et al. 2015), and Genyodectes serus from the

Aptian-Albian of Argentina (Rauhut 2004b).

Ceratosaurids were large theropods (6-8m long)

characterized by strongly elongated maxillary

teeth longer than the dentary height, and at least

Ceratosaurus showed a fused nasal horn, two

lacrimal horns, and osteoderms on the dorsal

midline of the animal (Marsh 1884a; Gilmore

1920; Madsen & Welles 2000; Rauhut 2004b).

Along with megalosaurids and allosaurids,

ceratosaurids were apex predators in the Late

Jurassic (Kimmeridgian-Tithonian) ecosystems

of Europe, North America, and possibly South

America and Africa (Henderson 1998; Bakker &

Bir 2004; Soto & Perea 2008; Rauhut 2011).

Abelisauroidea falls into two divergent sub-

clades, the Noasauridae and Abelisauridae (Wil-

son et al. 2003; Sereno et al. 2004; Carrano &

Sampson 2008; Pol & Rauhut 2012; Tortosa et

al. 2014; Figure 4). Noasaurids form a relatively

poorly known group of small, slender abelis-

auroids with forelimbs bearing well-developed

claws (Bonaparte 1991a; Carrano & Sampson

2008; Agnolín & Chiarelli 2010; Carrano et al.

2011). They are only known from the Creta-

ceous and may have already been present in

the Barremian-early Aptian of Argentina (Li-

gabueino andesi; Bonaparte 1996; Carrano &

Sampson 2008). Noasaurids are well-known in

the latest part of the Cretaceous of Gondwana,

having been unearthed in Santonian-Maas-

trichtian deposits in Argentina (Noasaurus

leali, Velocisaurus unicus; Bonaparte & Powell

1980; Bonaparte 1991b, 1996), Madagascar (Ma-

siakasaurus knopﬂeri; Carrano et al. 2002, 2011)

and India (Laevisuchus; Huene & Matley 1933).

Masiakasaurus knopﬂeri (Figure 8B), the best

known noasaurid taxon, shows the peculiar-

ity of having procumbent dentary teeth with a

constriction at the crown base and ﬂutes on the

lingual surface (Carrano et al. 2002, 2011).

Abelisauridae is a well-supported clade of

medium to large (5-9m long) stubby-armed

theropods with short rounded snouts, deep,

heavily sculptured skulls bearing bony protu-

berances and weakly recurved teeth (Bonaparte

1991a; Wilson et al. 2003; Carrano & Sampson

2008; Canale et al. 2009; Pol & Rauhut 2012).

The inclusion of Eoabelisaurus meﬁ (Pol &

Rauhut 2012) from the Aalenian-Bajocian of Pa-

tagonia within abelisaurids is subject of debate

(Pol & Rauhut 2012; Tortosa et al. 2014) and the

ﬁrst deﬁnitive Abelisauridae, Kryptops palaios,

comes from the Aptian-Albian of North Africa

(Sereno and Brusatte 2008). Abelisaurids were

not the dominant predators in Gondwanian

ecosystems in the Early Cretaceous and early

Late Cretaceous of South America and North

Africa, as they were dominated by the larger

spinosaurids and carcharodontosaurids during

that time (Holtz 2012; Novas et al. 2013). Fol-

lowing the extinction and/or decline of Spino-

sauridae and Carcharodontosauridae after the

Cenomanian-Turonian transition, abelisaurids

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PalArch Foundation 18

Figure 8. Skeletal reconstructions of three ceratosaurs. A) The ceratosaurid Ceratosaurus nasicornis; B) The noasaurid

Masiakasaurus knopﬂeri; C) The abelisaurid Majungasaurus crenatissimus. Reconstructions by Scott Hartman.

became apex predators in Africa, Western Eu-

rope, and South America in the latest part of the

Cretaceous (Buffetaut et al. 2005; Candeiro &

Martinelli 2005; Carrano et al. 2012; Novas et al.

2013; Tortosa et al. 2014; Csiki-Sava et al. 2015).

The best-known taxa are from the Campanian-

Maastrichtian of Europe and Gondwana, includ-

ing Majungasaurus crenatissimus (Sampson et

al. 1998; Sampson & Witmer 2007; Figure 8C)

from Madagascar, Aucasaurus garridoi (Coria et

al. 2002), Skorpiovenator bustingorryi (Canale et

al. 2009) and Carnotaurus sastrei (Bonaparte et

al. 1990; Carabajal 2011) from Argentina, Ra-

jasaurus narmadensis (Wilson et al. 2003) from

India, and Arcovenator escotae (Tortosa et al.

2014) from France.

Megalosauroidea

Tetanurae (Gauthier 1986), the most inclusive

clade containing Passer domesticus (Linnaeus

1758) but not Ceratosaurus nasicornis Marsh

1884a (Allain et al. 2012; Table 1), is diagnosed

by an antorbital tooth row, a moderately extend-

ed anterior ramus of the maxilla, a maxillary

fenestra piercing the lateral wall of the maxilla,

separated interdental plates, and a prominent

deltopectoral crest of the humerus (Carrano et

al. 2012). Several relatively complete basal tet-

anurans are known from the Early and Middle

Jurassic of China and Antarctica (i.e., ‘Dilopho-

saurus’ sinensis, Cryolophosaurus, and Mono-

lophosaurus). These primitive tetanurans are

recovered between basal averostrans and the re-

cent clade Orionides which comprises two major

radiations, the Megalosauroidea and Avethero-

Hendrickx et al., Non-Avian Theropods PalArch’s Journal of Vertebrate Palaeontology, 12, 1 (2015)

PalArch Foundation 19

poda (Carrano et al. 2012). The ﬁrst one, Mega-

losauroidea, currently gathers three subclades,

namely the Piatnitzkysauridae, Megalosauridae,

and Spinosauridae (Figure 4). Piatnitzkysauridae

is the sister group of Megalosauria, which is di-

vided into Megalosauridae and Spinosauridae.

Piatnitzkysaurids as currently known comprise

medium sized (5-6m long) American forms such

as Marshosaurus bicentesimus (Madsen 1976a;

Figure 9A) from the Kimmeridgian-Tithonian

of North-America, and Piatnitzkysaurus ﬂoresi

(Bonaparte 1986; Rauhut 2004a) and Condorrap-

tor currumili (Rauhut 2005) from the Toarcian-

Bajocian of Argentina (Cúneo et al. 2013). These

basal megalosauroids are characterized by a

maxilla with a short anterior ramus and vertical-

ly ridged interdental plates (Carrano et al. 2012).

Megalosauridae is a diverse clade of thero-

pods restricted to the Middle to Late Jurassic,

which suggests they went extinct at the Juras-

sic-Cretaceous boundary (Carrano et al. 2012).

Megalosaurids are medium to very large (4-10m

long) theropods exhibiting relatively elongate

skulls that lack cranial protuberances, and pow-

erful arms possibly bearing a large claw at digit

one (Hendrickx et al. 2015; Sadleir et al. 2008;

Benson 2010a; Allain et al. 2012; Carrano et al.

2012). The most primitive and one of the oldest

theropod embryos, found to date, from the Late

Kimmeridgian-Early Tithonian of Portugal have

been ascribed to this clade (Araújo et al. 2013).

Megalosaurids are known as early as the Bajo-

cian of England (Magnosaurus nethercombensis,

Duriavenator hesperis; Benson 2008a, 2010b) and

include forms from the Bajocian-Callovian of

England and France (Megalosaurus bucklandii,

Figure 9B; Dubreuillosaurus valesdunensis; Al-

lain 2002; Benson et al. 2008; Benson 2010a), the

Middle Jurassic of Africa (Afrovenator abakensis;

Sereno et al. 1996), the Late Jurassic of China

(Leshansaurus qianweiensis; Li et al. 2009), and

the Kimmeridgian-Tithonian of North-America

and Portugal (Torvosaurus tanneri, Torvosaurus

gurneyi; Britt 1991; Hendrickx & Mateus 2014a).

Sciurumimus albersdoerferi, a possible megalo-

saurid from the Kimmeridgian of Germany, is

the most complete megalosauroid discovered so

far (Rauhut et al. 2012). It is also currently the

most primitive theropod preserved with direct

evidence of ﬁlamentous integument, indicating

that protofeathers were already covering some

tetanurans early in their evolution (Rauhut et al.

2012).

Spinosauridae, the sister group of Mega-

losauridae, is a well-supported clade of highly

specialized theropods united by an elongated

crocodile-like skull, spatulate snout with sig-

moid alveolar margins, ﬂuted conical teeth with

minute or no denticles, and an hypertrophied

manual ungual (Charig & Milner 1997; Sereno

et al. 1998; Sues et al. 2002; Bertin 2010; Allain

et al. 2012; Ibrahim et al. 2014). These derived

anatomical features, associated with computer

modeling of the skull (Rayﬁeld et al. 2007; Cuff &

Rayﬁeld 2013), oxygen isotope ratios (Amiot

et al. 2010), morphofunctional analysis of the

mandibular articulation (Hendrickx et al. 2008)

and gut contents (Charig & Milner 1997; Buffe-

taut et al. 2004), suggest that spinosaurids were

at least partially piscivorous, while also feed-

ing on dinosaurs and pterosaurs. Spinosaurids

were large to very large theropods (8-17m long)

and include the largest terrestrial predators dis-

covered hitherto. They were also characterized

by elongated neural spines which evolved into

a bony sail in some members (e.g., Spinosaurus

aegyptiacus, Ichthyovenator laosensis; Stromer

1915; Allain et al. 2012; Ibrahim et al. 2014).

Spinosaurid teeth seem to be already present

in the Kimmeridgian-Tithonian of Tanzania

(Buffetaut 2011; but for a different opinion see

Rauhut 2011), yet the earliest deﬁnitive spino-

saurid is currently Baryonyx walkeri (Figure

9C) from the Barremian of England and Portu-

gal (Charig & Milner 1986, 1997; Mateus et al.

2011). Spinosauridae are also known from the

Aptian and/or Albian of Niger (Suchomimus te-

nerensis; Sereno et al. 1998; n.b., Suchomimus te-

nerensis most likely represents the same animal

as the non-diagnostic Cristatusaurus lapparenti

Taquet & Russell 1998 from the same deposits;

Carrano et al. 2012), Brazil (Angaturama limai,

Irritator challengeri; Kellner & Campos 1996;

Sues et al. 2002; n.b., these two taxa known

from non-overlapping cranial material recov-

ered from the same deposits may in fact rep-

resent the same taxon/individual; Sereno et al.

1998; Sues et al. 2002; Dal Sasso et al. 2005) and

South-eastern Asia (Ichthyovenator laosensis; Al-

lain et al. 2012). The most derived spinosaurid,

Spinosaurus aegyptiacus, comes from the Albi-

an-Cenomanian of North Africa (e.g., Stromer

1915; Taquet & Russell 1998; Buffetaut &

Ouaja 2002; Dal Sasso et al. 2005; Ibrahim et

al. 2014). Recent studies have shown that this

taxon had many adaptations for a semi-aquatic

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PalArch Foundation 20

Figure 9. Skeletal reconstructions of three megalosauroids. A) The piatnitzkysaurid Marshosaurus bicentissimus; B) The

megalosaurid Megalosaurus bucklandii; C) The spinosaurid Baryonyx walkeri. Reconstructions by Scott Hartman.

lifestyle, including short hind-limbs, downsized

pelvic girdle, ﬂat-bottomed pedal claws and sol-

id long bones (Ibrahim et al. 2014).

Despite the presence of the tetanuran Chilan-

taisaurus from the Turonian (or younger stage)

of China and considered to be a spinosaurid by

Allain et al. (2012; n.b., Chilantaisaurus is recov-

ered as a neovenatorid allosauroid by Benson

et al. 2010 and Carrano et al. 2012) as well as

isolated teeth tentatively assigned to Spinosau-

ridae from post-Cenomanian deposits of South

America and Asia (Salgado et al. 2009; Hone et

al. 2010; for a different opinion see Hasegawa

et al. 2010), spinosaurids seem to go extinct in

the early Late Cretaceous.

Allosauroidea

Avetheropoda (also known as ‘Neotetanurae’;

e.g., Sereno et al. 1994; Sereno 1998, 1999; Al-

lain et al. 2012; Table 1), the least inclusive clade

containing Allosaurus fragilis Marsh 1877 and

Passer domesticus (Linnaeus 1758) (Allain et al.

2012), is comprised of two major subclades: the

Allosauroidea, and the Coelurosauria (Figure

4). According to Carrano et al. (2012), avethe-

ropods differ from more primitive theropods

by possessing strongly curved chevrons, a

poorly developed ridge on the medial surface

of the ilium, and a subtriangular ﬂange-like ac-

cessory trochanter on the femur. Allosauroids

were dominant terrestrial predators in the Late

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PalArch Foundation 21

Jurassic, Early Cretaceous, and early Late Cre-

taceous worldwide. Allosauroids are currently

divided into four subclades: the Metriacantho-

sauridae, Allosauridae, Neovenatoridae, and

Carcharodontosauridae (Figure 4). Metriacan-

thosauridae (formerly known as ‘Sinraptori-

dae’; Carrano et al. 2012) is the most primitive

and contains forms from the Middle and Late

Jurassic of China such as Sinraptor dongi (Cur-

rie & Zhao 1993a), ‘Yangchuanosaurus’ hepi-

gensis, and Yangchuanosaurus shangyouensis

(Dong et al. 1983). These taxa, which are known

from exceptionally well-preserved skeletons,

share a maxilla with a promaxillary fenestra

larger than the maxillary fenestra, a pneumatic

recess on the lateral surface of the ascending

ramus, and the absence of an anterior ramus

(Currie & Zhao 1993a; Carrano et al. 2012). Me-

triacanthosaurus parkeri (Huene 1923) from

the Oxfordian of England is the only deﬁnitive

non-Asian metriacanthosaurid reported to date

(though Lourinhanosaurus antunesi from the

Kimmeridgian-Tithonian of Portugal may also

be referred to this clade; see Benson 2010a), and

Siamotyrannus isanensis (Buffetaut et al. 1996)

from the Barremian-Aptian of Thailand is the

only known metriacanthosaurid that survived

into the Cretaceous (Carrano et al. 2012).

Allosauridae, a more derived clade of allo-

sauroids and the sister-clade of Carcharodon-

tosauria, is a small group of Kimmeridgian-

Tithonian tetanurans comprising several North

American and Portuguese taxa, namely Allo-

saurus fragilis (Gilmore 1920; Madsen 1976b;

Chure 2000; Loewen 2010; Figure 10A), Allosau-

rus europaeus (Mateus et al. 2006), Allosaurus

n. sp. (Allosaurus jimmadseni sensu Chure 2000;

Chure et al. 2006; Loewen 2010), and Sauropha-

ganax maximus (Chure 1995). Allosaurids were

medium to large (8-10m long) theropods with

thin and dorsally-developed lacrimal horns, and

were one of the dominant predators in Late Ju-

rassic ecosystems of North America and Europe

(Chure 2000; Loewen 2010).

Carcharodontosauria falls into two sub-

clades, the Neovenatoridae and the Carchar-

odontosauridae (Carrano et al. 2012). It has

been debated whether Neovenatoridae is a

monospeciﬁc clade including the taxon Neove-

nator salerii (Brusatte et al. 2008; Figure 10B)

from the Hauterivian-Barremian of England

(Novas et al. 2013; Porﬁri et al. 2014), or a more

inclusive clade including Neovenator and meg-

araptorans (Benson et al. 2010; Carrano et al.

2012; Zanno & Makovicky 2013). According to

Benson et al. (2010), neovenatorids are united

by postcranial synapomorphies such as a short

and broad scapula and a pneumatic ilium. The

recent discovery of a relatively well-preserved

megaraptoran with cranial material, however,

seems to suggest a placement of Megaraptora

within Tyrannosauroidea (Porﬁri et al. 2014).

Though this is still an active debate in theropod

systematics, megaraptorans will be described in

the next section.

Carcharodontosauridae, on the other hand,

forms a well-supported clade comprising me-

dium to very large theropods (6-14m long)

characterized by a massive and deep skull

with sculptured facial bones, and cranial pro-

tuberances on the lacrimals and postorbitals

(Novas et al. 2005, 2013; Coria & Currie 2006;

Brusatte & Sereno 2007; Ortega et al. 2010; Cau

et al. 2013). The earliest carcharodontosaurid

is currently Veterupristisaurus milneri (Rauhut

2011) known from caudal vertebrae from the

Kimmeridgian-Tithonian of Tanzania. In the

Cretaceous, carcharodontosaurids became a

diversiﬁed clade of allosauroids distributed

worldwide. Due to their very large sized, car-

charodontosaurids were at the apex of the food

chain in most ‘mid’ Cretaceous ecosystems. The

best preserved carcharodontosaurids are Con-

cavenator corcovatus (Ortega et al. 2010) from

the Barremian of Spain, Acrocanthosaurus ato-

kensis (Harris 1998; Currie & Carpenter 2000;

Eddy and Clarke 2011) and Tyrannotitan chub-

utensis (Novas et al. 2005; Canale et al. 2015)

from the Aptian-Albian of North America and

Argentina, respectively, Carcharodontosaurus

saharicus (Rauhut 1995; Brusatte & Sereno

2007) from the Cenomanian of North Africa,

Giganotosaurus carolinii (Coria & Salgado 1995;

Calvo and Coria 1998; Figure 10C) and Ma-

pusaurus roseae (Coria & Currie 2006) from

the Cenomanian-?Santonian of Argentina, and

Shaochilong maortuensis (Brusatte et al. 2009,

2010b) from the Turonian of China (Carrano

et al. 2012). The carcharodontosaurid lineage

may have extended to the latest part of the Cre-

taceous in South America as material assigned

to Carcharodontosauridae have been reported

from the Campanian-Maastrichtian of Brazil

(e.g., Candeiro et al. 2012; Azevedo et al. 2013).

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PalArch Foundation 22

Figure 10. Skeletal reconstructions of three allosauroids. A) The allosaurid Allosaurus ‘jimmadseni’; B) The neovenatorid

Neovenator salerii; C) The carcharodontosaurid Giganotosaurus carolinii. Reconstructions by Scott Hartman.

Basal Coelurosauria and Tyrannosauroidea

Coelurosauria (Huene 1914a), the most inclu-

sive clade containing Passer domesticus (Lin-

naeus 1758) but not Allosaurus fragilis Marsh

1877, Sinraptor dongi Currie & Zhao 1993a, and

Carcharodontosaurus saharicus (Depéret & Sa-

vornin 1927) (Sereno 2005), is a well-supported

clade that contains a large diversity of herbivo-

rous and carnivorous non-avian theropods as

well as living birds. According to Turner et al.

(2012), members of this group differ from more

basal theropods by possessing a well-developed

medial shelf on the maxilla, a reversed L-shape

quadratojugal, and amphiplatyan cervical and

anterior dorsal vertebrae. Coelurosaur interre-

lationships are complex, including several well-

deﬁned coelurosaur groups nested in differ-

ent subclades (Figures 4-5). The oldest deﬁnite

coelurosaurs are known from the Bathonian

of Eurasia (Averianov et al. 2010; Rauhut et

al. 2010), though putative coelurosaur remains

have been described from the Early Jurassic

of China (Zhao & Xu 1998; Barrett 2009). The

majority of recent cladistic analyses on coeluro-

saurs recovered Tyrannosauroidea as the basal-

most clade of Coelurosauria (e.g., Dal Sasso &

Maganuco 2011; Senter et al. 2012b; Turner

et al. 2012; Godefroit et al. 2013a; Loewen et

al. 2013; Brusatte et al. 2014; Choiniere et al.

2014b; n.b., Tyrannosauroidea are found more

Hendrickx et al., Non-Avian Theropods PalArch’s Journal of Vertebrate Palaeontology, 12, 1 (2015)

PalArch Foundation 23

derived than Compsognathidae in Zanno et

al. 2009, Rauhut et al. 2010 and Novas et al.

2013). There are, however, several coelurosaur

taxa that fall outside Tyrannosauroidea, at the

very base of Coelurosauria (Figure 4). These

include Aorun zhaoi (Choiniere et al. 2014b)

and Zuolong sallei (Choiniere et al. 2010a) from

the Oxfordian-Callovian of China, Bicentenaria

argentina from the Cenomanian of Argentina

(Novas et al. 2012), and possibly Tanycolagreus

topwilsoni from the Kimmeridgian-Tithonian

of Wyoming and Tugulusaurus faciles from the

?Valanginian-Albian of China (Rauhut & Xu

2005). The latter two are also recovered as sister-

taxa among the Coeluridae, a clade recovered

at the base of Coelurosauria by Li et al. (2010),

but also at the base of the tyrannosauroid clade

(e.g., Dal Sasso & Maganuco 2011; Novas et al.

2012; Senter et al. 2012b; Brusatte et al. 2014),

or slightly more derived than tyrannosauroids

(Choiniere et al. 2014b).

Due to the iconic status of Tyrannosaurus

rex and numerous tyrannosauroid specimens,

tyrannosauroids are the most studied and best

known non-avian theropods (Brusatte et al.

2010d). The recent discovery of a large number

of basal and derived tyrannosauroids has dra-

matically increased the known diversity of this

group, resulting in a well-characterized phylo-

genetic sequence. Tyrannosauroids encompass

small to very large-bodied theropods (3-13m

long) diagnosed by premaxillary teeth signiﬁ-

cantly smaller than anterior maxillary teeth and

with a U-shaped cross-section, small premaxil-

lae with elongated nasal and maxillary (sub-

narial) processes, and fused nasals (Holtz 2004,

2012; Brusatte et al. 2010d). The discovery of

several well-preserved tyrannosauroids from

China has revealed that small to large bodied

primitive forms such as Dilong paradoxus (Xu et

al. 2004) and Yutyrannus huali (Xu et al. 2012)

were covered with ﬁlamentous integument.

Some recent phylogenetic analyses of Tyran-

nosauroidea suggest that three main subclades

radiated independently: the Proceratosauridae,

Megaraptora, and Tyrannosauridae (Figure 4).

The most basal clade, the Proceratosauridae,

comprises small-bodied tyrannosauroids char-

acterized by elaborated cranial crests (Brusatte

et al. 2010d; Figure 11A). Proceratosaurids orig-

inated in the Middle Jurassic of Eurasia, includ-

ing the taxa Proceratosaurus bradleyi (Rauhut

et al. 2010) and Kileskus aristotocus (Averianov

et al. 2010) from the Bathonian of England and

Siberia, respectively. Proceratosaurids are also

known from the Oxfordian of China (Guanlong

wucaii; Xu et al. 2006; Figure 11A), and the

youngest member is Sinotyrannus kazuoensis

(Ji et al. 2009) from the Aptian of China (Bru-

satte et al. 2010d).

Primitive non-proceratosaurid tyrannosau-

roids (i.e., non-proceratosaurid tyrannosauroids

more basal than Tyrannosauridae) encompass

several small to medium sized forms from the

Late Jurassic of Europe (Aviatyrannis jurassica,

Juratyrant langhami; Rauhut 2003b; Benson

2008b; Brusatte & Benson 2013) and North

America (Stokesosaurus clevelandi; Benson

2008b; Brusatte & Benson 2013), and the Early

Cretaceous of Europe (Eotyrannus lengi; Hutt et

al. 2001) and China (Dilong paradoxus, Yutyran-

nus huali, Xiongguanlong baimoensis; Xu et al.

2004, 2012; Li et al. 2010). They are also known

in the Late Cretaceous of North America (e.g.,

Dryptosaurus aquilunguis, Appalachiosaurus

montgomeriensis, Bistahieversor sealeyi; Carr et

al. 2005; Carr & Williamson 2010; Brusatte et al.

2011) and Asia (e.g., Raptorex kriegsteini, Alec-

trosaurus olseni; Mader & Bradley 1989; Sereno

et al. 2009).

Based on the recent description of a relative-

ly complete juvenile specimen of Megaraptor

namunhuaiquii, megaraptorans are thought to

have evolved from primitive tyrannosauroids

more derived than proceratosaurids (Novas et

al. 2013; Porﬁri et al. 2014). Megaraptorans are

gracile theropods characterized by an elongated

skull, and elongated and robust forelimbs with

enlarged thumb claws on digits I and II (Ben-

son et al. 2010; Porﬁri et al. 2014). They were

distributed widely across the globe as they en-

compass Fukuiraptor kitadaniensis (Azuma &

Currie 2000; Currie & Azuma 2006) from the

Barremian of Japan, Australovenator wintonen-

sis (Hocknull et al. 2009) from the Albian of

Australia, and Aerosteon riocoloradensis (Sere-

no et al. 2008) from the Campanian of North

Argentina. Megaraptorans seem also to extend

to the end of the Cretaceous, with Orkoraptor

burkei (Novas et al. 2008) from the Maastrich-

tian of Patagonia as the most recent member of

this clade.

Tyrannosaurids are the most derived and

the largest tyrannosauroids. Within Tyranno-

sauroidea, they show the derived features of

large body size (6-13m long), robust and broad

Hendrickx et al., Non-Avian Theropods PalArch’s Journal of Vertebrate Palaeontology, 12, 1 (2015)

PalArch Foundation 24

Figure 11. Skeletal reconstructions of three tyrannosauroids. A) The proceratosaurid Guanlong wucaii; B) The basal

tyrannosauroid Eotyrannus lengi; C) The tyrannosaurid Tyrannosaurus rex. Reconstructions by Scott Hartman.

skulls with powerful jaws bearing incrassate

teeth with long roots, and reduced forelimbs

ending in two functional ﬁngers (the third digit

is vestigial and does not carry phalanges; Currie

2003; Holtz 2004, 2012; Brusatte et al. 2010d).

Tyrannosaurids were apex predators in all Late

Cretaceous ecosystems of North America and

Asia. They were hypercarnivores and were able

to produce extremely powerful bite forces ca-

pable of crushing bone (Erickson et al. 1996;

Bates and Falkingham 2012). Tyrannosaurids

also possessed a higher degree of stereoscopic

vision than other non-avian theropods, and

their olfactory ratios are particularly high, sug-

gestive of a keen sense of smell (Stevens 2006;

Witmer & Ridgely 2009; Zelenitsky et al. 2009).

Studies have shown that they had accelerated

grow rates and underwent well-characterized

changes during ontogeny (Carr 1999; Erickson

et al. 2004; Horner & Padian 2004). The best

known tyrannosaurids are from the Campan-

ian-Maastrichtian of Asia and North-America

and include Albertosaurus sarcophagus, Gor-

gosaurus libratus, Daspletosaurus torosus, and

Tyrannosaurus rex (Figure 11C) from USA and

Canada (e.g., Russell 1970; Molnar 1991; Brochu

2003; Currie 2003), and Alioramus altai and Tar-

bosaurus baatar from Mongolia (e.g., Hurum &

Sabath 2003; Tsuihiji et al. 2011; Brusatte et al.

2012a).

Hendrickx et al., Non-Avian Theropods PalArch’s Journal of Vertebrate Palaeontology, 12, 1 (2015)

PalArch Foundation 25

Compsognathidae and Ornithomimosauria

Compsognathidae and Ornithomimosauria are

typically recovered as more derived than Tyran-

nosauroidea and more basal than Alvarezsau-

roidea, Therizinosauria, and Oviraptorosauria.

Several recent large scale cladistic analyses

have placed Compsognathidae and Ornitho-

mimosauria as the second and third basalmost

clades of coelurosaurs, respectively (e.g., Csiki

et al. 2010; Senter 2011; Turner et al. 2012;

Godefroit et al. 2013a; Choiniere et al. 2014b;

Figure 5). Compsognathids are characterized

by small body size (1-2.5m long), a gracile and

slender body, and an elongated skull with slen-

der jaws bearing ziphodont teeth (Figure 12A).

Many specimens are immature individuals re-

taining a primitive and unspecialized anatomy,

and Compsognathidae have sometimes been

thought to be paraphyletic, with some comp-

sognathid taxa recovered outside the clade in

phylogenetic analyses by Butler & Upchurch

(2007), Godefroit et al. (2013a), Choiniere et

al. (2014b) and others. Nevertheless, the clade

is strongly supported (i.e., united by 18 un-

ambiguous synapomorphies in Brusatte et al.,

2014), which is currently the largest and most

recent cladistic analyses performed on coeluro-

saurs. According to Brusatte et al. (2014), comp-

sognathids are diagnosed by a dentition with

some unserrated teeth, premaxillary teeth with

a subcircular cross-section, the presence of an

anterior ramus on the maxilla, a vertically ori-

ented pubis shaft, and ossiﬁed sternal plates. In

this study, compsognathids include Juravenator

starki (Chiappe & Göhlich 2010) and Compsog-

nathus longipes (Bidar et al. 1972; Ostrom 1978;

Peyer 2006; Figure 12A) from the Kimmerid-

gian-Tithonian of Germany and Germany and

France, respectively, Mirischia asymmetrica (Na-

ish et al. 2004) from the Albian of Brazil, as well

as Huxiagnathus orientalis (Hwang et al. 2004),

Sinocalliopteryx gigas (Ji et al. 2007a) and Sino-

sauropteryx prima (Currie & Chen 2001; Ji et al.

2007b) from the Barremian-early Aptian Yixian

Formation of China. Aristosuchus pusillus, from

the Barremian of England, and Scipionyx sam-

niticus, from the Albian of Italy, are also consid-

ered as compsognathids by some authors (e.g.,

Naish 2002, 2011; Dal Sasso & Maganuco 2011;

Loewen et al. 2013; Choiniere et al. 2014b). The

latter taxon is remarkable for being a hatchling

specimen preserving exquisitely fossilized soft

tissues and internal organs such as intestines,

muscles, and blood vessels (Dal Sasso & Maga-

nuco 2011). Compsognathid feeding behavior

is among the best known in non-avian thero-

pods, as stomach contents are preserved in both

specimens of Compsognathus (Bidar et al. 1972;

Ostrom 1976a; Evans 1994; Peyer 2006), Hux-

iagnathus (Hwang et al. 2004), Scipionyx (Dal

Sasso & Maganuco 2011), and two specimens

of Sinosauropteryx (Currie & Chen 2001; Ji et al.

2007b) and Sinocalliopteryx (Xing et al. 2012).

These reveal that compsognathids ingested

gastroliths and had an extremely diverse diet

composed of ﬁsh, lizards, non-avian theropods

(dromaeosaurids), primitive birds, and mam-

mals. Similar to more basal tetanurans, evidenc-

es of ﬁlamentous integument in well-preserved

compsognathids such as Sinosauropteryx (Cur-

rie & Chen 2001) and Juravenator (Chiappe &

Göhlich 2010) suggest that protofeathers par-

tially or extensively covered the body of these

basal coelurosaurs. A recent study on the fossil-

ized melanosomes in Sinosauropteryx has also

revealed that the tail of this animal had stripes

which exhibited chestnut to rufous (reddish-

brown) tones (Zhang et al. 2010).

Maniraptoriformes (Holtz 1995), the least

inclusive clade containing Passer domesti-

cus (Linnaeus 1758) and Ornithomimus velox

Marsh 1890 (Maryańska et al. 2002), is largely

composed of non-strictly-carnivorous thero-

pods that are partially or fully edentulous and/

or possess reduced lanceolate crowns, with a

few derived maniraptoriforms (i.e., dromaeo-

saurids) being secondarily carnivorous (Holtz

2012). The ﬁrst radiation of non-strictly car-

nivorous (i.e., herbivorous to omnivorous; see

Barrett 2005; Zanno & Makovicky 2011; Lee et

al. 2014) coelurosaurs were ornithomimosaurs.

The latter are small to very large (2-10m long)

lightly to heavily built theropods character-

ized by a low and delicate skull, slender neck,

elongate forehands bearing three non-raptorial

clawed ﬁngers, and in the ostrich-like ornitho-

mimids long powerful legs that were adapted

for rapid locomotion (Russell 1972; Nicholls

& Russell 1981; Makovicky et al. 2004; Barrett

2005; Kobayashi & Barsbold 2005a; Liyong et

al. 2012; Figure 12B). The jaws of basal orni-

thomimosaurs bear a large number of small

conical teeth, intermediate taxa possess small

teeth restricted to the anterior extremity of

the dentary and derived forms are fully eden-

tulous, possessing only a rhamphotheca (some

Hendrickx et al., Non-Avian Theropods PalArch’s Journal of Vertebrate Palaeontology, 12, 1 (2015)

PalArch Foundation 26

Figure 12. Skeletal reconstructions of three basal maniraptoriforms. A) The compsognathid Compsognathus longipes; B)

The ornithomimosaur Deinocheirus miriﬁcus; C) The basal maniraptoran Ornitholestes hermanni. Reconstructions by Scott

Hartman.

exhibit columnar structures that may have

been used as a ﬁlter-feeding system; Norell et

al. 2001; for an alternative hypothesis, see Bar-

rett 2005). Some derived ornithomimids pos-

sessed ﬁlamentous protofeathers and possibly

long shafted feathers (pennibrachium) on the

forearms, forming wings (Zelenitsky et al. 2012;

for a different opinion see Foth et al. 2014). Or-

nithomimosaurs originated in the earliest part

of the Cretaceous and the oldest and basalmost

member of the clade is Nqwebasaurus thwazi

from the Berriasian-Valanginian of South Af-

rica (Choiniere et al. 2012). Pelecanimimus poly-

odon (Pérez-Moreno et al. 1994), another basal

ornithomimosaur taxon from the Hauterivian

of Spain, possessed more than 200 unserrated

teeth on the jaws, which makes it the theropod

bearing the largest number of teeth. More de-

rived ornithomimosaurs with dentulous lower

jaws are mostly known from the Valanginian-

Albian of China such as Hexing qingyi (Liyong et

al. 2012), Beishanlong grandis (Makovicky et al.

2010), Shenzhousaurus orientalis (Ji et al. 2003),

and Harpymimus okladnikovi (Kobayashi &

Barsbold 2005b). Edentulous ornithomimo-

saurs are only known from the Upper Creta-

Hendrickx et al., Non-Avian Theropods PalArch’s Journal of Vertebrate Palaeontology, 12, 1 (2015)

PalArch Foundation 27

ceous of Asia and North America, and the best

known taxa are Garudimimus brevipes (Ko-

bayashi & Barsbold 2005a) and Sinornithomi-

mus dongi (Kobayashi and Lü 2003) from the

early Late Cretaceous of China, Ornithomimus

edmontonicus and Struthiomimus altus (Rus-

sell 1972) from the Campanian-Maastrichtian

of Canada, and Gallimimus bullatus (Osmólska

et al. 1972) and Deinocheirus miriﬁcus (Lee et

al. 2014; Figure 12B) from the Maastrichtian of

Mongolia. The latter was recently revealed to be

a very large omnivorous ornithomimosaur with

a deep jaw, tall neural spines, elongated fore-

limbs and short hind limbs. It was recovered

as a derived member of a new lineage of Asian

ornithomimosaurs known as Deinocheiridae

(Lee et al. 2014). Deinocheirids, which include

Beishanlong, Garudimimus and Deinocheirus,

do not seem to be adapted for speed, in con-

trast to cursorial ornithomimids such as Gal-

limimus, Struthiomimus and Ornithomimus that

are widely interpreted as fast runners (Russell

1972; Thulborn 1990; Lee et al. 2014).

Therizinosauria, Alvarezsauroidea and Ovirap-

torosauria

Maniraptora (Gauthier 1986), the most-inclu-

sive clade containing Passer domesticus (Lin-

naeus 1758) but not Ornithomimus edmon-

tonicus Marsh 1890 (Maryańska et al. 2002),

includes theropods characterized by a well-de-

veloped lateral process of the quadrate, a large

bony sternum with co-ossiﬁed sternal plates,

and a semilunate carpal (Holtz 2012; Turner et

al. 2012). Many maniraptorans convergently

acquired a retroverted pubis superﬁcially simi-

lar to ornithischians (Holtz 2012). Ornitholestes

hermanni (Osborn 1903; Carpenter et al. 2005;

Figure 12C) from the Upper Jurassic of North

America is either recovered as the basalmost

maniraptoran (e.g., Dal Sasso & Maganuco

2011; Novas et al. 2012; Senter et al. 2012b;

Turner et al. 2012; Foth et al. 2014) or as a basal

coelurosaur closely related to some compsog-

nathids (e.g., Godefroit et al. 2013a; Choiniere

et al. 2014b). The basalmost clade within Mani-

raptora is the Alvarezsauroidea (Figure 5). Al-

varezsauroids were small (1-2.5m long) coelu-

rosaurs characterized by a gracile and low skull

with large cranial openings, elongate rostrum,

and slender jaws bearing a large number of

teeth that are, at least for some crowns, lanceo-

late (Figure 13A). The forelimbs of alvarezsau-

roids bear three ﬁngers in which digit II and

III are reduced in size and were even lost in

some derived taxa (Perle et al. 1993; Chiappe

et al. 1998; Longrich & Currie 2009a; Choiniere

et al. 2010b, 2014a; Nesbitt et al. 2011; Xu et al.

2011b). The basalmost member is Haplocheirus

sollers from the Oxfordian of China; all alva-

rezsauroids more derived than Haplocheirus be-

long to Alvarezsauridae (Choiniere et al. 2010a,

2014a). Alvarezsaurids are restricted to the Late

Cretaceous of North-America, South-America,

Asia, and Europe (Naish and Dyke 2004; Lon-

grich & Currie 2009a; Agnolín et al. 2012; Xu

et al. 2013). They comprise taxa with a large

number of minute and lanceolate crowns, short

forelimbs bearing either a single ﬁrst digit, or a

hypertrophied thumb, in both case ended by a

massive claw used for digging, a pubis oriented

backward, and elongated hind limbs adapted

for cursoriality. The best known members are

Patagonykus puertai (Novas 1997a) from the

Turonian-Coniacian of Argentina, and the par-

vicursorines Xixianykus zhengi (Xu et al. 2010b)

and Linhenykus monodactylus (Xu et al. 2011b,

2013) from the Coniacian-Santonian and Cam-

panian of China, respectively, and Mononykus

olecranus (Perle et al. 1993, 1994), Shuvuuia de-

serti (Chiappe et al. 1998; Suzuki et al. 2002; Fig-

ure 13A), and Ceratonykus oculatus (Alifanov &

Barsbold 2009) from the Campanian-Maastrich-

tian of Mongolia. At least one member of this

group (i.e., Shuvuuia) possessed ﬁlamentous

integuments similar to those of more primitive

tetanurans (Schweitzer et al. 1999).

Therizinosaurs are small to very large (2-

10m long) ‘prosauropod’-like theropods char-

acterized by a small head bearing reduced and

basally constricted crowns, an elongated neck,

long and robust arms terminated by large

claws, broad abdomen and pelvis, and a rela-

tively vertical position of the body (Barsbold &

Perle 1980; Clark et al. 2004; Zanno 2010a, b;

Lautenschlager et al. 2014; Figure 13B). Therizi-

nosaurs seem to be restricted to North America

and Asia in the Cretaceous, yet the therizinosaur

Eshanosaurus deguchiianus, said to be found in

the Lower Lufeng Formation of the Yunnan

Province, China, may attest to the presence of

the clade back to the Lower Jurassic (Zhao &

Xu 1998; Xu et al. 2001a). However, given the

fact that the time separating this taxon from the

most basal therizinosaur is anomalous, an Early

Jurassic age of Eshanosaurus requires conﬁrma-

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PalArch Foundation 28

Figure 13. Skeletal reconstructions of three basal maniraptorans. A) The alvarezsauroid Shuvuuia deserti; B) The

therizinosauroid Nothronychus graffami; C) The oviraptorosaur Khaan mangas. Reconstructions by Scott Hartman.

tion (Kirkland et al. 2005; Barrett 2009). The

most primitive known member is currently Fal-

carius utahensis (Zanno 2006, 2010b) from the

Barremian of Utah. Jianchangosaurus yixianen-

sis (Pu et al. 2013) and Beipiaosaurus inexpectus

(Xu et al. 1999a) are two basal therizinosaurs

from the Early Cretaceous (Barremian?) of Chi-

na that are slightly more derived than Falcarius.

The body of these two primitive therizinosaurs

was covered with ﬁlamentous integument (Xu

et al. 2009a; Pu et al. 2013), which suggests that

most, if not all, therizinosaurs had protofeath-

ers. Therizinosaur taxa more derived than Ji-

anchangosaurus form the clade Therizinosau-

Hendrickx et al., Non-Avian Theropods PalArch’s Journal of Vertebrate Palaeontology, 12, 1 (2015)

PalArch Foundation 29

roidea (Pu et al. 2013). Jianchangosaurus and

therizinosauroids share a downturned anterior

extremity of the dentary, large apically inclined

denticles of the crowns, and an edentulous pre-

maxilla bearing a rhamphotheca (which may

not be present in Falcarius). Derived therizino-

sauroids (therizinosaurids sensu Zanno et al.

2009; Table 1) possess important basicranial

pneumaticity, long scythe-like manual unguals,

and a ﬂattened pubic shaft (Zanno 2010a). The

best known therizinosauroids are Alxasaurus

elesitaiensis (Russell & Dong 1993) from the

Albian of China, Nothronychus graffami (Zanno

et al. 2009; Figure 13B) from the Turonian of

Utah, Erlikosaurus andrewsi and Segnosaurus

galbinensis (Barsbold & Perle 1980; Barsbold

1983; Clark et al. 1994; Lautenschlager et al.

2014) from the Cenomanian-Turonian of Mon-

golia, and Neimongosaurus yangi (Zhang et al.

2001) from the Campanian-Maastrichtian of

China (Zanno 2010a).

The clade containing theropods more de-

rived than therizinosaurs, including Ovirapto-

rosauria and Paraves, has recently been named

Pennaraptora based on deﬁnitive evidence of

pennaceous feathers in multiple pennarap-

toran taxa (Foth et al. 2014). Oviraptorosauria

is a well-supported clade of small to large (1-

8m long) theropods easily recognized by their

short skulls with parrot-like beaks, forelimbs

with elongated manual ﬁngers, and short tails

(Clark et al. 2001; Osmólska et al. 2004; Bala-

noff et al. 2009; Longrich et al. 2010; Balanoff &

Norell 2012; Lamanna et al. 2014; Figure 13C).

Oviraptorosaurs are restricted to the Cretaceous

of Asia, North America and possibly South

America (Frey & Martill 1995; Frankfurt & Chi-

appe 1999; for a different opinion, see Agnolín

& Martinelli 2007), and most taxa come from

Campanian-Maastrichtian deposits. Members

of this clade were partially to strictly herbivo-

rous coelurosaurs who adopted an avian-like

brooding posture on their nests (Clark et al.

1999; Varricchio et al. 2008; Zanno & Makov-

icky 2011). Similar to ornithomimosaurs, basal

oviraptorosaurs retained teeth that were subse-

quently lost in more derived taxa; the majority

of oviraptorosaur taxa, which form the clade

Caenagnathoidea, were edentulous. The basal-

most oviraptorosaur is currently Incisivosaurus

gauthieri from the Aptian of China (Balanoff

et al. 2009). Incisivosaurus shows the primi-

tive condition of having dentulous maxillae

and dentaries, and the peculiarity of bearing

premaxillary teeth that are much larger than

the lateral teeth (Xu et al. 2002a; Balanoff et

al. 2009). Contemporaneous, yet more derived,

non-caenagnathoid oviraptorosaurs such as

Caudipteryx zoui and Similicaudipteryx yixi-

anensis from China retained only premaxillary

teeth, and several well-preserved specimens

possessed branching feathers such as remiges

on the forelimbs, and rectrices on the caudal

vertebrae (Ji et al. 1998; Zhou et al. 2000; He

et al. 2008; Xu et al. 2010a). This suggests that

some, if not all oviraptorosaurs had feathered

bodies and wings, yet they appear entirely

ﬂightless. Caenagnathoidea is divided into two

main subclades, Oviraptoridae and Caenag-

nathidae (Osmólska et al. 2004; Longrich et al.

2013; Lamanna et al. 2014). Caenagnathids are

characterized by fused dentaries and long, shal-

low pneumatized mandibles, whereas ovirapto-

rids had deep lower jaws and an external naris

extending back and over the antorbital fenes-

tra (Longrich et al. 2013). Oviraptorids such as

Khaan mckennai (Clark et al. 1999; Balanoff &

Norell 2012; Figure 13C) inhabited xeric envi-

ronments (i.e., deserts) whereas caenagnathids

occurred in ﬂuvial-dominated and costal ﬂood-

plain environments (Longrich et al. 2013). Taxa

from both clades convergently acquired cranial

crests, as shown in Citipati osmolskae (Clark

et al. 2002), Nemegtomaia barsboldi (Lü et al.

2004), and Anzu wyliei (Lamanna et al. 2014).

Paraves

The remaining maniraptorans, comprising birds

and two non-avian theropod clades traditionally

labeled deinonychosaurs, are grouped within

Paraves (Figure 5). The latter is deﬁned as the

most inclusive clade containing Passer domes-

ticus (Linnaeus 1758) but not Oviraptor philoc-

eratops (Holtz & Osmólska 2004). Deinonycho-

sauria, on the other hand, is either deﬁned as a

node-based clade containing the last common

ancestor of Troodon formosus and Velociraptor

mongoliensis and all of its descendants (Turner

et al. 2012), or the most-inclusive clade contain-

ing Dromaeosaurus albertensis but not Passer

domesticus (Linnaeus 1758) (Godefroit et al.

2013a). Deinonychosaurs are typically divided

into Dromaeosauridae and Troodontidae, thero-

pods that share a raptorial sickle-shaped claw on

the hyperextendable pedal digit II (Holtz 2012;

Turner et al. 2012). Deinonychosauria was con-

Hendrickx et al., Non-Avian Theropods PalArch’s Journal of Vertebrate Palaeontology, 12, 1 (2015)

PalArch Foundation 30

sidered a well-supported clade until recently

(e.g., Turner et al. 2012), but newly discovered

basal paravians and the description of addi-

tional specimens of Archaeopteryx (Mayr et al.

2005; Foth et al. 2014) have led to analyses that

ﬁnd troodontids more closely related to avialans

than to dromaeosaurids, rendering the taxon

Deinonychosauria paraphyletic or equivalent to

Dromaeosauridae, depending on the phylogenet-

ic deﬁnition given to this clade (e.g., Godefroit

et al. 2013a; Brusatte et al. 2014; Choiniere et al.

2014b; Foth et al. 2014; Figure 5).

Dromaeosaurids are the only deﬁnitively

carnivorous maniraptoriforms (with perhaps

the exception of Ornitholestes). They share un-

constricted ziphodont teeth and a hinge joint

(ginglymus) on the distal end of metatarsal

II that permits an extended range of motion

in the second toe and its hypertrophied and

highly modiﬁed claw (Norell & Makovicky

2004; Turner et al. 2012). Dromaeosaurids are

a widespread group of very small to large bod-

ied (0.6-7m long) paravians that were present

on all continents by the Late Cretaceous. Al-

though isolated teeth from the Late Jurassic

of Europe have been assigned to members of

this clade (e.g., Zinke 1998; Lubbe et al. 2009;

Hendrickx & Mateus 2014b) and the presence

of dromaeosaurids in the Jurassic is evidenced

by the appearance of closely related paravians

in the Late Jurassic (Figure 1), deﬁnitive drom-

aeosaurids currently range from the Barremian

(China) to the Maastrichtian (North America).

A large array of evidence indicates that some,

and most likely all Dromaeosauridae were cov-

ered with ﬁlamentous integuments, and at least

two dromaeosaurid taxa (i.e., Microraptor and

Changyuraptor) possessed four wings (i.e., pen-

naceous fore- and hind limbs) with branching

feathers like those seen in extant birds (Xu et al.

1999b, 2001b, 2003; Ji et al. 2001; Turner et al.

2007; Han et al. 2014). The majority of recent

phylogenetic analyses performed on paravians

typically recover three dromaeosaurid subclades:

Unenlagiinae, Microraptorinae, and Eudromaeo-

sauria (e.g., Senter et al. 2012b; Turner et al. 2012;

Brusatte et al. 2014; Choiniere et al. 2014b; Foth

et al. 2014). A different topology was obtained

by Agnolín & Novas (2013) who found Microrap-

torinae and Unenlagiinae outside Dromaeosauri-

dae and gathered within the new clade ‘Averap-

tora’ with Avialae, a conﬁguration not recovered

by other theropod workers.

Although Agnolín & Novas (2011, 2013)

have defended an avialan afﬁnity of unenlagi-

ines, it is commonly accepted that Unenlagiinae

was the ﬁrst dromaeosaurid radiation and is the

most basal lineage of Dromaeosauridae. These

primitive dromaeosaurids are characterized by

an elongate rostrum, unserrated teeth, and a

vertically oriented pubis (Gianechini & Apesteg-

uía 2011; Gianechini et al. 2011; Turner et al.

2012; Figure 14A). They are exclusively found

in the Upper Cretaceous of Gondwana, and are

mostly known from South America. The best

preserved forms are Buitreraptor gonzalezorum

from the Cenomanian (Makovicky et al. 2005;

Figure 13A), Unenlagia comahuensis from the

Turonian-Coniacian (Novas & Puerta 1997),

Austroraptor cabazai from the Maastrichtian

of Argentina (Novas et al. 2009), and Rahonavis

ostromi from the Maastrichtian of Madagascar

(Forster et al. 1998; Turner et al. 2012).

The remaining dromaeosaurids are distrib-

uted among three subclades, Microraptorinae

(‘Microraptoria’ sensu Senter et al. 2004, 2012b),

Velociraptorinae and Dromaeosaurinae (Turner

et al. 2012; Figure 5). As suggested by the ety-

mology, microraptorines were small to very

small (0.6-2m long) dromaeosaurids thought

to have aerial or subaerial abilities (i.e., gliding,

powered ﬂight, or other semi-aerial locomo-

tion) that are known from the Early to Late Cre-

taceous of China and North America (Xu et al.

2003; Longrich & Currie 2009b; Han et al. 2014).

The best preserved members of this clade, all

from the Early Cretaceous of Liaoning in China,

are Microraptor sp. (Hwang et al. 2002; Xu et al.

2003; O’Connor et al. 2011; Xing et al. 2013b),

Sinornithosaurus millenii (Xu et al. 1999b; Xu

and Wu 2001; Gong et al. 2010), Tianyuraptor

ostromi (Zheng et al. 2010), and Changyuraptor

yangi (Han et al. 2014). Hesperonychus eliza-

bethae, from the Campanian of Alberta, is the

youngest known microraptorine, and the only

one found outside China (Longrich & Currie

2009b). Velociraptorinae includes North Ameri-

can, Asian and possibly European dromaeosau-

rids, which are characterized by pleurocoels in

all dorsal vertebrae (Turner et al. 2012). Velo-

ciraptorines encompass the famous theropods

Velociraptor mongoliensis from the Campanian

of Mongolia (Sues 1977; Norell & Makovicky

1997, 1999; Barsbold and Osmólska 1999), Dei-

nonychus antirrhopus from the Aptian-Albian

of Montana (Ostrom 1969, 1976b), and Bambi-

Hendrickx et al., Non-Avian Theropods PalArch’s Journal of Vertebrate Palaeontology, 12, 1 (2015)

PalArch Foundation 31

Figure 14. Skeletal reconstructions of three basal paravians. A) The unenlagiine dromaeosaurid Buitreraptor gonzalezorum;

B) The velociraptorine dromaeosaurid Deinonychus antirrhopus; C) The troodontid Troodon formosus. Reconstructions by

Scott Hartman.

raptor feinbergi from the Campanian of Mon-

tana (Burnham et al. 2000; Burnham 2004).

While Balaur bondoc, from the Maastrichtian

of Romanian, may represent the only deﬁnitive

velociraptorine from Europe (Csiki et al. 2010;

Brusatte et al. 2013, 2014), two recent large

scale phylogenetic analyses on coelurosaurs re-

covered it as a basal avialan (i.e., Godefroit et

al. 2013a; Foth et al. 2014) and the position of

this taxon among paravians remains unclear.

Dromaeosaurinae, the remaining subclade of

dromaeosaurids, includes small to large-sized

theropods with a lateral dentition bearing me-

sial denticles, a ventrodorsally tall jugal process

of the maxilla, and a vertically oriented pubis

(Turner et al. 2012). This clade is mostly com-

prised by North American dromaeosaurid taxa

such as Utahraptor ostrommaysi from the Bar-

remian of Utah (Kirkland et al. 1993), Dromaeo-

saurus albertensis (Colbert & Russell 1969; Cur-

rie 1995; Figure 14B) and Atrociraptor marshalli

(Currie & Varricchio 2004) from the Campanian

Hendrickx et al., Non-Avian Theropods PalArch’s Journal of Vertebrate Palaeontology, 12, 1 (2015)

PalArch Foundation 32

of Alberta. Achillobator gigantibus (Perle et al.

1999) from the Cenomanian-Santonian of Mon-

golia also attests the presence of dromaeosau-

rines in central Asia in the Late Cretaceous.

Troodontidae is a clade of lightly built non-

avian maniraptorans with taxa that rank among

the smallest non-avian body sizes and the high-

est encephalization quotients (Makovicky &

Norell 2004; Lü et al. 2010; Zanno et al. 2011;

Tsuihiji et al. 2014). Troodontids share an an-

teroventrally inclined quadrate and jaws with

a large number of small, constricted teeth set

in an open groove in the dentary (Makovicky &

Norell 2004; Turner et al. 2012). The crowns

are unserrated in basalmost forms and bear

very large hooked denticles in derived taxa,

which suggests an herbivorous diet in primitive

troodontids and a carnivorous or omnivorous

diet in advanced forms bearing serrated teeth

(Currie 1987; Holtz et al. 1998; Currie & Dong

2001; Lü et al. 2010; Zanno & Makovicky 2011).

Troodontids are known from the Cretaceous

of Asia, North America, Europe, and possibly

from the Late Jurassic of China, depending on

the troodontid afﬁnities of newly discovered

forms such as Anchiornis, Xiaotingia, and Eos-

inopteryx (Makovicky & Norell 2004; Hu et al.

2009; Vullo & Néraudeau 2010; Xu et al. 2011a;

Turner et al. 2012; Godefroit et al. 2013b; Bru-

satte et al. 2014). Isolated teeth from the Late

Jurassic of North America and Portugal, and

the Late Cretaceous of India, have also been

assigned to Troodontidae (Chure 1994; Zinke

1998; Goswami et al. 2013). If Troodon formo-

sus from the Campanian of Canada is the most

famous troodontid and the ﬁrst to be discov-

ered (Russell 1948; Currie 1985, 1987; Currie &

Zhao 1993b; Figure 14C), the best preserved

troodontid taxa all come from the Cretaceous of

Asia. They include Sinusonasus magnodens (Xu &

Wang 2004), Mei long (Xu & Norell 2004; Gao

et al. 2012), and Sinovenator changii (Xu et al.

2002b) from the Early Cretaceous of China, and

Byronosaurus jaffei (Norell et al. 2000; Makov-

icky et al. 2003), Gobivenator mongoliensis (Tsui-

hiji et al. 2014), Saurornithoides mongoliensis

and Zanabazar junior (Barsbold 1974; Norell et

al. 2009) from the Campanian of Mongolia.

The recent discovery of a large number of

paravian taxa closely related to birds such as

Anchiornis huxleyi (Hu et al. 2009; Figure 15A),

Xiaotingia zhengi (Xu et al. 2011a), Aurornis xui

(Godefroit et al. 2013a), and Eosinopteryx brevi-

penna (Godefroit et al. 2013b), all from the Mid-

dle to Late Jurassic of the Tiaojishan Formation

of China, have brought new data to bear on the

relationships of the earliest avian theropods.

According to two of the most recent phyloge-

netic analyses (e.g., Godefroit et al. 2013a; Foth

et al. 2014), the latter taxa are gathered within

Avialae, the most-inclusive clade containing

Passer domesticus (Linnaeus 1758) but not

Dromaeosaurus albertensis Matthew & Brown

1922 or Troodon formosus Leidy 1856 (Gode-

froit et al. 2013a). Yet, these taxa were recovered

in the same clade at the base of Troodontidae in

another large scaled phylogenetic analyses (i.e.,

Brusatte et al. 2014), and their exact position

among paravians remains unsettled. For de-

cades, the most basal and earliest avialan taxon

was considered to be Archaeopteryx sp. (Figure

15B), but with the inclusion of these recently re-

ported paravians from the Tiaojishan Formation

into cladistic analyses, Archaeopteryx’s system-

atic position has become uncertain. Currently,

Archaeopteryx is either recovered as the basal-

most avialan (e.g., Turner et al. 2012; Agnolín &

Novas 2013; Brusatte et al. 2014a; Choiniere et

al. 2014b), a deinonychosaur closely related to

troodontids and dromaeosaurids (e.g., Xu et al.

2011a; Godefroit et al. 2013b; Xu & Pol 2014),

or an avialan theropod more derived than the

basalmost avialans Aurornis and Anchiornis

(e.g., Godefroit et al. 2013a; Foth et al. 2014).

The anatomical distinctions between non-avian

and avian theropods are, therefore, particularly

subtle and vary according to the phylogenetic

analysis performed by authors. For instance,

in one of the largest and most recent cladistic

analyses provided by Foth et al. (2014) on coe-

lurosaurs, avialan synapomorphies include the

presence of roots of dentary teeth subcircular in

cross-section, extensive contact between pubes,

humerus and femur subequal in thickness, and

dorsal margin of the antorbital fossa formed by

lacrimal and nasal. In another large scaled phy-

logenetic analysis on coelurosaurs performed

by Brusatte et al. (2014a), Avialae are diagnosed

by asymmetrical feathers on forelimbs, unfused

parietals, less than 26 caudal vertebrae, a dorsal

margin of the antorbital fossa formed by lacri-

mal and nasal, and a humerus longer than the

femur.

A similar situation occurs with Scansoriop-

terygidae and their unsettled phylogenetic po-

sition within Pennaraptora. Scansoriopterygids

Hendrickx et al., Non-Avian Theropods PalArch’s Journal of Vertebrate Palaeontology, 12, 1 (2015)

PalArch Foundation 33

Figure 15. Skeletal reconstructions of three basal avialan? theropods. A) The basal avialan Anchiornis huxleyi; B) The

archaeopterygid Archaeopteryx sp.; C) The scansoriopterygid Epidendrosaurus ninchengensis. Reconstructions by Ville

Sinkkonen for Anchiornis and Scott Hartman for Archaeopteryx and Epidendrosaurus.

Hendrickx et al., Non-Avian Theropods PalArch’s Journal of Vertebrate Palaeontology, 12, 1 (2015)

PalArch Foundation 34

form an aberrant subclade of very small-sized

maniraptorans (the only subadult specimen of

Scansoriopterygidae has a body length of less

than 30 centimeters; Zhang et al. 2008) char-

acterized by a short and high skull, a small

number of procumbent teeth restricted to the

anterior portion of the jaws, propubic pelvis,

and elongated ribbon-like tail-feathers (Zhang

et al. 2002, 2008; Agnolín & Novas 2013). Their

distinctive feature is, however, the slender and

hypertrophied manual digit III which suggests

climbing ability and arboreal habits (Zhang et

al. 2008) or gathering strategy (as the living

Ayes Ayes which uses its elongated ﬁngers to

pull bugs out of trees; Lhota et al. 2008). This

clade currently includes two or three taxa from

the Middle Jurassic Daohugou beds (Tiaojishan

Formation; Zhou et al. 2013) of Ningcheng, In-

ner Mongolia, China: Epidendrosaurus ninchen-

gensis (= Scansoriopteryx heilmanni; Padian

2004; Figure 15C) known from a partial skel-

eton (Zhang et al. 2002), Epidexipteryx hui, the

most complete scansoriopterygid preserving a

complete skull (Zhang et al. 2008), and possibly

Pedopenna daohugouensis known from a par-

tial right leg covered with pennaceous feathers

(Xu and Zhang 2005). Scansoriopterygids are

currently recovered as basal Oviraptorosauria

(Agnolín & Novas 2013; Brusatte et al. 2014),

basal Paraves (Godefroit et al. 2013a, b), and

basal Avialae (Zhang et al. 2008; Choiniere et al.

2010b; Novas et al. 2012; Senter et al. 2012b).

The clade is also found unresolved by some

workers (e.g., Turner et al. 2012).

Conclusions

Theropod dinosaurs form one of the most suc-

cessful and morphologically diverse groups

of tetrapods, surviving the Cretaceous-Paleo-

gene extinction event and radiating as birds

in the Cenozoic. Even before the K-Pg extinc-

tion non-avian theropods were an extremely

diverse group of archosaurs with complex

interrelationships. The adoption of cladistic

techniques in the 1980s was a major step in

the study of theropod phylogenetics; modern

analyses currently recover around 25 non-

avian theropod subclades, most in a ladder-

like organization. While a consensus of high-

er-level theropod relationships has emerged

and the systematics of non-avian theropods

seems to be relatively well understood, some

signiﬁcant points of contention remain. New-

ly discovered non-avian theropods will hope-

fully shed light on the systematic position of

herrerasaurids within saurischians, megarap-

torans within avetheropods, scansoriopteryg-

ids within pennaraptorans, and troodontids

within paravians. Though one might expect

few major changes in theropod relationships

in the future, large portions of theropod phy-

letic history remain obscure; future discover-

ies of theropods in the Jurassic of Australia or

the Cretaceous of Antarctica, where theropod

faunas are almost unknown, may change the

current view of non-avian theropod systemat-

ics dramatically.

Acknowledgements

We are particularly thankful of Ville Sink-

konen and Gregory Paul for allowing us to use

their skeletal reconstructions of theropods. We

also thank Steve Brusatte (University of Edin-

burg) and Oliver Rauhut (Ludwig-Maximilians-

University) for excellent critiques and helpful

recommendations on the manuscript, which

resulted in substantial improvement. We ac-

knowledge the use of Phylopic for the theropod

silhouettes, and thank Michael Bech Hussein,

Jaime Headden, Michael Keesey, William Park-

er, and Travis Tischler for providing their art-

works on Phylopic. A special thanks goes to the

Wikipaleo group on Facebook, archive.org and

biodiversitylibrary.org, which generously share

a huge number of old books and papers. Many

thanks to Jay Nair and Edio-Ernst Kischlat for

sharing scientiﬁc publications otherwise un-

available. This research was supported by the

Fundação para a Ciência e a Tecnologia (FCT)

scholarship SFRH/BD/62979/2009 (Ministério

da Ciência, Tecnologia e Ensino superior, Por-

tugal). C.H. dedicates this article to Philippe Ta-

quet.

Hendrickx et al., Non-Avian Theropods PalArch’s Journal of Vertebrate Palaeontology, 12, 1 (2015)

PalArch Foundation 35

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Taxon First deﬁnitional

author First phylogenetic deﬁnition Deﬁnition type Deﬁnition Deﬁnitional author

Abelisauridae Bonaparte and

Novas 1985

Novas 1997b Abelisaurus comahuensis, Carnotaurus sastrei,

Xenotarsosaurus bonapartei, Indosaurus

matleyi, Indosuchus raptorius, Majungasaurus

crenatissimus and all descendants of their com-

mon ancestor

Stem-based The most inclusive clade

containing Carnotaurus sas-

trei but not Noasaurus leali

Wilson et al. 2003

Abelisauroidea (Bonaparte and

Novas 1985) Bonaparte 1991a

Holtz 1994 Abelisaurids and those members of the

Ceratosaurus-Abelisauridae clade which shared

a more recent common ancestry than with the

North American genus [Ceratosaurus]

Node-based The least inclusive clade

containing Carnotaurus sas-

trei and Noasaurus leali

Sereno 2005

Afrovenatorinae Carrano et al.

2012

Carrano et al. 2012 All megalosaurids more closely related to

Afrovenator than to Megalosaurus

Stem-based The most inclusive clade

containing Afrovenator

abakensis but not Megalo-

saurus bucklandii

Carrano et al. 2012

Allosauria (Marsh 1878) Paul

1988

/ / Stem-based The most inclusive clade

containing Allosaurus fragi-

lis and Carcharodontosaurus

saharicus but not Sinraptor

dongi

New

Allosauridae Marsh 1878 Padian and Hutchin-

son 1997

Allosaurus and all Allosauroidea closer to it

than to Sinraptor

Stem-based The most inclusive clade

containing Allosaurus

fragilis but not Sinraptor

dongi, Carcharodontosau-

rus saharicus, and Passer

domesticus

Sereno 2005

Allosauroidea (Marsh 1878)

Currie & Zhao 1993a

Padian and Hutchin-

son 1997

Allosaurus and Sinraptor and all descendants

of their most recent common ancestor (Node-

based deﬁnition)

Stem-based The most inclusive clade

containing Allosaurus

fragilis but not Passer

domesticus

Sereno 2005

Hendrickx et al., Non-Avian Theropods PalArch’s Journal of Vertebrate Palaeontology, 12, 1 (2015)

PalArch Foundation 61

Taxon First deﬁnitional

author First phylogenetic deﬁnition Deﬁnition type Deﬁnition Deﬁnitional author

Alvarezsauridae Bonaparte

1991b

Sereno 1998 All ornithomimosaurs closer to Shuvuuia than

to Ornithomimus (Stem-based deﬁnition)

Node-based The least inclusive clade

containing Alvarezsaurus

calvoi and Mononykus

olecranus

Modiﬁed from

Choiniere et al.

2010b

Alvarezsauroidea Bonaparte

1991b

Livezey and Zusi

2007

Clade containing Patagonykus, Alvarezsaurus

and Mononykus

Stem-based The most inclusive clade

containing Alvarezsaurus

calvoi but not Passer domes-

ticus

Modiﬁed from

Choiniere et al.

2010b

Averostra Paul 2002 (sensu

Ezcurra 2006)

Paul 2002 [All] ceratosaurs, megalosaurs, and abelisaurs Node-based The least inclusive clade

containing Ceratosaurus

nasicornis and Passer do-

mesticus

Allain et al. 2012

Avetheropoda Paul 1988 Padian et al. 1999 The most recent common ancestor of Neor-

nithes and Allosaurus and all descendants of

that ancestor.

Node-based The least inclusive clade

containing Allosaurus fragi-

lis and Passer domesticus

Modiﬁed from Holtz

et al. 2004

Avialae Gauthier 1986 Gauthier 1986 Ornithurae plus all extinct maniraptorans

that are closer to Ornithurae than they are to

Deinonychosauria

Stem-based The most-inclusive clade

containing Passer domesti-

cus but not Dromaeosaurus

albertensis or Troodon

formosus

Maryańska et al.

2002

Baryonychinae (Charig & Mil-

ner 1986) Sereno et al. 1998

Sereno et al. 1998 All spinosaurids that are more closely related

to Baryonyx than to Spinosaurus

Stem-based The most inclusive clade

containing Baryonyx walkeri

but not Spinosaurus aegyp-

tiacus

Holtz et al. 2004

Brachyrostra Canale et al. 2009 Canale et al. 2009 All the abelisaurids more closely related to

Carnotaurus sastrei than to Majungasaurus

crenatissimus

Stem-based The most inclusive clade

containing Carnotaurus sas-

trei but not Majungasaurus

crenatissimus

Modiﬁed from Ca-

nale et al. 2009

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Taxon First deﬁnitional

author First phylogenetic deﬁnition Deﬁnition type Deﬁnition Deﬁnitional author

Caenagnathidae Sternberg 1940 Sues 1997 Caenagnathus pergracilis, Chirostenotes

elegans, “Elmisaurus rarus”, Caenagnathasia

martinsoni, and the most recent common an-

cestor of the aforementioned taxa (Node-based

deﬁnition)

Stem-based The most inclusive clade

containing Caenagnathus

collinsi but not Oviraptor

philoceratops

Maryańska et al.

2002

Caenagnathinae (Sternberg

1940) Paul 1988

Longrich et al. 2013 All species closer to Caenagnathus collinsi than

either Caenagnathasia martinsoni or Elmisau-

rus elegans

Stem-based The most inclusive clade

containing Caenagnathus

collinsi but not Elmisaurus

rarus

Modiﬁed from Lon-

grich et al. 2013

Caenagnathoidea (Sternberg

1940) Sereno 1998

Sereno 1998 Oviraptor, Caenagnathus, their most recent

common ancestor and all descendants

Node-based The least inclusive clade

containing Oviraptor philo-

ceratops and Caenagnathus

collinsi

Modiﬁed from

Sereno 2005

Carcharodontosauria (Stromer

1931) Benson et al. 2010

Benson et al. 2010 The most inclusive clade comprising Carchar-

odontosaurus saharicus and Neovenator salerii

but not Allosaurus fragilis or Sinraptor dongi

Stem-based The most inclusive clade

containing Carcharodon-

tosaurus saharicus and

Neovenator salerii but not

Allosaurus fragilis or Sinrap-

tor dongi

Benson et al. 2010

Carcharodontosauridae Stromer

1931

Sereno 1998 All allosauroids closer to Carcharodontosaurus

than to either Allosaurus, Monolophosaurus,

Cryolophosaurus, or Sinraptor

Stem-based The most inclusive clade

containing Carcharodon-

tosaurus saharicus but not

Neovenator dongi, Allosau-

rus fragilis or Sinraptor

dongi

Benson et al. 2010

Carcharodontosaurinae

(Stromer 1931) Carrano et al.

2012

Brusatte & Sereno

2008

The least inclusive clade containing Carcha-

rodontosaurus saharicus and Giganotosaurus

carolinii

Node-based The least inclusive clade

containing Carcharodon-

tosaurus saharicus and

Giganotosaurus carolinii

Brusatte & Sereno

2008

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Taxon First deﬁnitional

author First phylogenetic deﬁnition Deﬁnition type Deﬁnition Deﬁnitional author

Carnotaurinae Sereno 1998 Sereno 1998 All abelisaurids closer to Carnotaurus than to

Abelisaurus

Stem-based The most inclusive clade

containing Carnotaurus

sastrei but not Skorpiovena-

tor bustingorryi

Modiﬁed from

Sereno 1998

Caudipteridae Zhou and Wang

2000

/ / Stem-based The most inclusive clade

containing Caudipteryx zoui

but not Oviraptor philoc-

eratops and Caenagnathus

collinsi

New

Ceratosauria Marsh 1884b Rowe and Gauthier

1990

The group including Ceratosaurus nasicornis,

Dilophosaurus wetherilli, Liliensternus lilienster-

ni, Coelophysis bauri, Syntarsus rhodesiensis,

Syntarsus kayentakatae, Segisaurus halli, Sar-

cosaurus woodi, and all other taxa stemming

from their most recent common ancestor

Stem-based The most inclusive clade

containing Ceratosaurus

nasicornis but not Passer

domesticus

Sereno 2005 sensu

Holtz and Padian

1995

Ceratosauridae Marsh 1884b Rauhut 2004b Clade containing all ceratosaurs that are more

closely related to Ceratosaurus than to abelis-

aurids

Stem-based The most inclusive clade

containing Ceratosaurus na-

sicornis but not Carnotaurus

sastrei and Noasaurus leali

Modiﬁed from

Rauhut 2004b

Coelophysidae (Nopcsa 1928)

Paul 1988

Sereno 1998 Coelophysis, Procompsognathus, their most

recent common ancestor and all descendants

Node-based The least inclusive clade

containing Coelophysis

bauri and Procompsogna-

thus triassicus

Sereno 1998

Coelophysoidea (Nopcsa 1928)

Holtz 1994

Sereno 1998 All ceratosaurs closer to Coelophysis than to

Carnotaurus

Stem-based The most inclusive clade

containing Coelophysis

bauri but not Carnotaurus

sastrei, Ceratosaurus nasi-

cornis, and Passer domes-

ticus

Sereno 2005

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Taxon First deﬁnitional

author First phylogenetic deﬁnition Deﬁnition type Deﬁnition Deﬁnitional author

Coeluridae Marsh 1881 / / Stem-based The most inclusive clade

containing Coelurus fragilis

but not Proceratosaurus

bradleyi, Tyrannosaurus rex,

Allosaurus fragilis, Comp-

sognathus longipes, Ornitho-

mimus edmontonicus and

Deinonychus antirrhopus

New

Coelurosauria Huene 1914c Gauthier 1986 Birds and all other theropods that are closer to

birds than they are to Carnosauria

Stem-based The most inclusive clade

containing Passer domes-

ticus but not Allosaurus

fragilis, Sinraptor dongi

and Carcharodontosaurus

saharicus

Sereno 2005

Compsognathidae Cope 1871 Holtz et al. 2004 Compsognathus longipes and all taxa sharing

a more recent common ancestor with it than

with Passer domesticus

Stem-based The most inclusive clade

containing Compsognathus

longipes but not Passer

domesticus

Holtz et al. 2004

Deinocheiridae Osmólska and

Roniewicz 1970

Lee et al. 2014 Deinocheirus miriﬁcus and all taxa sharing a

more recent common ancestor with it than

with Ornithomimus velox

Stem-based The most inclusive clade

containing Deinocheirus

miriﬁcus but not Ornithomi-

mus velox

Lee et al. 2014

Deinonychosauria Colbert &

Russell 1969

Padian 1997 All maniraptorans closer to Deinonychus than

to birds (Stem-based deﬁnition)

Node-based The least inclusive clade

containing Troodon for-

mosus and Velociraptor

mongoliensis but not Passer

domesticus

Modiﬁed from

Sereno 2005

Hendrickx et al., Non-Avian Theropods PalArch’s Journal of Vertebrate Palaeontology, 12, 1 (2015)

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Taxon First deﬁnitional

author First phylogenetic deﬁnition Deﬁnition type Deﬁnition Deﬁnitional author

Dilophosauridae (Paul 1988)

Charig & Milner 1990

/ / Stem-based The most-inclusive clade

containing Dilophosaurus

wetherilli but not Coelo-

physis bauri, Ceratosaurus

nasicornis and Passer

domesticus

New

Dromaeosauridae (Matthew &

Brown 1922) Colbert & Russell

1969

Sereno 1998 All deinonychosaurs closer to Velociraptor than

to Troodon

Stem-based The most inclusive clade

containing Dromaeosaurus

albertensis but not Troodon

formosus, Ornithomimus

edmontonicus, and Passer

domesticus

Sereno 2005

Dromaeosaurinae Matthew &

Brown 1922

Sereno 1998 All dromaeosaurids closer to Dromaeosaurus

than to Velociraptor

Stem-based The most inclusive clade

containing Dromaeosau-

rus albertensis but not

Velociraptor mongoliensis,

Microraptor zhaoianus,

Unenlagia comahuensis and

Passer domesticus

Sereno 2005

Elmisaurinae (Osmólska 1981)

Currie 2000

/ / Stem-based The most inclusive clade

containing Elmisaurus rarus

but not Caenagnathus col-

linsi

New

Eudromaeosauria Longrich &

Currie 2009b

Turner et al. 2012 The node-based monophyletic group contain-

ing the last common ancestor of Saurornit-

holestes langstoni, Deinonychus antirrhopus,

Dromaeosaurus albertensis, and Velociraptor

mongoliensis, and all its descendants

Node-based The least inclusive clade

containing Saurornitholestes

langstoni, Deinonychus

antirrhopus, Dromaeosaurus

albertensis, and Velociraptor

mongoliensis

Modiﬁed from

Turner et al. 2012

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Taxon First deﬁnitional

author First phylogenetic deﬁnition Deﬁnition type Deﬁnition Deﬁnitional author

Herrerasauridae Benedetto 1973 Novas 1992 Herrerasaurus and Staurikosaurus and their

most recent common ancestor

Stem-based The most inclusive clade

containing Herrerasaurus

ischigualastensis but not

Passer domesticus

Sereno 2005

Jinfengopteryginae Turner et

al. 2012

Turner et al. 2012 A stem-based monophyletic group containing

Jinfengopteryx elegans, and all coelurosaurs

closer to it than to Troodon formosus, Passer

domesticus, and Sinovenator changii

Stem-based The most inclusive clade

containing Jinfengopteryx

elegans but not Troodon for-

mosus, Sinovenator changii

and Passer domesticus

Modiﬁed from

Turner et al. 2012

Majungasaurinae Tortosa et al.

2014

Tortosa et al. 2014 All the abelisaurids more closely related to

Majungasaurus crenatissimus than to Carnotau-

rus sastrei

Stem-based The most inclusive clade

containing Majungasaurus

crenatissimus but not Car-

notaurus sastrei

Modiﬁed from Tor-

tosa et al. 2014

Maniraptora Gauthier 1986 Gauthier 1986 All coelurosaurs that are closer to birds than

they are to Ornithomimidae

Stem-based The most-inclusive clade

containing Passer domesti-

cus but not Ornithomimus

velox

Maryańska et al.

2002

Maniraptoriformes Holtz 1995 Holtz 1996 The most recent common ancestor of Ornitho-

mimus and birds (i.e., The most recent common

ancestor of Arctometatarsalia and Manirap-

tora), and all descendants of that common

ancestor

Node-based The least-inclusive clade

containing Passer domesti-

cus and Ornithomimus velox

Maryańska et al.

2002

Megalosauria (Fitzinger 1843)

Bonaparte 1850

Allain et al. 2012 The most inclusive clade containing Spinosau-

rus aegyptiacus and Torvosaurus tanneri but

not Allosaurus fragilis, and Passer domesticus

(Stem-based deﬁnition)

Node-based The least inclusive clade

containing Megalosaurus

bucklandii and Spinosaurus

aegyptiacus

Modiﬁed from Al-

lain et al. 2012

Megalosauridae (Fitzinger 1843)

Bonaparte 1850

Allain 2002 Poekilopleuron? valesdunensis, Torvosaurus and

Afrovenator, and all descendants of their com-

mon ancestor (Node-based deﬁnition)

Stem-based The most inclusive clade

containing Megalosaurus

bucklandii but not Allosau-

rus fragilis, Spinosaurus

aegyptiacus, and Passer

domesticus

Holtz et al. 2004

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Taxon First deﬁnitional

author First phylogenetic deﬁnition Deﬁnition type Deﬁnition Deﬁnitional author

Megalosaurinae (Fitzinger 1843)

Carrano et al. 2012

Carrano et al. 2012 All megalosaurids closer to Megalosaurus than

to Afrovenator

Stem-based The most inclusive clade

containing Megalosaurus

bucklandii but not Afrovena-

tor abakensis

Carrano et al. 2012

Megalosauroidea (Fitzinger

1843) Walker 1964

Sereno 1998 Spinosaurus, Torvosaurus, their most recent

common ancestor and all descendants (Deﬁni-

tion given to Spinosauroidea)

Stem-based The most inclusive clade

containing Megalosaurus

bucklandii but not Passer

domesticus

Modiﬁed from Holtz

et al. 2004

Megaraptora Benson et al. 2010 Benson et al. 2010 The most inclusive clade comprising Megarap-

tor namunhuaiquii but not Chilantaisaurus

tashuikouensis, Neovenator salerii, Carchar-

odontosaurus saharicus or Allosaurus fragilis

Stem-based The most inclusive clade

containing Megaraptor

namunhuaiquii but not Chi-

lantaisaurus tashuikouensis,

Neovenator salerii, Carcha-

rodontosaurus saharicus or

Allosaurus fragilis

Benson et al. 2010

Megaraptoridae (Benson et al.

2010) Novas et al. 2013

Novas et al. 2013 A stem based clade including all theropods

closer to Megaraptor namunhuaiquii than to

Fukuiraptor kitadaniensis, Chilantaisaurus

tashuikouensis, Neovenator salerii, Carchar-

odontosaurus saharicus, Allosaurus fragilis,

Baryonyx walkeri, Tyrannosaurus rex, and

Passer domesticus

Stem-based The most inclusive clade

containing Megaraptor

namunhuaiquii but not Fu-

kuiraptor kitadaniensis, Chi-

lantaisaurus tashuikouensis,

Neovenator salerii, Carcha-

rodontosaurus saharicus,

Allosaurus fragilis, Baryonyx

walkeri, Tyrannosaurus rex,

and Passer domesticus

Modiﬁed from No-

vas et al. 2013

Metriacanthosauridae Paul 1988 Padian and Hutchin-

son 1997

Sinraptor and all Allosauroidea closer to it than

to Allosaurus (Deﬁnition given to Sinraptori-

dae)

Stem-based The most inclusive clade

containing Metriacantho-

saurus parkeri but not

Allosaurus fragilis, Carcha-

rodontosaurus saharicus, or

Passer domesticus

Modiﬁed from

Sereno 2005

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Taxon First deﬁnitional

author First phylogenetic deﬁnition Deﬁnition type Deﬁnition Deﬁnitional author

Metriacanthosaurinae (Paul

1988) Carrano et al. 2012

Carrano et al. 2012 All metriacanthosaurids more closely related to

Metriacanthosaurus than to Yangchuanosaurus

Stem-based The most inclusive clade

containing Metriacantho-

saurus parkeri but not Yang-

chuanosaurus shangyouensis

Modiﬁed from Car-

rano et al. 2012

Microraptorinae Senter et al.

2004

Sereno 2005 The most inclusive clade containing Microrap-

tor zhaoianus but not Dromaeosaurus alber-

tensis, Velociraptor mongoliensis, Unenlagia

comahuensis, Passer domesticus

Stem-based The most inclusive clade

containing Microraptor

zhaoianus but not Drom-

aeosaurus albertensis,

Velociraptor mongoliensis,

Unenlagia comahuensis, and

Passer domesticus

Sereno 2005

Mononykinae Chiappe et al.

1998

Chiappe et al. 1998 The common ancestor of Mononykus, Shuvuuia,

and Parvicursor, plus all their descendants

Node-based The least inclusive clade

containing Mononykus olec-

ranus and Shuvuuia deserti

Sereno 2005

Neoceratosauria Novas 1991 Holtz 1994 The most recent common ancestor of Cera-

tosaurus and Abelisauridae and all of its

descendants

Node-based The least inclusive clade

containing Ceratosaurus

nasicornis and Carnotaurus

sastrei

Modiﬁed from Holtz

1994

Neotheropoda Bakker 1986 Sereno 1998 Coelophysis, Neornithes, their most recent com-

mon ancestor and all descendants

Node-based The least inclusive clade

containing Coelophysis

bauri and Passer domesticus

Sereno 2005

Neovenatoridae Benson et al.

2010

Benson et al. 2010 The most inclusive clade comprising Neovena-

tor salerii but not Carcharodontosaurus sahari-

cus, Allosaurus fragilis or Sinraptor dongi

Stem-based The most inclusive clade

containing Neovenator sale-

rii but not Carcharodonto-

saurus saharicus, Allosaurus

fragilis or Sinraptor dongi

Benson et al. 2010

Noasauridae Bonaparte & Pow-

ell 1980

Wilson et al. 2003 The most inclusive clade containing Noasaurus

leali but not Carnotaurus sastrei

Stem-based The most inclusive clade

containing Noasaurus leali

but not Carnotaurus sastrei

Wilson et al. 2003

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Taxon First deﬁnitional

author First phylogenetic deﬁnition Deﬁnition type Deﬁnition Deﬁnitional author

Orionides Carrano et al. 2012 Carrano et al. 2012 Megalosauroidea, Avetheropoda, their most re-

cent common ancestor, and all its descendants

Node-based The least-inclusive clade

containing Megalosaurus

bucklandii, Allosaurus fragi-

lis and Passer domesticus

Modiﬁed from Car-

rano et al. 2012

Ornithomimidae Marsh 1890 Sereno 1998 All ornithomimosaurs closer to Ornithomimus

than to Erlikosaurus

Stem-based The most inclusive clade

containing Ornithomimus

velox but not Deinocheirus

miriﬁcus

Lee et al. 2014

Ornithomimosauria (Marsh

1890) Barsbold 1976a

Osmólska 1997 All bullatosaurs closer to Ornithomimus than to

Troodon

Stem-based The most inclusive clade

containing Ornithomimus

velox but not Allosaurus

fragilis, Tyrannosaurus rex,

Compsognathus longipes,

Alvarezsaurus calvoi, Ther-

izinosaurus cheloniformis,

Deinonychus antirrhopus,

Troodon formosus, and

Passer domesticus

Lee et al. 2014

Oviraptoridae Barsbold 1976b Sereno 1998 All oviraptorosaurs closer to Oviraptor than to

Caenagnathus

Stem-based The most inclusive clade

containing Oviraptor philo-

ceratops but not Caenagna-

thus collinsi

Maryańska et al.

2002

Oviraptorinae (Barsbold 1976b)

(Barsbold 1981)

Osmólska et al. 2004 Oviraptor philoceratops, Citipati osmolskae,

their most recent common ancestor, and all

descendants.

Node-based The least inclusive clade

containing Oviraptor

philoceratops and Citipati

osmolskae

Osmólska et al. 2004

Oviraptorosauria Barsbold

1976a

Barsbold 1997 Oviraptoridae and all taxa closer to Oviraptor

than to birds

Stem-based The most-inclusive clade

containing Oviraptor

philoceratops but not Passer

domesticus

Maryańska et al.

2002

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Taxon First deﬁnitional

author First phylogenetic deﬁnition Deﬁnition type Deﬁnition Deﬁnitional author

Paraves Sereno 1997 Sereno 1998 All maniraptorans closer to Neornithes than to

Oviraptor

Stem-based The most inclusive clade

containing Passer domesti-

cus but not Oviraptor philoc-

eratops

Holtz & Osmólska

2004

Parvicursorinae Karhu and

Rautian 1996

Choiniere et al.

2010b

The least inclusive clade containing Parvicur-

sor, Mononykus and their most recent common

ancestor (Node-based deﬁnition)

Stem-based The most inclusive clade

containing Parvicursor re-

motus but not Patagonykus

puertai

Xu et al. 2011b

Pennaraptora Foth et al. 2014 Foth et al. 2014 Clade including Oviraptor philoceratops, Dei-

nonychus antirrhopus and Passer domesticus

and all descendants of their most recent com-

mon ancestor

Node-based The least inclusive clade

containing Oviraptor

philoceratops, Deinonychus

antirrhopus and Passer

domesticus

Foth et al. 2014

Piatnitzkysauridae Carrano et

al. 2012

Carrano et al. 2012 All megalosauroids more closely related to

Piatnitzkysaurus than to either Spinosaurus or

Megalosaurus

Stem-based The most inclusive clade

containing Piatnitzkysaurus

ﬂoresi but not Spinosaurus

aegyptiacus and Megalosau-

rus bucklandii

Carrano et al. 2012

Proceratosauridae Rauhut et al.

2010

Rauhut et al. 2010 All theropods that are more closely related to

Proceratosaurus than to Tyrannosaurus, Al-

losaurus, Compsognathus, Coelurus, Ornithomi-

mus, or Deinonychus

Stem-based The most inclusive clade

containing Proceratosaurus

bradleyi but not Tyranno-

saurus rex, Allosaurus fragi-

lis, Compsognathus longipes,

Coelurus fragilis, Ornitho-

mimus edmontonicus and

Deinonychus antirrhopus

Rauhut et al. 2010

Scansoriopterygidae Czerkas

and Yuan 2002

Zhang et al. 2008 The least-inclusive clade containing Epidendro-

saurus ningchengensis and Epidexipteryx hui

Node-based The least-inclusive clade

containing Epidendrosaurus

ningchengensis and Epidexi-

pteryx hui

Zhang et al. 2008

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Taxon First deﬁnitional

author First phylogenetic deﬁnition Deﬁnition type Deﬁnition Deﬁnitional author

Spinosauridae Stromer 1915 Sereno 1998 All spinosauroids closer to Spinosaurus than to

Torvosaurus

Stem-based The most inclusive clade

containing Spinosaurus

aegyptiacus but not Torvo-

saurus tanneri, Allosaurus

fragilis, and Passer domes-

ticus

Sereno 2005

Spinosaurinae (Stromer 1915)

Sereno et al. 1998

Holtz et al. 2004 Spinosaurus aegyptiacus and all taxa sharing

a more recent common ancestor with it than

with Baryonyx walkeri

Stem-based The most inclusive clade

containing Spinosaurus ae-

gyptiacus but not Baryonyx

walkeri

Holtz et al. 2004

Tetanurae Gauthier 1986 Gauthier 1986 Birds and all other theropods closer to birds

than they are to Ceratosauria

Stem-based The most inclusive clade

containing Passer domes-

ticus but not Ceratosaurus

nasicornis

Allain et al. 2012

Therizinosauria (Maleev 1954)

Russell 1997

Russell 1997 Alxasaurus, Enigmosaurus, Erlikosaurus, Nan-

shiungosaurus, Segnosaurus, Therizinosaurus

and all others closer to them than to ovirapto-

rosaurs, ornithomimids, and troodontids

Stem-based The most inclusive clade

containing Therizinosaurus

cheloniformis but not Tyran-

nosaurus rex, Ornithomimus

edmontonicus, Mononykus

olecranus, Oviraptor philoc-

eratops or Troodon formosus

Zanno 2010b

Therizinosauridae Maleev 1954 Sereno 1998 All ornithomimosaurs closer to Erlikosaurus

than to Ornithomimus (Stem-based deﬁnition)

Node-based The least inclusive clade

containing Nothronychus

graffami, Segnosaurus

galbinensis, Erlikosaurus an-

drewsi and Therizinosaurus

cheloniformis

Modiﬁed from

Zanno et al. 2009

Therizinosauroidea (Maleev

1954) Russell & Dong 1993

Zhang et al. 2001 All coelurosaurs closer to Therizinosaurus than

to either Ornithomimus, Oviraptor, Velociraptor,

or Neornithes (Stem-based deﬁnition)

Node-based The least inclusive clade

containing Beipiaosaurus

inexpectus and Therizino-

saurus cheloniformis

Clark et al. 2004

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Taxon First deﬁnitional

author First phylogenetic deﬁnition Deﬁnition type Deﬁnition Deﬁnitional author

Theropoda Marsh 1881 Gauthier 1986 Birds and all saurischians that are closer to

birds than they are to sauropodomorphs

Stem-based The most inclusive clade

containing Passer domes-

ticus but not Saltasaurus

loricatus

Sereno 2005

Troodontidae Gilmore 1924 Varricchio 1997 Troodon, Sinornithoides, Saurornithoides,

Borogovia, and all coelurosaurs closer to them

than to ornithomimids, oviraptorosaurs, or

other well-deﬁned taxa

Stem-based The most inclusive clade

containing Troodon formo-

sus but not Velociraptor

mongoliensis, Ornithomimus

edmontonicus, and Passer

domesticus

Sereno 2005

Tyrannosauridae Osborn 1906 Holtz 2001 All descendants of the most recent common

ancestor of Tyrannosaurus and Aublysodon

Node-based The least inclusive clade

containing Tyrannosaurus

rex, Gorgosaurus libratus

and Albertosaurus sarcopha-

gus

Sereno 2005

Tyrannosaurinae (Osborn 1906)

Matthew & Brown 1922

Sereno 1998 All tyrannosaurids closer to Tyrannosaurus

than to either Albertosaurus, Daspletosaurus, or

Gorgosaurus

Stem-based The most inclusive clade

containing Tyrannosaurus

rex but not Gorgosaurus

libratus and Albertosaurus

sarcophagus

Sereno 2005

Tyrannosauroidea (Osborn

1906) Walker 1964

Sereno 1998 All maniraptorans closer to Tyrannosaurus

than to Neornithes

Stem-based The most inclusive clade

containing Tyrannosaurus

rex but not Ornithomimus

edmontonicus, Troodon

formosus, or Velociraptor

mongoliensis

Sereno 2005

Hendrickx et al., Non-Avian Theropods PalArch’s Journal of Vertebrate Palaeontology, 12, 1 (2015)

PalArch Foundation 73

Taxon First deﬁnitional

author First phylogenetic deﬁnition Deﬁnition type Deﬁnition Deﬁnitional author

Unenlagiinae Bonaparte 1999 Makovicky et al.

2005

All taxa closer to Unenlagia comahuensis than

to Velociraptor mongoliensis

Stem-based The most inclusive clade

containing Unenlagia coma-

huensis but not Velociraptor

mongoliensis, Dromaeosau-

rus albertensis, Microrap-

tor zhaoianus and Passer

domesticus

Sereno 2005

Preliminary investigation of Allosauroidea facial integument and the evolution of theropod facial armor

Article

Jan 2022

In amniotes, the growth of specialized facial integument, or external skin tissues, has a close relationship with the texture and morphology of underlying cranial bones. Osteological correlates of facial skin structures have been used before to reconstruct the integument of several extinct dinosaurian lineages but not for theropods from the clade Allosauroidea, whose members exhibit exceptionally rugose and heavily ornamented skulls. This study aims to investigate, in a preliminary sense, the facial integument of Allosauroidea by examining osteological correlates from high-resolution photographs of fossil material. We compared the inferred integument of allosauroids to analogous structures in modern animals in order to discuss potential biological and behavioral implications, with soft-tissue adaptations suggesting headbutting or sparring in some taxa. Allosauroids display an evolutionary trend for increasing cranial cornification throughout their history, and their most derived members exhibit greater development of facial armor than other tetanurans. We developed three hypotheses to explain the selective pressures that may have driven the independent evolution of this dermal armor in several unrelated theropod clades, namely abelisaurids, carcharodontosaurids, and tyrannosaurids. Empirical tests of paleoecological data did not support any of the three hypotheses of intraspecific conflict, competition from other carnivores, or dangerous prey items as the main pressure that drove the evolution of theropod facial armor. We suggest that more sampling of armor-faced theropods and their associated ecosystems, especially those from the Southern Hemisphere, is needed to reveal the reasons behind the convergent evolution of this integument.

Macroevolutionary patterns in the pelvis, stylopodium and zeugopodium of megalosauroid theropod dinosaurs and their importance for locomotor function

Article

Full-text available

Aug 2023

During the Mesozoic, non-avian theropods represented one of the most successful clades globally distributed, with a wide diversity of forms. An example is the clade Megalosauroidea, which included medium- to large-bodied forms. Here, we analyse the macroevolution of the locomotor system in early Theropoda, emphasizing the Megalosauroidea. We scored the Spinosaurus neotype in a published taxon-character matrix and described the associated modifications in character states, mapping them onto a phylogeny and using these to study disparity. In the evolution of Megalosauroidea, there was the mosaic emergence of a low swollen ridge; enlargement of the posterior brevis fossa and emergence of a posterodorsal process on the ilium in some megalosauroids; emergence of a femoral head oriented anteromedially and medially angled, and appearance of posterolaterally oriented medial femoral condyles in spinosaurids. The greatest morphological disparity is in the ilium of megalosaurids; the ischium seems to have a high degree of homoplasy; there is a clear distinction in the femoral morphospace regarding megalosauroids and other theropods; piatnitzkysaurids show considerable disparity of zeugopodial characters. These reconstructions of osteological evolution form a stronger basis on which other studies could build, such as mapping of pelvic/appendicular musculature and/or correlating skeletal traits with changes in locomotor function.

Morphological disparity and structural performance of the dromaeosaurid skull informs ecology and evolutionary history

Article

Full-text available

Apr 2024

Non-avialan theropod dinosaurs had diverse ecologies and varied skull morphologies. Previous studies of theropod cranial morphology mostly focused on higher-level taxa or characteristics associated with herbivory. To better understand morphological disparity and function within carnivorous theropod families, here we focus on the Dromaeosauridae, ‘raptors’ traditionally seen as agile carnivorous hunters. We applied 2D geometric morphometrics to quantify skull shape, performed mechanical advantage analysis to assess the efficiency of bite force transfer, and performed finite element analysis to examine strain distribution in the skull during biting. We find that dromaeosaurid skull morphology was less disparate than most non-avialan theropod groups. Their skulls show a continuum of form between those that are tall and short and those that are flat and long. We hypothesise that this narrower morphological disparity indicates developmental constraint on skull shape, as observed in some mammalian families. Mechanical advantage indicates that Dromaeosaurus albertensis and Deinonychus antirrhopus were adapted for relatively high bite forces, while Halszkaraptor escuilliei was adapted for high bite speed, and other dromaeosaurids for intermediate bite forces and speeds. Finite element analysis indicates regions of high strain are consistent within dromaeosaurid families but differ between them. Average strain levels do not follow any phylogenetic pattern, possibly due to ecological convergence between distantly-related taxa. Combining our new morphofunctional data with a re-evaluation of previous evidence, we find piscivorous reconstructions of Halszkaraptor escuilliei to be unlikely, and instead suggest an invertivorous diet and possible adaptations for feeding in murky water or other low-visibility conditions. We support Deinonychus antirrhopus as being adapted for taking large vertebrate prey, but we find that its skull is relatively less resistant to bite forces than other dromaeosaurids. Given the recovery of high bite force resistance for Velociraptor mongoliensis, which is believed to have regularly engaged in scavenging behaviour, we suggest that higher bite force resistance in a dromaeosaurid taxon may reflect a greater reliance on scavenging rather than fresh kills. Comparisons to the troodontid Gobivenator mongoliensis suggest that a gracile rostrum like that of Velociraptor mongoliensis is ancestral to their closest common ancestor (Deinonychosauria) and the robust rostra of Dromaeosaurus albertensis and Deinonychus antirrhopus are a derived condition. Gobivenator mongoliensis also displays a higher jaw mechanical advantage and lower resistance to bite force than the examined dromaeosaurids, but given the hypothesised ecological divergence of troodontids from dromaeosaurids it is unclear which group, if either, represents the ancestral condition. Future work extending sampling to troodontids would therefore be invaluable and provide much needed context to the origin of skull form and function in early birds. This study illustrates how skull shape and functional metrics can discern non-avialan theropod ecology at lower taxonomic levels and identify variants of carnivorous feeding. Supplementary Information The online version contains supplementary material available at 10.1186/s12862-024-02222-5.

A combined approach to identify isolated theropod teeth from the Cenomanian Kem Kem Group of Morocco: cladistic, discriminant, and machine learning analyses

Article

Mar 2024
J Vertebr Paleontol

The Kem Kem Group of Southeastern Morocco, North Africa, is well known for theropod remains, especially isolated teeth. Here, a collection of isolated theropod teeth is assessed for diversity using a combination of linear discriminant, phylogenetic, and machine learning analyses for the first time. The results confirm earlier studies on Kem Kem theropod diversity, with teeth referred to Abelisauridae, Spinosaurinae, and Carcharodontosauridae. A single tooth is ascribed to a non-abelisauroid ceratosaur or a megaraptoran and may represent the enigmatic averostran Deltadromeus. Spinosaurine teeth are clearly differentiated by all three methodologies, whereas abelisaurid and carcharodontosaurid teeth could only be distinguished by the machine learning and phylogenetic analyses. This study shows that a combination of independent methods is most effective at providing strong evidence on theropod dental diversity in a particular assemblage, and that cladistic and machine learning analyses are the most reliable approaches to identify isolated dinosaur teeth. The methodology used here is likely to yield results in other dinosaur assemblages where isolated teeth are more abundant than body fossils.

Desert Cretaceous Dinosaurs: The Botucatu Paleodesert and the Footprints Across the Dunes

Chapter

May 2024

Tetrapod fossil footprints are common in the aeolian sandstones from the Botucatu Formation in many Brazilian states, all around the edge of the Paraná Basin. This unit is represented by well-selected, reddish aeolian sandstones, related to an extensive dune field. Its age is Early Cretaceous (Berriasian-Barremian). The main ichnosite from the Botucatu Formation occurs in the interior of São Paulo State, in the Araraquara region, at the São Bento quarry, where slabs for covering and paving of public sidewalks, were extracted. This quarry presents a section of a large dune, about 20 m high and 100 m long, exhibiting foresets with a dip of 29° in the S-SW direction. Alongside the bedding surfaces there are abundant tracks attributed to mammals, Theropoda and Ornithopoda. These are herein grouped in four morphotypes, in addition to other kinds of isolated footprints difficult to classify. All these occurrences belong to bipedal and functionally tridactyl dinosaurs; no Sauropodomorpha tracks were found. To these theropod and ornithopod footprints and trackways from the eastern side of the Paraná Basin are added those of the western side, in the area of Nioaque, Mato Grosso do Sul State. The Botucatu paleodesert harbored a possibly endemic fauna, of bipedal carnivorous and herbivorous dinosaurs, which makes difficult a faunistic comparison with other localities, in other paleobiogeographic contexts.

Comparative osteology of non-avian dinosaurs and Aves, and the refinement of feathered flight

Preprint

Full-text available

Jun 2024

Ethan O'Callaghan-Linford

Aves (Dinosauria: Theropoda), the first and only clade of maniraptorans to have evolved powered flight, are the most speciose amniotes with over 18,000 known extant species that dominate the skies over all seven continents. They first evolved in the Middle Jurassic around 174-161.5 Ma, and have since represented nature’s greatest anecdote for success. Despite specious criticism, the connected phylogeny between non-avian dinosaurs and avian dinosaurs has been well-documented since 1868 with Thomas Henry Huxley’s description of the “urvogel” Archaeopteryx lithographica holotype from the Solnhofen Limestone, Germany, and is one of the most recognisable fossils in the world. The genus displays skeletal characteristics of both dinosaurian and neornithine nature—as well as the exceptional preservation of soft tissue plumage. Since then, workers have produced a plethora of theses that continue to elaborate on the phylogeny connecting dinosaurs and birds, debates over the origin of flapping-flight and gliding locomotion, and the origin of Neornithes. Particularly, osteological comparative anatomy. With this comes the conclusive assertion that the osteology of Aves is not only outstandingly similar to that of Maniraptora, but is of a direct causal result through modification. With emphasis placed on fossilised remains rather than the procedures of software-driven phylogenetic analysis, the present review will briefly detail the evidence (organised by apomorphy) of dinosaur-avian relationships, focusing on comparative osteology and feather evolution.

Reconstruction of the pelvic girdle and hindlimb musculature of the early tetanurans Piatnitzkysauridae (Theropoda, Megalosauroidea)

Article

Full-text available

Dec 2023
J ANAT

Piatnitzkysauridae were Jurassic theropods that represented the earliest diverging branch of Megalosauroidea, being one of the earliest lineages to have evolved moderate body size. This clade's typical body size and some unusual anatomical features raise questions about locomotor function and specializations to aid in body support; and other palaeobiological issues. Biomechanical models and simulations can illuminate how extinct animals may have moved, but require anatomical data as inputs. With a phylogenetic context, osteological evidence, and neontological data on anatomy, it is possible to infer the musculature of extinct taxa. Here, we reconstructed the hindlimb musculature of Piatnitzkysauridae ( Condorraptor , Marshosaurus , and Piatnitzkysaurus ). We chose this clade for future usage in biomechanics, for comparisons with myological reconstructions of other theropods, and for the resulting evolutionary implications of our reconstructions; differential preservation affects these inferences, so we discuss these issues as well. We considered 32 muscles in total: for Piatnitzkysaurus , the attachments of 29 muscles could be inferred based on the osteological correlates; meanwhile, in Condorraptor and Marshosaurus , we respectively inferred 21 and 12 muscles. We found great anatomical similarity within Piatnitzkysauridae, but differences such as the origin of M . ambiens and size of M . caudofemoralis brevis are ev. Similarities were evident with Aves, such as the division of the M . iliofemoralis externus and M . iliotrochantericus caudalis and a broad depression for the M . gastrocnemius pars medialis origin on the cnemial crest. Nevertheless, we infer plesiomorphic features such as the origins of M . puboischiofemoralis internus 1 around the “cuppedicus” fossa and M . ischiotrochantericus medially on the ischium. As the first attempt to reconstruct muscles in early tetanurans, our study allows a more complete understanding of myological evolution in theropod pelvic appendages.

Turtle skull development unveils a molecular basis for amniote cranial diversity

Article

Full-text available

Nov 2023

Amniote skulls display diverse architectural patterns including remarkable variations in the number of temporal arches surrounding the upper and lower temporal fenestrae. However, the cellular and molecular basis underlying this diversification remains elusive. Turtles are a useful model to understand skull diversity due to the presence of secondarily closed temporal fenestrae and different extents of temporal emarginations (marginal reduction of dermal bones). Here, we analyzed embryos of three turtle species with varying degrees of temporal emargination and identified shared widespread coexpression of upstream osteogenic genes Msx2 and Runx2 and species-specific expression of more downstream osteogenic genes Sp7 and Sparc in the head. Further analysis of representative amniote embryos revealed differential expression patterns of osteogenic genes in the temporal region, suggesting that the spatiotemporal regulation of Msx2 , Runx2 , and Sp7 distinguishes the temporal skull morphology among amniotes. Moreover, the presence of Msx2- and/or Runx2 -positive temporal mesenchyme with osteogenic potential may have contributed to their extremely diverse cranial morphology in reptiles.

New caenagnathid (Theropoda: Oviraptorosauria) dinosaur specimens from middle and upper Campanian strata of West Texas

Article

Nov 2023

Late Triassic small and medium sized vertebrates of Jameson Land Basin, East Greenland

Thesis

Aug 2021

Valerian J. P. Jésus

Several paleontological field expeditions of Danish-US team took place in the Jameson Land Basin, in East Greenland. Their objective was to recover vertebrate fossils for understanding the evolution of the vertebrate fauna of the region and its dynamics with the surrounding through geological times. In the Triassic, the basin was located at 41° N in the northern rim of Pangea and bordered in the North by the Boreal Sea. During Late Triassic, the region was an ephemeral to perennial lake systems in dry steppe-like climate, crossed by rivers. This work focus on the microvertebrate remains recovered by expeditions from 1991 to 2016. The specimens have been photographed and listed in a catalogue. 867 specimens out of 993 could be identified. The main groups of Vertebrate are all present (Chondrichthyes, Actinopterygii, Amphibia, Reptilia, Mammalia) with new taxa never described nor reported in Greenland, such as Lissodus, Parvodus, Rhomphaiodon, Gyrolepis, Nothosauridae and the first batrachian ilium found in the region. The revision of two sphenodontians jaws fragment as Clevosauridae, with in association of the previously mentioned taxa that are highly documented in Europe and Asia, confirms a relationship between faunas of Greenland and Eurasia during the Late Triassic.

The osteology of Masiakasaurus knopfleri, a small abelisauroid (Dinosauria: Theropoda) from the Late Cretaceous of Madagascar

Article

Full-text available

Sep 2002

We describe the osteology of the new small theropod dinosaur Masiakasaurus knopfleri, from the Late Cretaceous Maevarano Formation of northwestern Madagascar. Approximately 40% of the skeleton is known, including parts of the jaws, axial column, forelimb, pelvic girdle, and hind limb. The jaws of Masiakasaurus are remarkably derived, bearing a heterodont, procumbent dentition that is unknown elsewhere among dinosaurs. The vertebrae are similar to those of abelisauroids in the reduction of the neural spine, lack of pleurocoelous fossae on the centrum, and extensively pneumatized neural arch. The limb skeleton is relatively gracile and bears numerous abelisauroid synapomorphies, including a rounded humeral head, peg-and-socket iliac-pubic articulation, prominent femoral medial epicondyle, expanded tibial cnemial crest, and double-grooved pedal unguals. The femora and tibiae show evidence of dimorphism. More specific features shared between Masiakasaurus, the Argentine Noasaurus, and the Indian Laevisuchus suggest that these taxa form a clade (Noasauridae) within Abelisauroidea. This is supported by a cladistic phylogenetic analysis of 158 characters and 23 theropod taxa. Additionally, Ceratosauria is rendered paraphyletic in favor of a sister-taxon relationship between Neoceratosauria and Tetanurae that is exclusive of Coelophysoidea. The unique dental and jaw specializations of Masiakasaurus suggest deviation from the typical theropod diet. Finally, the distribution of noasaurids further supports a shared biogeographic history between South America, Madagascar, and India into the Late Cretaceous.

The Origin of Birds and the Evolution of Flight

Article

Jul 2017

One of the most salient advances in vertebrate paleontology in recent decades has been the settling of the question of the origin of birds, a problem that has vexed evolutionary biologists since well before Darwin. To be sure, the consensus is not unanimous, and many details of this branch of the phylogenetic tree are yet to be worked out, but we now have a much clearer picture of this problem than we had a decade ago. Less settled, but equally stimulating, has been the controversy over the origin of flight in birds and other flying vertebrates. Was there a gliding stage? Did flight begin from the ground up or from the trees down? Were birds initially arboreal? What selective pressures drove the ancestors of birds to take advantage of the aerial opportunity?

The Fossils of the South Downs: Or, Illustrations of the Geology of Sussex

Book

Mar 2012

Gideon Algernon Mantell

Gideon Mantell (1790–1852) was an English physician and geologist best known for pioneering the scientific study of dinosaurs. After an apprenticeship to a local surgeon in Sussex, Mantell became a member of the Royal College of Surgeons in 1811. He developed an interest in fossils, and in 1822 his discovery of fossil teeth which he later identified as belonging to an iguana-like creature he named Iguanadon spurred research into ancient fossils. This volume, first published in 1822, contains Mantell's early research into fossil remains of southern England. Arranged according to rock type and strata, Mantell provides descriptions and analysis of fossil tropical plants, fish, molluscs and what he described as 'an animal of the lizard tribe, of enormous magnitude'. This volume was the first published work to describe a collection of dinosaur remains and provides a fascinating view of early palaeontological research before the formation of the discipline.

The systematic position of Baryonyx walkeri , in the light of Gauthier's reclassification of the Theropoda

Chapter

Nov 1990

In recent years dinosaurs have captured the attention of the public at an unprecedented scale. At the heart of this resurgence in popular interest is an increased level of research activity, much of which is innovative in the field of palaeontology. For instance, whereas earlier palaeontological studies emphasized basic morphologic description and taxonomic classification, modern studies attempt to examine the role and nature of dinosaurs as living animals. More than ever before, we understand how these extinct species functioned, behaved, interacted with each other and the environment, and evolved. Nevertheless, these studies rely on certain basic building blocks of knowledge, including facts about dinosaur anatomy and taxonomic relationships. One of the purposes of this volume is to unravel some of the problems surrounding dinosaur systematics and to increase our understanding of dinosaurs as a biological species. Dinosaur Systematics presents a current overview of dinosaur systematics using various examples to explore what is a species in a dinosaur, what separates genders in dinosaurs, what morphological changes occur with maturation of a species, and what morphological variations occur within a species.

The osteology of Shaochilong maortuensis, a carcharodontosaurid (Dinosauria: Theropoda) from the Late Cretaceous of Asia

Article

Jan 2010

Large-bodied theropod dinosaurs from the Early-mid Cretaceous of the northern continents (Laurasia) are poorly known. One of the most complete and intriguing theropods from this interval is Shaochilong maortuensis Hu, 1964 from the Turonian (< 92 Ma) Ulansuhai Formation of Inner Mongolia, China. The phylogenetic placement of Shaochilong has long been a subject of debate, as it has been referred to several disparate theropod groups (e.g., Megalosauridae, Allosauridae, Tyrannosauroidea, Maniraptora). In a recent taxonomic reassessment, Shaochilong was identified as the first Asian member of Carcharodontosauridae, a clade of allosauroid theropods that was once thought to be restricted to Gondwana and includes some of the largest terrestrial predators to ever live. However, the characters supporting such a placement were only briefly discussed, and a full anatomical description of Shaochilong has yet to be presented. We provide a detailed osteological description of the lectotype and paralectotype series, show that Shaochilong is a small-bodied and short-snouted carcharodontosaurid, and highlight numerous cranial features shared with other carcharodontosaurids. We argue that the vicariant hypothesis of allosauroid biogeography, in which lineages split in concert with the fragmentation of Pangaea, is poorly supported. Finally, large-scale patterns of theropod evolution and faunal replacement are discussed, and it is argued that allosauroids persisted as large-bodied predators later in the Cretaceous than previously thought.

The origin and early evolution of birds

Article

Feb 1998

Birds evolved from and are phylogenetically recognized as members of the theropod dinosaurs; their first known member is the Late Jurassic Archaeopteryx, now represented by seven skeletons and a feather, and their closest known non-avian relatives are the dromaeosaurid theropods such as Deinonychus. Bird flight is widely thought to have evolved from the trees down, but Archaeopteryx and its outgroups show no obvious arboreal or tree-climbing characters, and its wing planform and wing loading do not resemble those of gliders. The ancestors of birds were bipedal, terrestrial, agile, cursorial and carnivorous or omnivorous. Apart from a perching foot and some skeletal fusions, a great many characters that are usually considered ‘avian’ (e.g. the furcula, the elongated forearm, the laterally flexing wrist and apparently feathers) evolved in non-avian theropods for reasons unrelated to birds or to flight. Soon after Archaeopteryx, avian features such as the pygostyle, fusion of the carpometacarpus, and elongated curved pedal claws with a reversed, fully descended and opposable hallux, indicate improved flying ability and arboreal habits. In the further evolution of birds, characters related to the flight apparatus phylogenetically preceded those related to the rest of the skeleton and skull. Mesozoic birds are more diverse and numerous than thought previously and the most diverse known group of Cretaceous birds, the Enantiornithes, was not even recognized until 1981. The vast majority of Mesozoic bird groups have no Tertiary records: Enantiornithes, Hesperornithiformes, Ichthyornithiformes and several other lineages disappeared by the end of the Cretaceous. By that time, a few Linnean ‘Orders’ of extant birds had appeared, but none of these taxa belongs to extant ‘families’, and it is not until the Paleocene or (in most cases) the Eocene that the majority of extant bird ‘Orders’ are known in the fossil record. There is no evidence for a major or mass extinction of birds at the end of the Cretaceous, nor for a sudden ‘bottleneck’ in diversity that fostered the early Tertiary origination of living bird ‘Orders’.

The coelophysoid Lophostropheus airelensis, gen. nov.: a review of the systematics of “Liliensternus” airelensis from the Triassic–Jurassic outcrops of Normandy (France)

Article

Mar 2007

In the early 1990s a theropod dinosaur found close to the Triassic-Jurassic boundary of France was assigned to a second species of the genus Liliensternus: L. airelensis (Moon Airel Formation). This contribution reveals that common features that purportedly unite “L.” airelensis with L. liliensterni are more widely distributed among coelophysoids and basal dinosaurs than it was thought. A cladistic analysis reveals that “L.” airelensis is more closely related to the Coelophysidae than to L. liliensterni. A feature that supports this systematic arrangement includes a supraacetabular crest forming a well-developed ridge continuous with the lateral margin of the brevis fossa, with nondistinct notch between both structures. The new genus Lophostropheus, gen. nov., is therefore erected to include the species L. airelensis. Thus, the new combination Lophostropheus airelensis is proposed.

New information on Stokesosaurus, a tyrannosauroid (Dinosauria: Theropoda) from North America and the United Kingdom

Article

Sep 2008

Roger B. J. Benson

A partial postcranial skeleton from the Late Jurassic (Tithonian) of Dorset, England represents a new species of the theropod dinosaur Stokesosaurus, Stokesosaurus langhami. S. langhami is a member of Tyrannosauroidea, showing a distinct median vertical ridge on the lateral surface of the ilium, a prominent shelf medial to the preacetabular notch, a pronounced ischial tubercle, and a tibia that is elongate relative to the femur. One of only two definitive Jurassic tyrannosauroids known from more than isolated elements, it is the largest Jurassic tyrannosauroid reported to date and provides additional evidence for the presence of relatively small-or medium-sized basal tyrannosauroids in Asia, North America, and Europe during the Late Jurassic. The occurrence of Stokesosaurus in the Tithonian of the UK and USA and the absence of tyrannosauroids in contemporaneous west African faunas supports the hypothesis of a paleobiogeographic link during the Late Jurassic between North America and Europe, to the exclusion of Africa.

A new species of Carcharodontosaurus (Dinosauria: Theropoda) from the Cenomanian of Niger and a revision of the genus

Article

Dec 2007

The systematics and taxonomy of the large allosauroid Carcharodontosaurus are reviewed and a new species of this theropod is described from the Late Cretaceous (Cenomanian) Echkar Formation of Niger, which is roughly coeval with Cenomanian beds elsewhere in northern Africa. The type species, C. saharicus, was based originally on a pair of isolated teeth from Algeria, to which was referred a partial skull and skeleton from Egypt. All of this material was either lost or destroyed more than 50 years ago. Thus, a neotype is designated for C. saharicus, a well-preserved cranium from the Kem Kem beds in Morocco. Newly discovered material from Niger is distinct from that of C. saharicus, justifying the erection of a distinct southern species. C. iguidensis, n. sp. is represented by cranial (maxilla, braincase, dentary, lacrimal, teeth) and postcranial (vertebrae) remains and grew to the same large body size as C. saharicus. The skull displays several autapomorphies, including a laterally reduced external antorbital fossa, a maxillary anteromedial process that is broadly arched toward the midline, a prominent horizontal crest on the medial aspect of the maxilla, and a deeply invaginated fossa on the laterosphenoid. Unlike C. saharicus the ventral rim of the external antorbital fossa does not protrude laterally, the anteromedial process does not demarcate a deep ventral fossa, and the frontal-lacrimal suture is not elevated. The new material provides evidence of differentiation among Cenomanian-age faunas from the Sahara and supports a close relationship between Carcharodontosaurus, Giganotosaurus, and Acrocanthosaurus.

Highly incomplete taxa and the phylogenetic relationships of the theropod dinosaur Juravenator starki Richard J. Butler Department of Palaeontology , The Natural History Museum , Cromwell Road, London, SW7 5BD, United Kingdom E-mail: R.Butler@nhm.ac.uk & Paul Upchurch Department of Earth Sciences , University College London , Gower Street, London, WC1E 6BT, United Kingdom E-mail: P.Upchurch@ucl.ac.uk

Article

Mar 2007

An overview of non-avian theropod discoveries and classification

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