Dinosaur Field Guide Supplement 2nd Ed

The Dinosaur Field Guide Supplement
September 2010 December 2015
By, Zachary Perry (ZoPteryx)
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Disclaimer:
This supplement is intended to be a companion for Gregory S. Pauls impressive work

The Princeton Field Guide to Dinosaurs, and as such, exhibits some similarities in format, text, and
taxonomy. This was done solely for reasons of aesthetics and consistency between his book and this
supplement. The text and art are not necessarily reflections of the ideals and/or theories of Gregory S.
Paul.
The author of this supplement was limited to using information that was freely available from
public sources, and so more information may be known about a given species then is written or
illustrated here. Should this information become freely available, it will be included in future
supplements.
For genera that have been split from preexisting genera, or when new information about a
genus has been discovered, only minimal text is included along with the page number of the
corresponding entry in The Princeton Field Guide to Dinosaurs or a previous supplement. Genera
described solely from inadequate remains (teeth, claws, bone fragments, etc.) are not included, unless
the remains are highly distinct and cannot clearly be placed into any other known genera; this includes
some genera that were not included in Gregory S. Pauls work, despite being discovered prior to its
publication.
All artists are given full credit for their work in the form of their last name, or lacking this, their
username, below their work. Modifications have been made to some skeletal restorations for aesthetic
reasons, but none affecting the skeleton itself, except where noted. Should any artist want their piece
modified or removed from this supplement, they may contact the author. All life restorations are by the
author and based on the accompanying skeletal diagram. All artwork corresponds to the entry
immediately above it.
The author apologizes in advance for any typos, overlooked information, and inconsistencies
that may be present in this and future supplements.
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Dinosaurs
Baso-Dinosaurs
Nyasasaurus parringtoni
2.4 m (8 ft) TL, 15 kg (30 lb)
Fossil Remains: Partial skeleton.
Anatomical Characteristics: Three sacral vertebrae; neck vertebrae elongated, hollow, possibly pitted
as in theropods; long deltopectoral crest on humerus.
Age: Middle Triassic, Anisian.
Distribution: Tanzania; Manda Formation.
Habits: Growth apparently more rapid than herrerasaurs. Was likely omnivorous and capable of
running faster than most contemporary animals.
Notes: Tentatively placed within dinosauria. May be a common ancestor of both ornithischia and
saurischia, although some analyses have concluded that it is most likely a theropod. If it is not a
dinosaur, then it is the most derived protodinosaur yet known. First remains mistakenly placed in much
later Thecodontosaurus. It is not certain that the two known specimens belong under the same genus or
species.
Theropods
Herrerasaurs
Eoraptor lunensis
See Page 68
Notes: Possibly a very basal sauropodomorph or perhaps simply a basal saurischian.
Sanjuansaurus gordilloi
2.5 m (9 ft) TL, 30 kg (60 lb)
Fossil Remains: Minority of skull and partial skeleton.
Anatomical Characteristics: Standard for baso-theropods.
Age: Late Triassic, Carnian.
Distribution: Northern Argentina; Ischigualasto.
Habitat: Seasonally well-watered forests, including dense stands of giant conifers.
Habits: Generalist predator.
Notes: Prey included Pisanosaurus, Panphagia, and Eoraptor. Main enemies Herrerasaurus and large
thecodonts.
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Alcober & Martinez
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Derived Non-Avepods
Eodromaeus murphi
1.2 m (4 ft) TL, 5 kg (10 lb)
Fossil Remains: Majority of skull and skeleton.
Anatomical Characteristics: Relatively standard for baso-therapods, but overall more gracile and with a
less robust pubis than herrerasaurs.
Age: Late Triassic, Early Carnian.
Habitat: Seasonally well-watered forest, including dense stands of giant conifers.
Habits: Hunted small game.
Notes: More advanced than typical herrerasaurs but less advanced than avepods. Shared its habitat
with similar sized Eoraptor and larger Herrerasaurus, was likely prey of the latter. Prey included
Panphagia and Pisanosaurus.
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Sereno & Abraczinskas
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Avepods
Baso-Avepods
Daemonosaurus chauliodus
1.5 m (5 ft) TL, 5 kg (10 lb)
Fossil Remains: Complete skull and minority of skeleton.
Anatomical Characteristics: Snout short and sloping. Eyes large. Teeth in upper jaw elongated. Upper
jaw indented near tip and slightly longer than lower jaw. Teeth at front of upper and lower jaws are
procumbent.
Age: Late Triassic, Rhaetian.
Distribution: New Mexico; Chinle.
Habitat: Well-watered forest.
Habits: Probably preyed on small game.
Notes: More advanced than herrerasaurs, but may or may not be avepods. Shared its habitat with
Coelophysis, Chindesaurus, and Tawa.
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Sues et al.
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Tawa hallae
2.2 m (7 ft) TL, 13 kg (30 lb)
Fossil Remains: Complete skull and several skeletons, juvenile to adult.
Anatomical Characteristics: Overall build very similar to coelophysoids, but arms longer. Pelvis short
like herrerasaurs.
Age: Late Triassic, Early Norian.
Distribution: New Mexico; Chinle.
Habitat: Well-watered forests, including dense stands of giant conifers.
Habits: Capable of hunting small and medium game. Skeletons of various ages found together may
suggest some sort of group behavior.
Notes: Likely competed with Coelophysis and Chindesaurus. Enemies included large thecodonts. More
advanced than herrerasaurs, but less advanced than coelophysoids; may or may not be avepods.
Nesbitt et al.
http://www.deviantart.com/art/Tawa-the-perfect-intermediate-441064057
Hartman
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Tachiraptor admirabilis
1.5 m (5 ft) TL, 5 kg (10 lb)
Fossil Remains: Minority of skeleton.
Anatomical Characteristics: Insufficient information.
Age: Early Jurassic, Hettangian.
Distribution: Venezuela; La Quinta.
Notes: Potential prey included Laquintasaura.
Coelophysoids
Lepidus praecisio
1 m (3 ft) TL, 1 kg (2 lb)
Fossil Remains: Minority of skull and skeleton.
Age: Late Triassic, Early Norian.
Distribution: Texas; Dockum Group.
Notes: If this is a coelophysoid, it is the earliest known member.
Panguraptor (or Coelophysis) lufengensis
1.5 m (5 ft) TL, 5 kg (10 lb)
Fossil Remains: Majority of skull and partial skeleton.
Anatomical Characteristics: Snout and arms relatively short.
Distribution: Southern China; lower Lufeng.
Notes: More closely related to Coelophysis than to Syntarsus, suggesting that all should either be
lumped into Coelophysis, or Syntarsus should be elevated back to the genus level. Main enemy
Sinosaurus, potential prey included early mammals and young prosauropods.
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Studziski
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Zupaysaurus rougieri
See Page 75
Notes: Research indicates that Zupaysauruss large crests may actually be displaced lacrimal bone.
Sinosaurus triassicus
See Page 76
Notes: Dilophosaurus sinensis has been found to be a junior synonym of this genus and species.
Abelisauroids
Baso-Abelisauroids
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Fosterovenator churei
Adult size not certain
Age: Late Jurassic, Middle Tithonian.
Distribution: Wyoming; upper Morrison.
Habitat: Semiarid with open floodplain prairies and riverine forests.
Notes: Classification uncertain, probably the first known basal abelisauroid from the northern
hemisphere. May include remains assigned to Elaphrosaurus? unnamed species. Shared its habitat with
the much larger Allosaurus and Torvosaurus, amongst many other dinosaurs.
Dahalokely tokana
3.5 m (12 ft) TL, 40 kg (85 lb)
Anatomical Characteristics: Robustly constructed.
Age: Late Cretaceous, Turonian.
Distribution: Madagascar; Diego Basin.
Notes: Taxonomic placement uncertain due to incompleteness of remains. May be a large basal
noasaurid, as some evidence suggests, or a small abelisaurid. At the time of Dahalokelys existence,
Madagascar and India were linked together as a large island.
Abelisaurids
Eoabelisaurus mefi
6 m (20 ft) TL, 750 kg (1,600 lb)
Fossil Remains: Partial skull and majority of skeleton.
Anatomical Characteristics: Skull tall and arms shortened, but still functional. Vertebral spines over
hips and base of tail form shallow ridge.
Age: Middle Jurassic, Late Aalenian or Early Bajocian.
Distribution: Southern Argentina; Caadn Asfalto.
Notes: Oldest known abelisaurid. Shared its habitat with Piatnitzkysaurus, Condorraptor, and a variety
of sauropods.
Rauhut
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Kryptops palios
See Page 78
Notes: Portions of skeleton may belong to Eocarcharia.
Pycnonemosaurus nevesi
See Page 79
Notes: Correct spelling of Pycnoneosaurus.
Majungasaurus crenatissimus
See Page 80
Anatomical Characteristics: Neck robust and fairly long, shorter legs result in a low-slung appearance.
Headden
Rahiolisaurus gujaratensis
7.5 m (25 ft) TL, 2 tonnes
Fossil Remains: Minority of several skeletons found in association.
Anatomical Characteristics: Leg long and gracile.
Age: Late Cretaceous, Maastrichtian.
Distribution: Western India; Lameta.
Habits: Hunted titanosaur sauropods and ankylosaurs.
Notes: May be the same as contemporary Indosuchus raptorius. Shared its habitat with Rajasaurus.
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Vitakridrinda sulaimani
Distribution: Pakistan; Pab.
Notes: Appears fairly standard for an abelisaurid, but too little known to be certain.
Arcovenator escotae
6 m (20 ft) TL, 750 kg (1,600 lb)
Anatomical Characteristics: Bony brow ridges well developed.
Age: Late Cretaceous, Late Campanian.
Distribution: France; Lower Argiles Rutilantes.
Habitat: Large subtropical island with large rivers.
Notes: Potential prey included titanosaurs and rhabdodonts.
Noasaurids
Velocisaurus unicus
1.5 m (5 ft) TL, 15 kg (30 lb)
Anatomical Characteristics: Middle toe enlarged, outer toe atrophied. Claws short and straight.
Age: Late Cretaceous, Santonian.
Distribution: Northern Argentina; Bajo de la Carpa.
Habits: Peculiar foot anatomy, unlike any other known dinosaur, likely an adaptation for running.
Notes: Prey may have included Alvarezsaurus and flightless birds. Shared its habitat with Bonitasaura.
Averostrans
Elaphrosaurs
Ceratosaurs
Genyodectes serus
Fossil Remains: Minority of skull.
Anatomical Characteristics: Snout robust, teeth long and dagger-like.
Age: Early Cretaceous, Aptian.
Distribution: Southern Argentina; Cerro Barcino.
Notes: Classification uncertain, but arrangement of teeth most like that of Ceratosaurus. Shared its
habitat with Tyrannotitan and Chubutisaurus.
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Camarillasaurus cirugedae
3 m (10 ft) TL, 70 kg (150 lb)
Age: Early Cretaceous, Early Barremian.
Distribution: Spain; Camarillas.
Notes: The only Cretaceous age non-abelisauroid ceratosaur known from Eurasia thus far. Prey
included hypsilophodonts and iguanodonts. Exact classification uncertain, may be an elaphrosaur.
Tetanurans
Baso-Tetanurans
Kayentavenator elysiae
Fossil Remains: Minority of skeleton, juvenile.
Age: Early Jurassic, Pliensbachian.
Distribution: Arizona; Kayenta.
Habitat: Well-watered forest, likely surrounded by arid regions.
Notes: May be one of the earliest tetanurans or an advanced coelophysoid. Shared its habitat with
Dilophosaurus.
Chilesaurus diegosuarezi
3 m (10 ft) TL, 70 kg (150 lb)
Fossil Remains: Majority of skull and several partial skeletons, juvenile to adult.
Anatomical Characteristics: Skull small and snout at least moderately short, though its exact length is
uncertain. Teeth are elongated, unserrated, and slightly procumbent. A short beak may have been
present. Neck and body long and gracile. Arms moderately long, hands bear only two functional
fingers; third finger greatly reduced and likely vestigial. Hand claws are slightly hooked. Pubis slightly
retroverted. Legs moderately long. Hallux enlarged and weight bearing, as in unrelated therizinosaurs.
Age: Late Jurassic, Late Tithonian.
Distribution: Chile; Toqui.
Habits: Likely omnivorous, browsing on plant matter as well as catching insects and small vertebrates.
Notes: The earliest confirmed example of herbivory in a theropod, this unusual dinosaur bears a
mixture of traits seen in ornithischians, basal sauropodomorphs, basal theropods, coelurosaurs, and
non-dinosaurian shuvosaurids. Had some of remains not been found in full articulation, they would
have likely been classified as many different taxa. Exact classification uncertain, may be close to
elaphrosaurs, basal avepods, or, conversely, to coelurosaurs. Either way, a rather large ghost lineage
must still be hidden in the fossil record. Eshanosaurus deguchiianus, an enigmatic species from the Early
Jurassic of China, bears a lower jaw and procumbent teeth that are quite similar to that of Chilesaurus,
and therefore could be related.
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Novas et al.
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Megalosaurs
Cruxicheiros newmanorum
9 m (30 ft) TL, 2 tonnes
Age: Middle Jurassic, Bathonian.
Distribution: Central England; Chipping Norton Limestone.
Notes: Originally placed in Megalosaurus. Taxonomic placement uncertain, may be a basal carnosaur.
Leshansaurus qianweiensis
7 m (21 ft) TL, 700 kg (1,500 lb)
Fossil Remains: Partial skull and skeleton.
Anatomical Characteristics: Snout fairly wide toward front.
Age: Middle to Late Jurassic, Bathonian to Callovian.
Distribution: Central China; Dashanpu, Shangshaximiao.
Habitat: Heavily forested.
Habits: Potential prey included sauropods and stegosaurs. Purpose of widened snout unclear, may
have allowed for a larger bite when attempting to wound large prey.
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Notes: Originally classified as a sinraptorid, is more likely a megalosaur. Snout shows some similarities
to Afrovenator.
Torvosaurus gurneyi
Anatomical Characteristics: More robust and with fewer teeth than T. tanneri.
Age: Late Jurassic, Late Kimmeridgian.
Distribution: Portugal; Lourinh.
Habitat: Large seasonally dry island with open woodlands.
Habits: Big game hunter, potential prey included sauropods and stegosaurs.
Notes: Theropod eggs and embryos found in the region likely belong to this species.
Sciurumimus albersdoerferi
Fossil Remains: Nearly complete juvenile skull and skeleton, external fibers.
Anatomical Characteristics: In juveniles: Snout subtriangular. Tail fairly long. Simple protofeathers
cover much of body, particularly long over base of tail.
Age: Late Jurassic, Early Tithonian.
Distribution: Southern Germany; Painten.
Habitat: Semi-arid islands.
Habits: Likely hunted small game. Adults diet unknown.
Notes: The first non-coelurosaur found with definitive feathers. It is unknown if these feathers were
retained into maturity. Although superficially similar to Juravenator starki, the two appear to be quite
distinct from one another. Not all researchers agree that Sciurumimus is a megalosauroid, preferring to
classify it as a very basal coelurosaur. As physical features can change with maturity, any exact
classification is tentative. Skeletal shown here is only partially restored.
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Spinosaurs
Notes: Fragmentary remains suggest presence in Australia. Other fragments suggest that derived
spinosaurs may have existed as early as the Late Jurassic.
Ichthyovenator laosensis
8.5 m (28 ft) TL, 1.7 tonnes
Anatomical Characteristics: Vertebral spines moderately long but show an abrupt decrease in height
over hips, and then rise again over base of tail. This gap appears to be natural, but it is unclear if it
would have been visible in life.
Distribution: Laos; Grs suprieurs.
Habitat: Coastal river delta.
Notes: The first definitive spinosaur from Asia. Probably includes Siamosaurus suteethorni, which is
based on inadequate remains. Shared its habitat with sauropods and iguanodonts.
Spinosaurus aegyptiacus
See Page 88 15 m (49 ft) TL, 10 tonnes
Fossil Remains: Minority of skulls and partial skeletons, juvenile and adult.
Anatomical Characteristics: Snout pitted with receptor sites, neck fairly long, hand rather long, tall
(over 2 m tall) skin-covered sail over body, pelvis small, legs barely longer than arms. Toes bear
flattened claws and the hallux is elongated and lowered, so feet likely webbed in life. Limb bones are
quite dense internally.
Habits: Spent most of its time in the water; either walking on the bottom when in the shallows or
paddling with feet when in deeper water, paddling motion likely accompanied by horizontal undulation
of the tail. Sensory pits on snout, like those of crocodilians, suggest that Spinosaurus was a tactile
feeder of aquatic prey; probably either a sit-and-wait predator of passing game or moved slowly along
feeling with snout, akin to a stork. Though sail likely played a role in thermoregulation, greatly
exaggerated size indicates a primary function of communication to other members of its species,
perhaps allowing rivals to assess the size and strength of each other while the majority of the body was
submerged. Far forward center of gravity and short legs imply that quadrupedal movement was
employed on land, but this is not certain, and a slow bipedal gate is perhaps more likely.
Notes: Current skeletal model is a composite of Egyptian and Moroccan material from specimens of
varying maturity, thus it raises numerous concerns regarding reliability. Recent studies indicate the
Moroccan material can safely be regarded as that of a spinosaur, possibly Spinosaurus, but no material
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overlaps the original Egyptian remains for comparison, so these could also belong to Sigilmassasaurus
and/or an unknown third taxa. Nevertheless, the basic anatomy outlined above still likely holds true, as
all of the aforementioned species are close relatives. Proposed quadrupedal movement on lands has
received numerous criticisms as the theropod pectoral girdle and forelimbs are not designed to support
weight or move in a fashion conducive to walking. Short hips and legs are scaled up from the remains of
a Moroccan juvenile; as ontogenic changes in dinosaurs can be profound, its possible that the hips and
legs in a mature adult would have been proportioned more normally.
Ibrahim et al.
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Sigilmassasaurus (or Spinosaurus) brevicollis
Fossil Remains: Neck vertebrae, possibly an unknown portion of skull and skeleton.
Age: Late Cretaceous, Early Cenomanian.
Distribution: Morocco; Kem Kem.
Habitat: Coastal mangroves, lagoons, and floodplains.
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Habits: Likely an aquatic or semi-aquatic predator akin to its relatives.

Notes: Resurrected genus split from Spinosaurus aegyptiacus, as its cervical vertebrae have been
demonstrated to be distinct. Includes remains assigned to Spinosaurus maroccanus and possibly Ernst
Stromers destroyed Spinosaurus B. Other spinosaurid remains from the Kem Kem, such as a partial
rostrum, pelvis, and hind limbs, cannot be directly compared to either Spinosaurus or Sigilmassasaurus
due to a lack of overlapping elements. However, the presence of additional cervical vertebrae distinct
from Sigilmassasaurus suggests that at least one other species of giant spinosaurid, possibly
Spinosaurus, was contemporaneous with Sigilmassasaurus. Whether the two avoided competition via
niche partitioning or if there was simply enough prey to go around remains to be seen.
Oxalaia (or Spinosaurus/Sigilmassasaurus) quilombensis
Distribution: Eastern Brazil; Alcntara.
Notes: Known from only a snout tip and a few other fragments. May belong to one of the better known
giant North African spinosaurs.
Avetheropods
Baso-Avetheropods
Carnosaurs
Carnosaur Miscellanea
Allosauroids
Sinraptorids
Metriacanthosaurus parkeri
8 m (25 ft) TL, 1 tonne
Age: Late Jurassic, Callovian.
Distribution: Southern England; Oxford Clay.
Notes: Classification uncertain. May be a more derived carnosaur.
Allosaurids
Allosaurus lucasi
Fossil Remains: Partial skull and minority of skeleton, juvenile and adult.
Anatomical Characteristics: Skull more robustly constructed then other Allosaurus species.
Age: Late Jurassic, Tithonian.
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Distribution: Colorado; upper Morrison.

Habitat: Semiarid with open floodplain prairies and riverine forests.
Habits: Big game hunter.
Notes: May include remains assigned to Allosaurus atrox.
Lourinhanosaurus (or Allosaurus) antunesi
See Page 96
Notes: Eggs with embryos found nearby likely belong to this species. Classification uncertain, may be a
basal coelurosaur.
Carcharodontosaurids
Veterupristisaurus milneri
Age: Late Jurassic, Late Kimmeridgian/Early Tithonian.
Distribution: Tanzania; middle Tendaguru.
Habitat: Coastal, seasonally dry with heavier vegetation further inland.
Notes: Fairly large in size. If this is a carcharodontosaurid, it is the earliest known.
Aerosteon riocolloradensis
See Page 99
Notes: Correct spelling of Aerosteons.
Eocarcharia dinops
See Page 97
Notes: A pelvis assigned to Kryptops may belong to this species.
Becklespinax altispinax
Fossil Remains: A few vertebrae.
Anatomical Characteristics: At least some vertebral spines relatively tall.
Age: Early Cretaceous, Valanginian.
Distribution: Southeast England; Hastings Beds.
Notes: Exact placement of tall spined vertebrae uncertain, initially thought to be above the shoulders
but may actually be closer to the hips as in Concavenator. Taxonomic placement questionable, could be
another type of allosauroid or even a spinosaurid. Shared its habitat with Hylaeosaurus and
iguanodonts.
Concavenator corcovatus
6 m (20 ft) TL, 500 kg (1,000 lb)
Fossil Remains: Majority of skull and skeleton, skin impressions.
Anatomical Characteristics: Vertebrae just before hips rise sharply to form a sail-like structure
extending over hips and likely connecting to second smaller rise in vertebrae at base of tail.
Alternatively, these may have formed separate hump-like structures. Bony knobs along back of ulna may
be quill nodes. Broad subrectangular scales line the underside of the tail, bird-like scutes cover the feet,
and the underside of the feet possessed pads.
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Age: Early Cretaceous, Barremian.

Distribution: Spain; Calizas de al Huergina.
Habitat: Well-watered forest and floodplains.
Habits: Sail or hump almost certainly had a display function, but potential mechanical use is unclear.
May have been used in thermoregulation and/or as a site for muscle attachments. Preyed included
sauropods, iguanodonts, and Pelecanimimus.
Notes: Bony knobs on ulna may actually be muscle attachment points, as their arrangement is not
consistent with those of normal quill nodes.
Sinkkonen
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Sauroniops pachytholus
Anatomical Characteristics: Skull roof robust in structure.
Distribution: Morocco; Kem Kem Beds.
Habitat: Coastal region with arid interior.
Notes: Could potentially be a large abelisaurid, but carcharodontosaurid identity is more likely.
Neovenatorids
Notes: Subgroup megaraptora, which includes all members except Neovenator and Chilantaisaurus,
may actually be basal coelurosaurs close to tyrannosauroids. All are left within neovenatorids at this
time until the taxonomy of the group can be resolved.
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Megaraptor namunhuaiquii
See Page 99
Fossil Remains: Minority of skeletons, adult. Majority of skull and skeleton, juvenile.
Anatomical Characteristics: In large juveniles: Skull narrow and subtriangular, teeth relatively small.
Arms well developed with slender hand claws, legs long and gracile.
Habits: Likely a pursuit predator that used its teeth and claws to wound smaller prey.
Notes: Juvenile remains suggest placement of this species and relatives within the tyrannosauroids.
Until mature specimens can be found to test this theory, it has been left in its traditional position within
neovenatorids.
Get Away Trike! Blog
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Siats meekerorum
Fossil Remains: Partial skeleton, large juvenile.
Anatomical Characteristics: Fairly robust.
Age: Early Creatceous, Early Cenomanian.
Distribution: Utah; Upper Cedar Mountain.
Habitat: Semiarid open woodland.
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Habits: Big game hunter.

Notes: Establishes the presence of neovenatorids in North America and shows that they were capable
of reaching the same sizes as carcharodontosaurids. Potential prey included nodosaurs, iguanodonts,
and basal hadrosaurs.
Datanglong guangxiensis
Age: Early Cretaceous.
Distribution: Eastern China; Xinlong.
Notes: Originally classified as a basal allosauroid, is more likely a neovenatorid or perhaps a basal
tyrannosaur.
Coelurosaurs
Tyrannosauroids
Baso-Tyrannosauroids
Guanlong wucaii
See Page 93
Notes: Almost certainly a tyrannosauroid related to Proceratosaurus and Kileskus.
Proceratosaurus bradleyi
See Page 123
Notes: Most likely a basal tyrannosauroid close to Guanlong and Kileskus.
Kileskus aristotocus
3 m (10 ft) TL, 70 kg (150 lb)
Anatomical Characteristics: May have had a large midline crest.
Age: Middle Jurassic, Bathonian.
Distribution: Central Russia; Itat.
Notes: Closest relative Proceratosaurus, and like it, probably a basal tyrannosauroid.
Juratyrant langhami
See Page 100
Notes: New genus for Stokesosaurus langhami.
Yutyrannus huali
9 m (30 ft) TL, 2.5 tonnes
Fossil Remains: Complete skull and nearly complete skeletons, external fibers, juvenile and adult.
Anatomical Characteristics: Body standard for baso-tyrannosauroid. Low wavy midline crest on snout.
Brow hornlets prominent. Arms well developed. Simple feathers up to 20 cm (8 inches) long covered
most of body and limbs.
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Distribution: Northeast China; Yixian.

Habitat: Well-watered forests and lakes.
Habits: Midline crest and brow hornlets were for display. Long feathers were likely for insulation in the
cool Yixian climate. Two adults found together with a juvenile suggest this species may have lived in
small family groups. Shows a standard tyrannosauroid growth pattern; limbs shorten and skull becomes
more robust as age progresses.
Notes: The largest dinosaur known to preserve direct evidence of feathers. The largest known predator
in its habitat, prey included a variety of smaller therapods, ceratopsians, and ornithopods. Some
features of the skull resemble those of allosauroids.
Anton
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http://scotthartman.deviantart.com/art/Beautiful-feathered-tyrant-head-comparison-294353870
Hartman
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Derived Tyrannosaurs
Raptorex kriegstenis
See Page 101
Notes: Found to be from younger sediments than the Yixian, most likely from the Nemegt. Also shown
to most likely be a young juvenile, rather than a small adult. These factors combine to suggest that
Raptorex is actually the juvenile of a larger tyrannosaur species, possibly Tarbosaurus.
Tyrannosaurids
Qianzhousaurus (or Alioramus) sinensis
Fossil Remains: Complete skull and partial skeleton.
Anatomical Characteristics: Snout very long, low, and shallow. Crenulated midline crest on snout.
Distribution: Southeastern China; Nanxiong.
Habits: Likely preferred smaller game, despite large size.
Notes: Closely related to, if not a mature specimen of, Alioramus, and with them likely represents a
unique East Asian radiation of long-snouted tyrannosaurids. Some studies have found these longsnouted tyrannosaurs to be the sister group of true tyrannosaurids, rather than actually members.
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Lythronax argestes
7.5 m (25 ft) TL, 2.5 tonnes
Anatomical Characteristics: Skull short and stout. Body robust.
Age: Late Cretaceous, Early Campanian.
Distribution: Utah; Wahweap.
Habits: Likely an ambush predator adapted to tackling robust game, such as ceratopsians.
Notes: The earliest member of the lineage that would lead to Tyrannosaurus.
http://scotthartman.deviantart.com/art/All-hail-the-King-of-Gore-412767668
Hartman
Loewen et al.
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Teratophoneus curriei
Fossil Remains: Partial skulls and skeletons.
Anatomical Characteristics: Snout short and number of teeth reduced.
Distribution: Utah; Kaiparowits.
Habitat: Riverine forests and floodplains.
Habits: Short snout probably an adaptation for hunting well protected game, such as ceratopsians.
Notes: Prey included Kritosaurus, Utahceratops, Kosmoceratops, and Hagryphus. Along with
Bistahieversor, likely represents a unique lineage of tyrannosaurids in southwestern North America.
Carr et al.
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http://scotthartman.deviantart.com/art/Teratophoneus-the-monster-killer-268423426
Hartman
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Zhuchengtyrannus magnus
Fossil Remains: Partial skull.
Anatomical Characteristics: Robustly constructed. Teeth bear extensive serrations.
Age: Late Cretaceous, Campanian.
Distribution: Eastern China; Wangshi Group.
Notes: Prey included Sinoceratops, Zhuchengceratops, Tanius and Tsintaosaurus.
Nanuqsaurus hoglundi
Age: Late Cretaceous, Late Maastrichtian.
Distribution: Northern Alaska; Prince Creek.
Habitat: Polar forest with warm humid summers and cold dark winters.
Notes: Originally considered a species of Albertosaurus/Gorgosaurus, is instead more closely related to
tyrannosaurines such as Tyrannosaurus. Potential prey included pachycephalosaurs, ceratopsians, and
hadrosaurs.
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Ornithomimosaurs
Baso-Ornithomimosaurs
Hexing qingyi
1.1 m (3.5 ft) TL, 5 kg (12 lb)
Anatomical Characteristics: Snout narrow, tip of upper jaw curves downward. Tiny teeth restricted to
front of jaws.
Age: Early Cretaceous, Valanginian or Early Barremian.
Distribution: Northeast China; lower Yixian.
Notes: The smallest known ornithomimosaur. Main enemy Sinornithosaurus.
Jin et al.
ZP
Deinocheirus mirificus
See Page 112 11 m (36 ft) TL, 6 tonnes
Fossil Remains: Complete skull and majority of skeletons, gastroliths.
Anatomical Characteristics: Snout elongated, upper jaw shallow and terminates in a fairly broad
squared-off beak, lower jaw very robust and terminates in a short cropping beak. Cheeks likely present
along posterior half of jawline, no teeth present. Neck and arms are proportional normal for this large
an ornithomimosaur, digits end in blunt hooked claws. Relatively narrow body is tilted upright by large
hips and bears tall robust vertebral spines over midsection and hips, likely culminating in a camel-like
hump in life. Legs relatively short, short toes end in blunt claws, hallux absent. Last few vertebrae of tail
fused into a pygostyle, may have supported a small feather fan.
Habits: Small fish bones in stomach hint at an omnivorous lifestyle in and around swampy terrain.
Hadrosaur-like beak, lack of teeth, robust lower jaw but weak bite force, and presence of many
gastroliths suggest that the majority of this species diet was composed of soft low-growing plant matter
with animal matter likely being taken opportunistically.
Notes: Bizarre anatomy likely adaptations to supporting its supersized lifestyle, such as the wide beak
to maximize eating efficiency and tall vertebral spines lined with stiffening ligaments that helped
support the neck and stomach. Closest relatives appear to be Garudimimus and Beishanlong, together
they form a sister-group to the ornithomimids. One skeleton shows evidence of being fed upon by
Tarbosaurus. Evolution of a pygostyle may be independent of that of more advanced coelurosaurs.
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Lee et al.
ZP
Lee et al.
ZP
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27
Lee et al.
Ornithomimids
Qiupalong henanensis
Anatomical Characteristics: Smaller than other derived ornithomimids.
Age: Late Cretaceous, probably Campanian.
Distribution: Central China; Qiupa.
Notes: Closest relatives appear to be North American forms like Struthiomimus. Main enemy
Luanchuanraptor.
Anserimimus planinychus
See Page 113
Notes: Correct spelling of Ansermimus.
Struthiomimus (or Ornithomimus) sedens
See Page 117
Anatomical Characteristics: Short protofeathers found on neck, back, and legs; longer pennaceous
feathers on lower arm of adults. Legs devoid of feathers from mid-thigh downward, scales apparently
absent. Underside of the tail bare as well, however, long protofeathers along the top and sides of the
tail likely hung down and obscured this. A flap of skin connected the upper thigh to the torso region, a
reduced version of what is seen in modern birds.
Notes: Long arm feathers, only found in adults, appear ragged and most likely used for display.
Tototlmimus (or Struthiomimus) packardensis
4 m (12 ft) TL, 170 kg (370 lb)
Anatomical Characteristics: Legs long.
Distribution: Mexico; Packard Shale.
Notes: The most southerly known North American ornithomimosaur.
Maniraptors
Compsognathids
Juravenator starki
See Page 118
Anatomical Characteristics: Small scales cover most of tail, legs, and snout. Simple protofeathers at
base of tail and above hips, most likely covered more of body.
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28
Sinocalliopteryx gigas
See Page 121
Habits: Known to have consumed smaller dinosaurs, including Sinornithosaurus and a small
ornithischian, and the early bird Confuciusornis. Stomachs containing multiple prey items suggest that
Sinocalliopteryx had a very high metabolism. Stomachs also found to contain small gastroliths.
Maniraptor Miscellanea
Aorun zhaoi
Fossil Remains: Complete skull and partial skeleton, juvenile.
Anatomical Characteristics: Head subtriangular. Thumb claw larger and more curved compared to
other claws on hand.
Age: Late Jurassic, Early Oxfordian.
Distribution: Western China; Shishugou.
Notes: Difficult to classify due to the immaturity of the specimen. A very basal coelurosaur, may not be
a maniraptor, but it is seemingly more advanced than tyrannosauroids and possibly ornithomimosaurs.
Main enemies included Sinraptor, Monolophosaurus, Guanlong, and Zuolong; it does not appear to be a
juvenile of any of these.
Zuolong salleei
3 m (10 ft) TL, 70 kg (150 lb)
Anatomical Characteristics: More robust than compsognathids, snout subtriangular.
Distribution: Western China; Shishugou.
Notes: Main enemy Yangchuanosaurus (=Sinraptor) dongi.
Choiniere et al.
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ZP
Bicentenaria argentina
3 m (10 ft) TL, 60 kg (130 lb)
Fossil Remains: Majority of skull and partial skeleton. Numerous unevaluated remains.
Anatomical Characteristics: Snout subtriangular, hand short, legs long.
Age: Late Cretaceous, Middle Turonian.
Distribution: Western Argentina; Portezuelo.
Habitat: Well-watered woodlands with short dry season.
Habits: A pursuit predator of small- and medium-sized game.
Notes: Little available information on this genus, despite apparent completeness of remains. Site
locality contains a number of individuals that have yet to be excavated or thoroughly examined. May be
a basal tyrannosauroid. Main enemy Megaraptor. Prey included Macrogryphosaurs and young
titanosaurs.
ZP
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ZP
Scansoriopterygids
Epidexipteryx hui
See Page 147
Notes: Almost certainly a scansoriopterygid rather than an oviraptorosaur.
Yi qi
0.5 m (1.7 ft) TL, 0.4 kg (0.85 lb)
Fossil Remains: Complete skull and majority of skeleton, impressions of feathers and wing membranes.
Anatomical Characteristics: Skull subtriangular, jaws slightly downturned, peg-like procumbent teeth
restricted to the ends of jaws. Nostrils retracted from snout tip, eyes large. Arms very long. Hand very
long and strongly asymmetric due to hyper-elongation of third finger, fingers not fused to each other.
Claws moderately hooked, more so on first finger. A slightly bent bony or cartilaginous prong, identified
as a styliform, projects from the back of the wrist; exact orientation uncertain. Legs fairly long. Feet and
tail unknown. The feathers have a paintbrush-like structure, with many smaller filaments emerging
from a central flattened structure that accounts for the majority of the feathers length. Feathering
extends from just behind the nostrils, down both sides of the neck, throughout the body, down the arms
to the center of the hand, and down along the legs to at least the ankles. Feathers are longest about the
neck, upper arm, and lower leg. Patches of striated membranous skin are preserved between the
fingers and between the third finger and the styliform, likely forming a gliding membrane, the exact
shape of which is uncertain. The striations in the membrane may be stiffening fibers or merely folds on
the skin. Large melanosomes are present in the feathers of head, neck, and legs, likely corresponding to
darker coloration in these regions.
Distribution: Northeast China; Tiaojishan.
Habitat: Warm, well-watered forest.
Habits: Likely a nocturnal or crepuscular insectivore or omnivore. Long legs suggest it fed on the
ground, but climbing ability undoubtedly present as it would have needed elevated perches in order to
start its glides; its pectoral anatomy was not well suited for powered flight. Gliding would be a very
energy efficient way to search large areas for suitable foraging sights, such as fallen logs, where its
downturned snout and procumbent teeth would be useful in stripping bark in search of prey. Since the
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31
elongated third finger was incorporated into the flight membrane, it was probably not used to probe
crevices like an aye-aye.
Notes: Unique wing structure of Yi likely represents an independent evolution of volant tendencies to
that of birds, perhaps one of several amongst maniraptors. The incorporation of the elongated fingers
into the flight membrane may have been a driving factor of their hyper-elongation in this group; perhaps
all long-fingered scansoriopterygids had gliding membranes of some sort. Shared its habitat with
Anchiornis, Pedopenna, Xiaotingia, Eosinopteryx, Scansoriopteryx, Epidexipteryx, Aurornis, and
Tianyulong. There is a possibility that Yi is the adult form of one of the other scansoriopterygid species it
shared its habitat with, Scansoriopteryx and Epidexipteryx, the juvenile nature for the former perhaps
making it the most likely suspect.
Xu et al.
ZP
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Alvarezsaurs
Alvarezsaurids
Albinykus baatar
0.6 m (2 ft) TL, 1 kg (2 lb)
Fossil Remains: Complete legs and pelvis.
Anatomical Characteristics: Bones in feet fused together.
Distribution: Mongolia; Javkhlant.
Notes: Preserved in a sitting position reminiscent of modern birds.
Alnashetri cerropoliciensis
1 m (3.3 ft) TL, 3.5 kg (8 lb)
Age: Late Cretaceous, Cenomanian.
Distribution: Western Argentina; Candeleros.
Habitat: Well-watered woodlands with short dry season.
Notes: Main enemy Ekrixinatosaurus. Shared its habitat with Buitreraptor.
Bonapartenykus ultimus
2.5 m (8 ft) TL, 25 kg (50 lb)
Fossil Remains: Minority of skeleton, egg shell fragments.
Age: Late Cretaceous, Late Campanian or Early Masstrichtian.
Distribution: Western Argentina; Allen.
Notes: Shared its habitat with Saltasaurus.
Heptasteornis andrewsi
Distribution: Romania; Maastrichtian Snpetru.
Notes: Although the remains are very fragmentary, they are almost certainly those of a small
alverezsaur, the first known from Europe.
Linhenykus monodactylus
0.6 m (2 ft) TL, 1 kg (2 lb)
Anatomical Characteristics: Completely lacked remnants of other fingers, neck relatively long.
Distribution: Northern China; Wulansuhai.
Notes: Despite derived hands, this is still a relatively basal alvarezsaurid.
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Xu et al.
Xixianykus zhangi
0.4 m (1.3 ft) TL, 0.25 kg (0.5 lb)
Anatomical Characteristics: Pubis strongly retroverted, foot strongly compressed from side to side.
Distribution: Eastern China; Majiacun.
Notes: The oldest known derived alvarezsaurid.
Avepectorans
Deinonychosaurs
Deinonychosaur Miscellanea
Pneumatoraptor fodori
0.7 m (2.5 ft) TL, 0.6 kg (1.3 lb)
Distribution: Hungary; Csehbnya.
Habits: Hunted small game.
Notes: May be a dromaeosaurid or troodontid. Shared its habitat with Hungarosaurus.
Anchiornis huxleyi
See Page 128
Notes: Considered a primitive avian by some researchers.
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Xiaotingia zhengi
0.5 m (1.7 ft) TL, 0.5 kg (1.1 lb)
Fossil Remains: Complete skull and majority of skeleton.
Anatomical Characteristics: Overall very similar to Archaeopteryx, but arm and especially hand shorter.
Teeth more robust. Hyperextendable toe well developed, but sickle claw not large.
Distribution: Northeast China; probably Tiaojishan.
Habits: Flight ability inferior to that of Archaeopteryx, if present at all. Diet included small game and
possibly seeds.
Notes: Originally thought to be an archaeopterygid, probably more closely related to Anchiornis. Exact
origin of fossil unclear, slightly more derived deinonychosaur traits may lend toward the younger Yixian
Formation. Considered a primitive avian by some researchers.
Xu et al.
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ZP
Eosinopteryx brevipenna
0.3 m (1 ft) TL, 0.25 kg (0.5 lb)
Fossil Remains: Complete skull and majority of skeleton, feathers.
Anatomical Characteristics: Snout short, eyes large. Arms long, with well-developed symmetrical
primary feathers; orientation of wing bones and shape of feathers made flapping flight improbable.
Toes slender with unhooked claws, second toe not hyperextendable, hallux semireversed, and lower leg
lacked feathers. Tail relatively short and may have lacked a pennaceous feather fan.
Age: Late Jurassic, Oxfordian.
Habitat: Well-watered subtropical forest.
Habits: Lack of feathers on feet and inability to flap indicates a more terrestrial existence than other
small feathered deinonychosaurs of its age; legs and feet well suited for running. Likely a nocturnal
predator of small game.
Notes: Originally classified as a basal troodont, rather than a basal deinonychosaur. Lack of a feather
fan at the end of its tail could be gender or age related, or a retained basal trait. Shared its habitat with
Anchiornis, Xiaotingia, Aurornis, and Tianyulong.
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Herran
ZP
Pamparaptor micros
0.7 m (2 ft) TL, 1 kg (2 lb)
Anatomical Characteristics: Foot construction primitive.
Distribution: Southern Argentina; Portezuelo.
Notes: Originally considered to be the same as Neuquenraptor. Exact classification uncertain, shows
some similarities to troodontids.
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Balaur bondoc
2 m (6 ft) TL, 20 kg (40 lb)
Anatomical Characteristics: Robustly constructed. Outer finger vestigial, claws on remaining fingers
strongly hooked. Pubis highly retroverted. Hyperextendable hallux faces forward and bears sickle claw
along with inner toe.
Age: Late Cretaceous, Early Maastrichtian.
Distribution: Romania; Sebe.
Habitat: Forested island.
Habits: Robust legs, reduced hands, and retroverted pubis suggest that this species ate more plant
matter than most other deinonychosaurs. Probably also an opportunist of small game.
Notes: Originally thought to be velociraptorine dromaeosaurid, however, recent studies suggest that
this is species is in fact a large and highly unusual member of a lineage closer to birds that also includes
such species as Anchiornis and Xiaotingia. Unusual anatomy likely the result insular isolation.
Ellison
Archaeopterygids
Aurornis xui
0.5 m (1.7 ft) TL, 0.5 kg (1.1 lb)
Fossil Remains: Complete skull and skeleton, feathers.
Anatomical Characteristics: Head subtriangular with highly pointed snout and small teeth. Neck
relatively short. Body not deep. Pelvis short and pubis nearly vertical. Arm and hand very long, but
unable to flap. Legs similar in length to forelimbs, ankles relatively short. Second toe not
hyperextendable, hallux semireversed. Tail moderately long. Body covered in short feathers. Exact
arrangement of symmetrical primary feathers uncertain, may have lacked feathers on feet and ankles.
Habits: Probably incapable of powered flight, some gliding possible. Fed on small game.
Notes: Classification as a basal archaeopterygid not certain. The most derived non-avian known, or the
most primitive avian known, depending on the position of Archaeopteryx and other basal
deinonychosaurs. Shared its habitat with Anchiornis, Xiaotingia, Eosinopteryx, and Tianyulong.
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Herran
ZP
Dromaeosaurids
Microraptorinae
Sinornithosaurus millenii
See Page 132
Anatomical Characteristics: Mottled coloration composed of rusty red, black, and gray.
Notes: Despite popular claims, was not venomous.
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Microraptor zhaoianus
See Page 133
Anatomical Characteristics: First few teeth of lower jaw slightly procumbent. Two streamer-like
feathers at end of tail. All feathers entirely glossy black, however, some individuals show color banding
on primary feathers.
Habits: Known to have preyed on small mammals, birds, and fish. Streamer feathers on tail and glossy
coloration suggest that Microraptor may have been fairly social like many glossy black birds today, such
as starlings, crows, and grackles. Differences in coloration amongst individuals likely represent sexual or
age differences.
Notes: Likely includes M. hanqingi.
Changyuraptor yangi
1.2 m (4 ft) TL, 5 kg (10 lb)
Fossil Remains: Complete skull and skeleton, feathers.
Anatomical Characteristics: Tail long with long feathers (up to 30 cm) running its length. Long primary
feathers on arms and legs extending onto feet.
Age: Early Cretaceous, Early Aptian.
Habitat: Well-watered forest and lakes.
Habits: Small game hunter, able to attack larger prey than Microraptor. Aerial abilities likely inferior to
those of Microraptor due to its larger size and long tail feathers.
Notes: Possible descendent of Microraptor. Shared its habitat with a wide variety of other dinosaurs.
Dromaeosaurid Miscellanea
Zhenyuanlong suni
2 m (6 ft) TL, 20 kg (44 lb)
Fossil Remains: Complete subadult skull and majority of skeleton, feathers.
Anatomical Characteristics: Skull subrectangular. Arms rather short. Legs fairly long, though the femur
is rather short. Simple feathers, up to 1 cm long, cover the body and neck, full extent uncertain. Long
pennaceous feathers (over 30 cm) sprout from the lower arm and along the second finger over most of
its length, considerably smaller coverts overlap the large primary feathers. Wings asymmetrical in
overall shape, appear to lack alular feathers. More pennaceous feathers lined the entire length of the
tail, growing longer toward the end (at least 15 cm). Legs apparently devoid of feathers, though this
could be an artifact of preservation. Color banding present on wing coverts.
Habits: A terrestrial predator of small game.
Notes: Despite similar proportions, this does not appear to be a specimen of Tianyuraptor. Exact
relationship to other Yixian dromaeosaurids uncertain, may form a short-armed clade with Tianyuraptor
within microraptorinae, or it may be closer to more advanced dromaeosaurs.
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Skull modified after Sinkkonen

Tianyuraptor body modified after Anton
ZP
Boreonykus certekorum
Distribution: Alberta; Wapiti.
Habitat: Riverine forest with warm humid summers and cool winters.
Notes: Classification not certain, may or may not be a dromaeosaur. Shared its habitat with
Pachyrhinosaurus.
Dromaeosaurines
Yurgovuchia doellingi
2.5 m (8 ft) TL, 25 kg (55 lb)
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Distribution: Utah; upper Cedar Mountain.
Habitat: Short wet season, otherwise semiarid with floodplain prairies and open woodlands, and
riverine forest.
Notes: Shared its habitat with a variety of iguanodonts, nodosaurs, basal therizinosaurs, and basal
hadrosaurs. Main enemy Utahraptor.
Dakotaraptor steini
5.5 m (18 ft) TL, 200 kg (450 lb)
Fossil Remains: Minority of skeletons.
Anatomical Characteristics: Proportions more gracile than other giant dromaeosaurs. Teeth heavily
serrated. Sickle claw large. Quill knobs present on ulna, indicative of relatively large wing feathers.
Distribution: South Dakota; Hell Creek.
Habits: Predator of small and medium sized game, such as ornithomimids, Thescelosaurus,
Pachycephalosaurus, and the young of large ceratopsians and hadrosaurs. Better suited to running than
other large dromaeosaurs, better able to pursue prey over longer distances. Main competition was
likely juvenile Tyrannosaurus.
Notes: Robust and gracile morphs appear to be present in adults of this species, probably indicative of
sexual dimorphism. Exact classification uncertain. Shared its habitat with the considerably smaller
Acheroraptor, among many other dinosaurs. A purported furcula assigned to this species more likely
belongs to a softshell turtle.
Velociraptorines
Linheraptor (or Tsaagan) exquisitus
2 m (7 ft) TL, 15 kg (30 lb)
Anatomical Characteristics: Very similar to Velociraptor, slightly more robust overall.
Habitat: Desert with dunes and oases.
Habits: Generalist predator.
Notes: Apparently more closely related to Tsaagan than either is to Velociraptor. Shared its habitat
with at least one of the raptors mentioned above, as well as Protoceratops, oviraptorids, and
Saurornithoides.
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Headden
ZP
Pyroraptor olympius
1.3 m (4 ft) TL, 5 kg (10 lb)
Age: Late Cretaceous, Late Campanian to Early Maastrichtian.
Distribution: France; Grs Reptiles.
Notes: Shared its habitat with sauropods.
Acheroraptor temertyorum
2.5 m (8 ft) TL, 25 kg (55 lb)
Anatomical Characteristics: Snout more robust than relatives.
Distribution: Montana; Hell Creek.
Habitat: Well-watered forests and floodplains.
Notes: First velociraptorine known from North America.
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Saurornitholestes sullivani
Distribution: New Mexico; Kirtland.
Habitat: Seasonal woodland with several large rivers.
Unenlaginines
Rahonavis ostromi
See Page 138
Notes: Considered a primitive avian by some researchers.
Troodonts
Geminiraptor suarezarum
2 m (7 ft) TL, 20 kg (40 lb)
Anatomical Characteristics: Snout not slender. Teeth robust.
Habitat: Short wet season, otherwise semiarid with floodplain prairies, open woodlands, and riverine
forests.
Notes: The oldest confirmed troodontid from North America. Shared its habitat with Eolambia,
Hippodraco, and Iguanacolossus.
Byronosaurus jafferi
See Page 141
Notes: Hatchling skulls, originally assigned to Velociraptor, have been found oviraptorid nests. How this
would come to be is uncertain, most likely the oviraptorid preyed upon the hatchling troodontids and
fed them to its own young. It has been suggested, however, that Byronosaurus may have been a nest
parasite, in which case the newly hatched troodontids may have preyed on their sibling oviraptorids.
Xixiasaurus henanensis
Anatomical Characteristics: Snout long and shallow.
Habits: Preyed upon small game, possibly fished.
Notes: Closely related to Byronosaurus. Together they probably represent a unique East Asian radiation
of troodontids.
Linhevenator (or Saurornithoides?) tani
2 m (7.5 ft) TL, 20 kg (40 lb)
Anatomical Characteristics: Arms short and robust. Sickle claw large.
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44

Distribution: Northern China; Bayan Mandahu.
Habits: Possibly hunted larger game than most troodontids.
Notes: May be congeneric with Saurornithoides.
Talos sampsoni
2 m (7 ft) TL, 15 kg (30 lb)
Notes: Potential prey included Kritosaurus and Hagryphus. Main enemy Teratophoneus.
Gobivenator mongoliensis
1.6 m (5 ft) TL, 3 kg (7 lb)
Anatomical Characteristics: Snout narrow, otherwise standard for large troodontid.
Age: Late Cretaceous, late Campanian.
Distribution: Mongolia; Djadochta.
Habits: Small game hunter, possible took plant matter on occasion.
Notes: The most complete large troodontid known. Potential prey included alvarezsaurs. Shared its
habitat with Saurornithoides and velociraptorine dromaeosaurs.
Tsuihiji et al.
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ZP
Oviraptorosaurs
Ningyuansaurus wangi
1 m (3 ft) TL, 3 kg (7 lb)
Fossil Remains: Partial skull and majority of skeleton, feathers.
Anatomical Characteristics: Skull more elongated than other oviraptorosaurs, also with more teeth and
a straighter lower jaw than usual. Body short and deep, pelvis not large. Arms and hands short, outer
finger reduced. Legs very long, hallux semireversed. Tail long by oviraptorosaur standards. May have
had a feather fan at the end of its tail.
Distribution: Northeast China; upper Yixian.
Habits: Known to have eaten seeds, so likely omnivorous. Main defense high speed.
Notes: The most basal known oviraptorosaur, as favored by the long tail and relatively high number of
teeth which are oriented like those of protarchaeopterygids. However, unlike protarchaeopterygids, it
has long legs, short arms, and a long skull which are more similar to caudipterygids. Alternatively, it
could be an indeterminate basal deinonychosaur.
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Studziski
ZP
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Omnivoropterygids
See Page 144
Notes: Most authorities consider omnivoropterygids to be birds distantly related to oviraptorosaurs.
Protarchaeopterygids
Epidexipterygids
Notes: Despite similarities in skull construction, these are almost certainly scansoriopterygids.
Caudipterygids
Avimimids
Caenagnathids
Gigantoraptor erlianensis
See Page 152
Notes: Probably a basal caenagnathid rather than an oviraptorid.
Epichirostenotes curreii
See Page 151
Notes: New genus and species for Chirostenotes? unnamed species.
Leptorhynchos gaddisi
1.5 m (5 ft) TL, 15 kg (30 lb)
Fossil Remains: Partial skull and minority of skeletons.
Anatomical Characteristics: Lower jaw short and broad. Tip of beak wide and upturned.
Distribution: Texas; Aguja.
Habits: Upturned beak may be an adaptation for rooting in soil for plant and animal matter.
Notes: Shared its habitat with at least one other caenagnathid species and a variety of other dinosaurs.
Oddly shaped beak and presence of close relatives in the same habitat suggests niche partitioning via
differences in feeding strategies within caenagnathids. The name Leptorhynchos is currently occupied
by a plant, and will need to be changed in the future.
Anzu wyliei
See Page 151
3.5 m (11 ft) TL, 200 kg (440 lb)
Notes: New genus and species for Caenagnathus? unnamed species.
Oviraptorids
Ajancingenia yanshini
2 m (7 ft) TL, 40 kg (85 lb)
Fossil Remains: Majority of skulls and skeletons.
Anatomical Characteristics: Beak short and notably hooked. Hand short.
Distribution: Mongolia; Baruungoyot.
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Habitat: Semidesert with dunes and oases.

Notes: New genus for Ingenia yanshini. Remains placed in this genus from other formations are likely
the growth stages of related forms. Considerable controversy surrounds the publication and use of this
new name; it may not withstand the test of time.
Headden
Machairasaurus leptonychus
Anatomical Characteristics: Hand not large, claws relatively long and slender.
Distribution: Northern China; Bayan Mandahu.
Habits: Long claws suitable for pulling down branches.
Notes: Main enemies Velociraptor and Linhevenator.
Luoyanggia liudianensis
1.5 m (5 ft) TL, 15 kg (30 lb)
Fossil Remains: Minority skull and skeleton.
Anatomical Characteristics: Beak of lower jaw pointed.
Age: Early Late Cretaceous.
Distribution: Eastern China; Mangchuan.
Notes: Probably a rather primitive oviraptorid.
Yulong mini
Fossil Remains: Complete skull and majority of skeletons, embryos to subadults; eggs.
Anatomical Characteristics: Much smaller than other derived oviraptorids. Snout not as high as in
other oviraptorids and no crest present, but these features could be age related. Upper beak pointed
and well developed.
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Distribution: Central China; Qiupa.

Habits: Gathered in groups to nest like other oviraptorids.
Notes: Possibly the smallest derived oviraptorid. Proportions of legs similar at all ages, suggesting no
change in feeding strategy upon maturity. Shared its habitat with Qiupalong. Main enemy
Luanchuanraptor.
L et al.
Wulatelong gobiensis
1.5 m (5 ft) TL, 20 kg (45 lb)
Anatomical Characteristics: Lower jaw deep. Hands and claws slender.
Notes: Possibly the most basal known oviraptorid.
Headden
Banji long
Fossil Remains: Complete skull.
Anatomical Characteristics: Relatively short crest bears horizontal striations. Nostrils elongated and
follow curvature of crest. Small by oviraptorid standards.
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Distribution: Southern China; Nanxiong.

Notes: Purpose of striations unclear, may have acted as an attachment point for a keratin extension.
Xu
ZP
Ganzhousaurus nankangensis
Anatomical Characteristics: Lower jaw shallow.
Notes: Appears to have had a mix of primitive and advanced features. Shared its habitat with several
other oviraptorids and Gannansaurus.
Jiangxisaurus ganzhouensis
1.5 m (5 ft) TL, 20 kg (45 lb)
Fossil Remains: Partial skull and majority of skeleton.
Anatomical Characteristics: Lower jaw not as downturned as relatives. Thumb robust.
Notes: Shared its habitat with several other oviraptorids.
Nankangia jianxiensis
2 m (7 ft) TL, 40 kg (85 lb)
Anatomical Characteristics: Lower jaw deep and U-shaped.
Habits: Probably more herbivorous than other oviraptorids.
Notes: Appears to be a relatively basal oviraptorid. Shared its habitat with several other oviraptorids.
Haunansaurus (or Citipati) ganzhouensis
2.5 m (8 ft) TL, 75 kg (160 lb)
Anatomical Characteristics: Low subrectangular crest tilted forward. Upper beak not strongly hooked
and projects further than lower beak. Hand fairly short, claws moderately hooked.
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51
Age: Late Cretaceous, Late Campanian or Early Maastrichtian.

Distribution: Southeast China; Nanxiong.
Habits: Possibly more herbivorous than relatives.
Notes: Despite coexisting with at least four other oviraptorids, its closest relative appears to be Citipati.
Jesse
ZP
Therizinosauroids
Notes: Fragmentary remains suggest presence in Latest Cretaceous North America and Africa.
Baso-Therizinosaurs
Martharaptor greenriverensis
4 m (12 ft) TL, 100 kg (220 lb)
forests.
Notes: Likely a close relative or descendant of Falcarius.
Jianchangosaurus yixianensis
Fossil Remains: Complete skull and majority of skeleton with feathers, juvenile.
Anatomical Characteristics: Skull somewhat robust with a well-developed upper beak and a tooth
arrangement like that of ornithischians. Hand not as elongated as Falcarius and lunate carpal not as well
developed. Tapering band-like feathers at least over shoulders. It is not certain that the feet belong to
this specimen.
Habits: Likely a mid-level browser.
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Notes: More derived than Falcarius and larger than Beipiaosaurus when fully grown, it shared its
habitat with the latter species as well as a wide variety of other dinosaurs. Relatively short neck and
some other features may be juvenile traits. Main enemy Yutyrannus.
Pu et al.
ZP
Alxasaurids
Therizinosaurids
Nothronychus mckinleyi (=grafmani?)
See Page 158
Anatomical Characteristics: Well-developed salt glands in the nasal region.
Habits: Prominent salt glands may hint at a diet composed mostly of saltmarsh plants.
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Sauropodomorphs
Baso-Sauropodomorphs
Pampadromaeus barberenai
1.5 m (5 ft) TL, 10 kg (25 lb)
Anatomical Characteristics: Skull fairly large, front teeth heavily serrated. Arm short. Lower leg long.
Distribution: Southern Brazil; Santa Maria.
Habits: Possibly more omnivorous than other baso-sauropods. Long lower legs indicate that it was a
good runner.
Notes: Possibly the most primitive sauropodomorph known. Prey of Staurikosaurus.
Cabreira et al.
ZP
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Prosauropods
Notes: Remains from South Africa suggest some prosauropods were capable of reaching or exceeding
the size of the largest theropods, possibly making them the largest bipedal animals of all time.
Chromogisaurus novasi
2 m (6.5 ft) TL, 25 kg (50 lb)
Habitat: Seasonally well-watered forests, including dense stands of conifers.
Notes: Main enemies were herrerasaurs and large thecodonts.
Chuxiongosaurus lufengensis
Fossil Remains: Complete skull.
Anatomical Characteristics: Snout subtriangular.
Notes: Appears to be the most basal known prosauropod from the Lufeng.
L et al.
Sarahsaurus aurifrontanalis
4.3 m (14 ft) TL, 200 kg (450 lb)
Fossil Remains: Complete skull and majority of skeletons.
Anatomical Characteristics: Snout subtriangular, upper jaw slightly longer than lower. Hands unusually
large and robust.
Age: Early Jurassic, Sinemurian or Pliensbachian.
Distribution: Arizona; Kayenta.
Habitat: Partially arid.
Habits: Large hands possibly adapted for digging up roots.
Notes: Among the most primitive prosauropods known from North America.
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Colbert & Rowe
ZP
Seitaad ruessi
3 m (10 ft) TL, 70 kg (150 lb)
Age: Early Jurassic, Pliensbachian.
Distribution: Utah; upper Glen Canyon.
Notes: A basal prosauropod.
Sertich & Loewen
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56
Efraasia (or Plateosaurus) minor

See Page 166
Notes: Split from Plateosaurus (=Sellosaurus) gracilis. Many mostly complete specimens are awaiting
further study.
Glacialisaurus hammeri
Age: Early Jurassic, Sinemurian or Pliensbachian.
Distribution: Central Antarctica; Hanson.
Habitat: Polar forest with warm, daylight-dominated summers and cold, dark winters.
Notes: May be another species of Plateosaurus, but too little is known to be certain. Main enemy
Cryolophosaurus.
Jaklapallisaurus asymmetrica
Age: Late Triassic, Late Norian or Early Rhaetian.
Distribution: Central India; Upper Maleri.
Leonerasaurus taquetrensis
Anatomical Characteristics: Neck possibly more stiff than in other prosauropods.
Age: Early Jurassic, probably Pliensbachian or Sinemurian.
Distribution: Southern Argentina; Leoneras.
Notes: A highly derived prosauropod.
Nambalia roychowdhurii
Fossil Remains: Partial skeletons.
Age: Late Triassic, Late Norian or Early Rhaetian.
Distribution: India; Upper Maleri.
Notes: Two of the skeletons, a smaller and a larger individual, were found together.
Xixiposaurus suni
Anatomical Characteristics: Snout slants sharply downward.
Age: Early Jurassic, Hettangian and/or Sinemurian.
Distribution: Southwest China; Lower Lufeng.
Notes: A derived prosauropod.
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Unaysaurus (or Plateosaurus) tolentinoi

2.5 m (8 ft) TL, 40 kg (80 lb)
Anatomical Characteristics: Head shallow, subrectangular. Arm relatively short.
Age: Late Triassic, Norian.
Distribution: Southern Brazil; Caturrita.
Habits: Short arm indicates that this species was more bipedal than similar prosauropods.
Notes: Establishes the presence of Plateosaurus-type prosauropods in the Southern Hemisphere. May
include Teyuwasu barbarenai. Shared its habitat with Guaibasaurus.
Elias
Leyesaurus marayensis
2.5 m (8 ft) TL, 40 kg (80 lb)
Fossil Remains: Majority of skull and minority of skeleton.
Anatomical Characteristics: Skull subrectangular.
Age: Early Jurassic.
Distribution: Western Argentina; Quebrada de Barro.
Notes: Appears to be a relative of Massospondylus.
Apaldetti et al.
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Eucnemesaurus entaxonis
6 m (20 ft) TL, 500 kg (1,000 lb)
Anatomical Characteristics: Legs robust.
Age: Late Triassic, Late Carnian or Early Norian.
Distribution: Southeast Africa; Lower Elliot.
Habitat: Arid.
Notes: Shared its habitat with Eucnemosaurus fortis, Melanorosaurus, and Blikanasaurus.
Sefapanosaurus zastronensis
Fossil Remains: Minority of several skeletons.
Age: Late Triassic or Early Jurassic.
Distribution: South Africa; Elliot.
Habitat: Arid.
Habits: More quadrupedal than most prosauropods.
Notes: Shared its habitat with a variety of other prosauropods, early sauropods, heterodontosaurids,
and coelophysoids.
Aardonyx celestae
Fossil Remains: Partial skulls and skeletons, juveniles.
Anatomical Characteristics: Elastic cheeks greatly reduced, yet snout still narrow, nasal openings large.
Limbs relatively robust. Orientation of limbs suggest more quadrupedal than other prosauropods, but
still capable of slow bipedal motion.
Distribution: South Africa; Upper Elliot.
Habitat: Some regions arid.
Habits: Lack of large cheeks widened gape, allowing for bulk feeding similar to sauropods.
Notes: A highly derived prosauropod, already showing adaptations for quadupedalism and bulk feeding.
Yates
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ZP
Arcusaurus pereirabdalorum
Age: Early Jurassic, Hettangian to Sinemurian.
Habitat: Semi-arid.
Notes: Thought to be a derived prosauropod, but too little is known to be certain.
Sauropods
Vulcanodonts
Antetonitrus ingenipes
See Page 171 8 m (25 ft) TL, 1.5 tonnes
Pulanesaura eocollum
Fossil Remains: Minority of skull and several skeletons, juvenile to adult.
Anatomical Characteristics: Neck fairly flexible.
Age: Early Jurassic, Late Hettangian or Sinemurian.
Habitat: Arid.
Habits: Could high-browse with less effort than the prosauropods it cohabitated with.
Eusauropods
Cetiosaurs
Nebulasaurus taito
Age: Middle Jurassic.
Distribution: Southern China; Zhanghe.
Notes: Known only from a braincase. Closest relative appears to be Spinophorosaurus. Shared its
habitat with Yuanmousaurus and Eomamenchisaurus.
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Mamenchisaurids
Notes: Euhelopids are likely basal titanosaurs and therefore not particularly closely related to
mamenchisaurids.
Tonganosaurus hei
Age: Early Jurassic.
Distribution: Central China; Yimen.
Notes: If this is a mamenchisaurid, it is the earliest known.
Huangshanlong anhuiensis
Age: Middle Jurassic.
Distribution: Eastern China; Hongqin.
Notes: Only known from an arm, but most likely belongs with mamenchisaurids.
Xinjiangtitan shanshanesis
Anatomical Characteristics: Hindlimbs relatively short.
Age: Middle Jurassic, Aalenian.
Distribution: Northwestern China; Qigu.
Notes: One of the first gigantic sauropods.
Qijianglong guokr
Anatomical Characteristics: Neck very long.
Distribution: Central China; Suining.
Notes: A relatively basal mamenchisaurid.
Turiasaurs
Zby atlanticus
Age: Late Jurassic, Late Kimmeridgian.
Distribution: Portugal; Lourinh.
Habitat: Large, seasonally dry island with open woodlands.
Notes: Main enemies included Allosaurus, and Torvosaurus.
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Neosauropods
Diplodocoids
Rebbachisaurids
Comahuesaurus windhauseni
Age: Early Cretaceous, Aptian or Albian.
Distribution: Western Argentina; Lohan Cura.
Notes: Previously thought to be a species of Limaysaurus, appears to be a more basal rebbachisaurid.
Demandasaurus darwini
Age: Early Cretaceous, Late Barremian or Early Aptian.
Distribution: Spain; Castrillo la Reina.
Notes: Closest relative appears to be Nigerasaurus.
Tataouinea hannibalis
Anatomical Characteristics: Skeleton shows extensive pneumaticity.
Age: Early Cretaceous, Early Albian.
Distribution: Tunisia; Ain el Guettar.
Habitat: Tropical floodplain surrounded by arid regions.
Notes: Demonstrates that sauropods could, at least occasionally, exhibit levels of skeletal pneumaticity
previously known only amongst birds, theropods, and pterosaurs. Shared its habitat with a variety of
large theropods.
Dicraeosaurids
Diplodocids
Diplodocines
Kaatedocus siberi
Fossil Remains: Complete skull and minority of skeleton.
Anatomical Characteristics: Skull shallow.
Age: Late Jurassic, Kimmeridgian.
Distribution: Wyoming; lower Morrison.
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62
Habitat: Short wet season, otherwise semiarid with open floodplain prairies and riverine forests.
Habits: Primarily a high-level browser.
Notes: Originally placed in Barosaurus lentus.
Leinkupal laticauda
Anatomical Characteristics: Broad tail vertebrae.
Age: Early Cretaceous, Late Berriasian.
Distribution: Southern Argentina; Bajada Colorada.
Notes: The youngest diplodocine diplodocid and the first from South America.
Apatosaurines
Galeamopus hayi
See Page 190
Notes: New genus for Diplodocus hayi. Not as derived as true Diplodocus species.
Brontosaurus excelsus
See Page 192
Notes: Resurrected genus for Apatosaurus excelsus.
Brontosaurus parvus
See Page 192
Notes: Resurrected genus for Apatosaurus parvus.
Brontosaurus yahnahpin
Fossil Remains: Majority of skeleton.
Age: Late Jurassic, Kimmeridgian.
Distribution: Wyoming; lower Morrison.
Habitat: Short wet season, otherwise semiarid with open floodplain prairies and riverine forests.
Notes: Formerly referred to Apatosaurus and Eobrontosaurus.
Macronarians
Macronarian Miscellanea
Haestasaurus becklesii
Fossil Remains: Minority of skeleton, casts of skin.
Anatomical Characteristics: Non-overlapping hexagonal scales, up to 2.5 cm wide, cover upper arm and
decrease in size toward the elbow.
Age: Early Cretaceous, Valanginian.
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63
Notes: Previously placed in Pelorosaurus, Morosaurus, and Cetiosaurus. Shared its habitat with a
spinosaurid, Becklespinax, Hylaeosaurus, Barilium, Hypselospinus, and at least one other macronarian
sauropod.
Camarasaurids
Cathetosaurus lewisi
See Page 199
Notes: New genus for Camarasaurus lewisi.
Brontomerus mcintoshi
Fossil Remains: Minority of skeletons, juvenile and adult.
Anatomical Characteristics: Hips unusually large, probably to support large leg muscles.
Age: Early Cretaceous, Late Aptian or early Albain.
Distribution: Utah; lower Cedar Mountain.
riverine forest.
Habits: Defenses included kicking with hind legs.
Notes: Relationship to other sauropods unclear. Juvenile and adult found in association.
Titanosauriformes
Titanosauriform Miscellanea
Liubangosaurus hei
Distribution: Southern China; Napai.
Notes: May be congeneric with Fusuisaurus, but too little is known to be certain.
Brachiosaurids
Astrophocaudia slaughteri
Age: Early Cretaceous, Albian.
Distribution: Texas, Oklahoma; Glen Rose, Antlers.
Habitat: Floodplain with coastal swamps and marshes.
Notes: Portions of remains originally placed in Pleurocoelus and/or Paluxysaurus.
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64
Padillasaurus leivaensis
Distribution: Colombia; Paja.
Habitat: Coastal forest.
Notes: The first definitive brachiosaurid from South America. Indicates that the group persisted into
the Cretaceous in both North and South America.
Euhelopids
Notes: These are almost certainly more closely related to titanosaurs than they are to mamenchisaurs.
Gannansaurus sinensis
Fossil Remains: Vertebrae.
Notes: Shows that euhelopid sauropods survived to the end of the Cretaceous. Shared its habitat with
a variety of oviraptorids.
Titanosaurids
Baso-Titanosaurs
Agustinia ligabuei
See Page 205
Notes: Some researchers have concluded that Agustinias bizarre armor is actually just bone that
became warped and eroded during the fossilization process.
Tambatitanis amicitiae
Distribution: Japan; Sasayama.
Notes: Probably part of unique East Asian radiation of titanosaurs.
Yunmenglong ruyangensis
Anatomical Characteristics: Neck relatively long.
Age: Late Early Cretaceous, Aptian or Albian.
Distribution: Central China; Haoling.
Notes: Closest relative appears to be Qiaowanlong.
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65
Rukwatitan bisepultus
Age: Middle Cretaceous.
Distribution: Tanzania; Galula.
Notes: Suggests that unique titanosaur lineages occurred in northern and southern Africa during this
part of the Cretaceous.
Lithostrotians
Atacamatitian chilensis
Age: Late Cretaceous, probably Campanian or Maastrichtian.
Distribution: Chile; Tolar.
Notes: If adult, this was a relatively small titanosaur.
Atsinganosaurus velauciensis
Distribution: France; Grs Reptiles.
Notes: Shared its habitat with other sauropods and Pyroraptor.
Drusilasaura deseadensis
Age: Late Cretaceous, Cenomanian or Turonian.
Distribution: Southern Argentina; Bajo Barreal.
Notes: Shared its habitat with larger Elaltitan.
Elaltitan lilloi
30+ m (100+ ft) TL, 80+ tonnes
Age: Late Cretaceous, Late Cenomanian or Early Turonian.
Distribution: Southern Argentina; Bajo Barreal.
Notes: Among the largest of sauropods.
Fukuititan nipponensis
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66

Distribution: Japan; Kitadani.
Notes: Shared its habitat with Fukuiraptor and Fukuisaurus.
Narambuenatitan palomoi
Fossil Remains: Partial skull and skeleton, large juvenile.
Anatomical Characteristics: Snout not short.
Distribution: Northern Argentina; Anacleto.
Notes: Main enemies Abelisaurus and Aerosteon.
Panamericansaurus schroederi
Distribution: Central Argentina; Allen.
Habitat: Semiarid coastline.
Notes: Closely related to Aeolosaurus.
Petrobrasaurus puestohernandezi
Anatomical Characteristics:
Distribution: Southern Argentina; Plottier.
Rugocaudia cooneyi
Distribution: Montana; Cloverly.
riverine forests.
Notes: Suggests that sauropods may have persisted in North America throughout the Cretaceous.
Some researches consider this to be a dubious genus. Main enemy Acrocanthosaurus.
Tapuiasaurus macedoi
Anatomical Characteristics: Head long, shallow, bony nostrils strongly retracted to above orbits but
fleshy nostrils probably still near front of snout, which is broad, rounded, lower jaws short; pencilshaped teeth limited to front of jaws, head flexed downward relative to neck. Skeleton robust.
Distribution: Southeast Brazil; Quiric.
Habits: Probably capable of browsing at most levels.
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Notes: Grooves on the front of the jaws and retracted nostrils may be indicative of a well-developed
external cropping beak on both the upper and lower jaw. If present, such structures may have been
widespread amongst titanosaurs. The earliest known advanced titanosaur.
Sanches
Yongjinglong datangi
Anatomical Characteristics: Scapula very large.
Age: Late Early Cretaceous.
Distribution: Northern China; Hekou Group.
Notes: Shared its habitat Daxiatitan and Huanghetitan.
Overosaurus paradasorum
Distribution: Central Argentina; Anacleto.
Notes: Shared its habitat with a variety of other sauropods. Main enemies abelisaurids and Aerosteon.
Brasilotitan nemophagus
Anatomical Characteristics: Lower jaw strongly squared-off, neck relatively short.
Age: Late Cretaceous, probably Campanian or Maastrichtian.
Distribution: Southern Brazil; Adamantina.
Habits: Probably a low-level grazer.
Notes: Related to Bonitasaura, but does not appear to have possessed a beak.
Normanniasaurus genceyi
Distribution: France; Poudingue Ferrugineux.
Notes: A relatively basal titanosaur.
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Dreadnoughtus schrani
26+ m (85+ ft) TL, 50+ tonnes
Fossil Remains: Minority of skull and majority of skeletons.
Anatomical Characteristics: Skeleton well pneumatisized, shorter limbs than most other gigantic
titanosaurs, neck proportionally long.
Age: Late Cretaceous, late Campanian or early Maastrichtian.
Distribution: Argentina; Cerro Fortaleza.
Habitat: Large rivers present.
Habits: Likely a generalist browser, able to feed on both groundcover and in the treetops.
Notes: The most complete gigantic titanosaurs discovered thus far. The largest known specimen is
skeletally immature, but likely still an adult capable of breeding. Specimens show signs of scavenging by
small theropods.
Lacovara et al.
Quetecsaurus rusconii
Distribution: Western Argentina; Cerro Lisandro.
Habitat: Swamps present.
Vahiny depereti
Distribution: Madagascar; Maevarano.
Habitat: Seasonally dry floodplain with coastal swamps and marshes.
Notes: Shared its habitat with Rapetosaurus, to which it was not closely related. Main enemy
Majungasaurus.
Saltasaurs
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69
Ornithischians
Baso-Ornithischians
Pisanosaurus mertelli
See Page 214
Notes: This may be a dinosauriforme silesaurid, rather than a true dinosaur.
Laquintasaura venezuelae
1 m (3.5 ft) TL, 1 kg (2 lb)
Anatomical Characteristics: Skull more robust than other baso-ornithischians.
Distribution: Venezuela; La Quinta.
Notes: Likely prey of Tachiraptor.
Barrett et al.
Genasaurs
Thyreophorans
Scelidosaurs
Bienosaurus lufengensis
Notes: Main enemy Sinosaurus triassicus. Shared its habitat with prosauropods.
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70
Eurypods
Stegosaurs
Huayangosaurids
Stegosaurids
Miragaia (or Dacentrurus) longicollum
See Page 223
Notes: Several features of this taxons postcranial anatomy match well with whats known of
Dacentrurus.
Hesperosaurus (or Stegosaurus) mjosi
See Page 224
Notes: Two plate types appear to be present in adults, one with relatively wider plates and the other
with relatively taller plates. As intermediates are unknown, this is likely indicative of sexual dimorphism.
Ankylosaurs
Minmids
Kunbarrasaurus ieversi
See Page 227
Distribution: Northeast Australia, Allaru Mudstone.
Notes: New genus and species for Minmi? unnamed species.
Leahey et al.
Chuanqilong chaoyangensis
Adult size uncertain
Anatomical Characteristics: Tail club probably absent, full extent of armor uncertain.
Distribution: Northeast China; Jiufotang.
Notes: Closely related to Liaoningosaurus, with which it shared its habitat. Like Liaoningosaurus,
possibly not a minmid.
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Polacanthians
Taohelong jinchengensis
Anatomical Characteristics: Irregularly shaped osteoderms on sacral shield.
Age: Early Cretaceous.
Distribution: Central China; Hekou Group.
Notes: Closely related to Polacanthus.
Horshamosaurus rudgwickensis
Distribution: Southeast England; Weald Clay.
Notes: Classification uncertain, may be a primitive ankylosaurid or nodosaurid. Shared its habitat with
Baryonyx and iguanodonts. Formerly considered a species of Polacanthus.
Ankylosaurids
Crichtonpelta benxiensis
Anatomical Characteristics: Prominent squamosal and jugal horns angled backwards. Presence of tail
club unknown.
Age: Late Cretaceous, Cenomanian or Turonian.
Distribution: Northeast China; Sunjiawin.
Notes: Probably includes a skeleton assigned to Crichtonsaurus bohlini, with which it coexisted and was
formerly considered synonymous with.
Zhongyuansaurus luoyangensis
Anatomical Characteristics: Skull shallow, lacked tail club.
Distribution: Eastern China; unknown.
Notes: Possibly the basal most ankylosaurid. If not, then it may be a polacanthian or a basal nodosaur.
Zaraapelta nomadis
Fossil Remains: Majority of skull and minority of skeletons.
Anatomical Characteristics: Double-row of osteoderms on top of skull, conical hornlets on the upper
rim of orbits, four prominent postorbital horns. Beak blunt. Elongated osteoderm runs majority of the
length of lower jaw. Tail club present but shape uncertain.
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Distribution: Mongolia; Baruungoyot.

Habitat: Seasonal woodland.
Notes: Closely related to Tarchia and Saichania, with which it shared its environment. Highly
ornamented cranium possibly a sexual display.
Arbour et al.
Anodontosaurus lambei
See Page 234
Notes: Resurrected genus and species for Euplocephalus unnamed species.
Scolosaurus cutleri
Fossil Remains: Majority of skull and minority of skeletons.
Anatomical Characteristics: Hornlets at back of skull and under eyes are large, three-sided, and
triangular; eye sockets large. Osteoderms have a smooth surface. Limbs robust. Tail club modest-sized
and heart-shaped when viewed from above.
Distribution: Montana; Upper Two Medicine.
Habitat: Seasonally dry upland woodlands.
Notes: Split from Euoplocephalus tutus. Skull of one specimen shows evidence of being stepped on.
Shared its habitat with Edmontonia and a wide variety of other dinosaurs. Probably includes Oohkotokia
horneri.
Dyoplosaurus acutosquameus
Anatomical Characteristics: Tail club very narrow.
Distribution: Alberta; Lower Dinosaur Park.
Habitat: Well-watered, forested floodplain with coastal swamps and marshes, cool winters.
Notes: Split from Euoplocephalus tutus.
Ziapelta sanjuanensis
5.5 m (18 ft) TL, 2.5 tonnes
Anatomical Characteristics: Large postorbital horns. Six large spines on cervical rings.
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Habitat: Moderately watered floodplain woodlands, coastal swamps and marshes.
Notes: Shared its habitat with Nodocephalosaurus. Main enemy Bistahieversor.
Ahshislepelta minor
Anatomical Characteristics: Osteoderms have a knobby texture.
Notes: Smallest ankylosaurid in region. Main enemy Bistahieversor.
Nodosaurids
Propanoplosaurus marylandicus
Fossil Remains: Natural cast of a young juvenile including most of the body and head.
Anatomical Characteristics: Only osteoderms on head well developed.
Age: Early Cretaceous, Late Aptian.
Distribution: Maryland; Patuxent.
Notes: Reduced armor is likely a condition only present in young juveniles.
Tatankacephalus cooneyorum
Fossil Remains: Partial skull and minority of skeleton.
Anatomical Characteristics: Skull domed and with a ridge along back edge.
Age: Early Cretaceous, Late Aptian or Early Albian.
forests.
Notes: Placement uncertain. Shared its habitat with Sauropelta, Tenontosaurus, and Deinonychus.
Dongyangopelta yangyanensis
4.5 m (15 ft) TL, 1.4 tonnes
Anatomical Characteristics: Pelvic shield fused with moderately-sized, smooth, rounded osteoderms;
other osteoderms have a very rough surface indicating they once supported larger attachments.
Age: Early Late Cretaceous, Cenomanian.
Distribution: East China; Chaochuan.
Notes: Shared its habitat with fellow nodosaurid Zhejiangosaurus.
Europelta carbonensis
4.5 m (15 ft) TL, 1.4 tonnes
Fossil Remains: Majority of skull and partial skeletons.
Anatomical Characteristics: Nostrils rather large, moderately sized jugal ridges present. Subrectangular
osteoderms cover neck gradually becoming more oval shaped toward hips. Three moderately sized
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74
shoulder spines, largest toward front. Plate-like osteoderms along sides of hips and tail have a rounded
margin.
Distribution: Spain; Escucha.
Notes: Oldest European nodosaurid, suggests that nodosaurids replaced polacanthian ankylosaurs in
Europe during the latter part of the Cretaceous, just as they did in North America. Shared its habitat
with Proa.
Kirkland et al.
http://scotthartman.deviantart.com/art/Europe-s-shield-429476594
Hartman
ZP
Kirkland et al.
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Heterodontosauriforms
Heterodontosaurids
Pegomastax africana
1.2 m (4 ft) TL, 3.5 kg (7.5 lb)
Anatomical Characteristics: Beak and lower jaw robust.
Age: Early Jurassic, Hettangian or Sinemurian.
Habitat: Arid.
Notes: Shared its habitat with Heterodontosaurus, which it was not particularly closely related to. Prey
of Coelophysis.
Manidens condorensis
0.75 m (2.5 ft) TL, 0.8 kg (1.8 lb)
Anatomical Characteristics: Snout shorter than relatives, short fangs present.
Age: Middle Jurassic, Aalenian or Early Bathonian.
Distribution: Northern Argentina; Caadn Asfalto.
Notes: Main enemies Piatnitzkysaurus, Condorraptor, and Eoabelisaurus. Shared its habitat with many
sauropods.
Pol et al.
Tianyulong confuciusi
See Page 240
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Marginocephalians
Pachycephalosaurids
Acrotholus audeti
1.5 m (5 ft) TL, 10 kg (20 lb)
Fossil Remains: Skull dome.
Anatomical Characteristics: Skull heavily reinforced despite small size.
Age: Late Cretaceous, Late Santonian.
Distribution: Alberta; Milk River.
Notes: The earliest fully domed pachycephalosaurid known from North America.
Amtocephale (or Stegoceras) gobiensis
Fossil Remains: Skull dome.
Age: Late Cretaceous, Turonian or Santonian.
Distribution: Mongolia; Bayan Shireh.
Notes: Main enemy Achillobator. Shared its habitat with Garudimimus.
Stegoceras? novomexicanum
1.5 m (5 ft) TL, 10 kg (20 lb)
Fossil Remains: Minority of skulls.
Distribution: New Mexico; Fruitland, Kirtland.
Habitat: Floodplain woodlands.
Notes: It is not certain that this species belongs in Stegoceras.
Ceratopsians
Chaoyangosaurs
Hualianceratops wucaiwanensis
1 m (3 ft) TL, 6 kg (13 lb)
Anatomical Characteristics: Skull subtriangular and broad, lower jaw very robust.
Distribution: Northwest China; Lower Shishugou.
Habits: Likely bipedal. Fed on tough plant matter.
Notes: Classification as a chaoyangosaur uncertain, but if accurate, suggests that chaoyangosaurs and
psittacosaurids formed a sister group to all other ceratopsians. Main enemies Aorun, Guanlong, and
Zuolong. Also shared its habitat with Limusaurus, Haplocheirus, stegosaurs, basal titanosaurs,
mamenchisaurs, allosauroids, and other basal ceratopsians.
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77
Paxceratopsians
Psittacosaurids
Notes: An adult preserved alongside numerous young juveniles, apparently buried while in a burrow,
may indicate parental care and possibly communal nesting/rearing.
Psittacosaurus unnamed species?
See Page 246
Anatomical Characteristics: Predominately brownish in color with some form of camouflage-patterning
likely present. Stomach, back of legs, and underside of tail lighter than dorsal color. Coloration of chest
somewhere between the dorsal and ventral colors. Horns and facial bosses darker.
Notes: Likely preferred dense regions of forest where its disruptive camouflage and countershading
would be most effective.
Neoceratopsians
Mosaiceratops azumai
1.3 m (4 ft) TL, 20 kg (40 lb)
Anatomical Characteristics: Upper beak moderately hooked, lower beak fairly straight. Lacks
premaxillary teeth. Frill incipient. Arms fairly long.
Age: Late Cretaceous, stage uncertain.
Distribution: Eastern China; Xiaguan.
Habits: Probably semi- or mostly bipedal.
Notes: Exact classification uncertain. Thought to be a transitional form between psittacosaurids and
neoceratopsians, indicating that premaxillary teeth evolved at least twice amongst ceratopsians.
However, some feature suggest that this species may fall within psittacosaurids, or even well within
neoceratopsians close to Bagaceratops.
Zheng, Jin, & Xu
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78
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Aquilops americanus
0.6 m (2 ft) TL, 2 kg (4 lb)
Fossil Remains: Majority of skull, possibly juvenile.
Anatomical Characteristics: Beak of upper jaw strongly hooked with a distinct knob on its anterior
edge, which may have been enlarged with keratin to form a short horn or crest. Beak of lower jaw
relatively straight and pointed. Rear of skull flares outwards giving a highly triangular profile when
viewed from above.
riverine forests.
Habits: Low-level browser. Beak ornamentation likely for display, but could have also assisted in
rooting in soil for food.
Notes: The oldest and most primitive ceratopsian known from North America. Its closest relatives
appear to be from Asia, indicating multiple faunal exchanges between Asia and North America during
the Cretaceous, probably via an intermittent land bridge across the Bering Sea. Main enemy
Deinonychus.
Farke et al.
ZP
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79
Farke et al.
Protoceratopsids
Archaeoceratops yujingziensis
0.5 m (1.7 ft) TL, 2 kg (4 lb)
Anatomical Characteristics: Insufficient information
Distribution: Central China; Xinminbao.
Habitat: Well-watered forest and lakes.
Notes: Potential synonymous with A. oshimai, with which it shared its environment.
Ajkaceratops kozmai
1.1 m (3.5 ft) TL, 10 kg (20 lb)
Distribution: Hungary; Csehbnya.
Notes: The first ceratopsian known form Europe. Shared its habitat with Hungarosaurus.
Gryphoceratops morrisoni
0.5 m (1.7 ft) TL, 2 kg (4 lb)
Age: Late Cretaceous, Late Santonian.
Distribution: Alberta; Milk River.
Notes: The smallest ceratopsian known from North America.
Koreaceratops hwaseongensis
0.9 m (3 ft) TL, 10 kg (20 lb)
Anatomical Characteristics: Vertebral spines over tail form an ascending sail, tallest near the end.
Distribution: South Korea; Tando Beds.
Notes: Purpose of sail-tail unclear, probably not used for swimming, more likely for display.
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80
Unescoceratops koppelhusae
1.3 m (4.3 ft) TL, 20 kg (40 lb)
Anatomical Characteristics: Teeth highly rounded.
Distribution: Alberta; Dinosaur Park.
Habitat: Well-watered forests.
Habits: Purpose of unusual teeth unclear, may have been for cracking nuts or eating gritty foods.
Zhuchengceratops inexpectus
2.5 m (8 ft) TL, 175 kg (380 lb)
Anatomical Characteristics: Lower jaw deep.
Notes: Shared its habitat with Ischioceratops (which it may be a senior synonym of), Sinoceratops,
Tanius, and Tsintaosaurus. Prey of Zhuchengtyrannus.
Ischioceratops zhuchengensis
2.5 m (8 ft) TL, 175 kg (380 lb)
Anatomical Characteristics: Tail not as short as some of its relatives. Ischium exhibits a peculiar bend
and a perforation halfway along its length.
Notes: Potentially synonymous with Zhuchengceratops inexpectus, which is known from the same
locality, but lacks overlapping elements to be certain. Also shared its habitat with Tsintaosaurus, Tanius,
Sinoceratops, and Zhuchengtyrannus.
Protoceratops andrewsi
See Page 254
Notes: Nests containing numerous hatchlings indicate some degree of parental care early in their lives.
Ceratopsids
Centrosaurines
Xenoceratops (or Albertaceratops) foremostensis
Anatomical Characteristics: Modest sized brow horns curve forward, nasal boss low. Frill
subhorizontal, concave, and slightly indented; a pair of flattened spines adorn top with another smaller
pair on sides, small hornlets between.
Age: Late Cretaceous, Middle Campanian.
Distribution: Alberta; Foremost.
Notes: The oldest centrosaurine known from Alberta.
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81
Medusaceratops (or Albertaceratops) lokii

Fossil Remains: Partials skulls.
Anatomical Characteristics: Nasal boss low and narrow, frill subvertical, back rim bore a pair of massive,
sideways-arcing spikes, smaller sideways-arcing spikes rim sides.
Distribution: Alberta; Judith River.
Habitat: Well-watered, forested floodplain with coastal swamps and marshes and drier upland
woodlands.
Habits: Spikes rimming frill may have been used in defense.
Notes: Originally placed in Albertaceratops nesmoi, may be the same genus or species.
Wendiceratops (or Albertaceratops) pinhornensis
Anatomical Characteristics: Nasal horn fairly long, strongly flattened laterally. Frill subhorizontal with
subtriangular hornlets that increase in size toward the top of the frill margin. Largest hornlets near top
strongly curve downward.
Distribution: Alberta; Oldman.
Notes: Shared its habitat with Stegoceras and Albertaceratops. Main enemy Daspletosaurus.
Evans & Ryan

Nasutoceratops (or Albertaceratops) titusi
4.5 m (15 ft) TL, 1.2 tonnes
Anatomical Characteristics: Frill short and nearly horizontal with small hornlets lining rim and at base of
median bar. Brow horns moderately long, point forward, curving inward and up. Nasal region strongly
expanded upward, nasal boss nearly absent.
Habits: Inward curving horns better suited for conflicts within own species than in defense.
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82
Notes: Shared its habitat with Kosmoceratops and Utahceratops, along with a variety of other
dinosaurs. Main enemy Teratophoneus.
Sampson et al.
ZP
Sampson et al.
Spinops (or Centrosaurus) sternbergorum
Anatomical Characteristics: Nasal horn medium-sized, brow horns small. Frill subhorizontal, two large
spikes on back rim, two smaller downward curving spikes in front of these.
Distribution: Alberta; upper Oldman or lower Dinosaur Park.
Habitat: Well-watered, forested floodplain with coastal swamps and marshes.
Notes: May or may not be a distinct genus from Centrosaurus. Main enemy Daspletosaurus.
Coronosaurus (or Centrosaurus) brinkmani
See Page 260
Notes: New genus for Centrosaurus brinkmani, as found to phylogenetically closer to Styracosaurus
than Centrosaurus.
Rubeosaurus (or Centrosaurus) ovatus
See Page 262
Notes: New genus for Styracosaurus ovatus.
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83
Pachyrhinosaurus (or Centrosaurus) perotorum

Fossil Remains: Partials skulls, juvenile and adult.
Anatomical Characteristics: In mature adults, very large nasal boss possesses a raised central area and
nearly connects with smaller bosses over eyes; heavily pitted surface of bosses suggest they were
enlarged with keratin in life. Frill subhorizontal with two sideways arcing blades at the top of the rim,
central region and rim lined with small hornlets. Beak relatively blunt.
Distribution: Alaska; Prince Creek.
Habitat: Polar forest with warm humid summers and cold dark winters.
Notes: Main enemy Nanuqsaurus.
Fiorillo and Tykoski

Sinoceratops zhuchengensis
Anatomical Characteristics: Nasal horn medium-sized, brow horns very small. Frill subvertical, rimmed
with flattened triangular spines that curve forward, bony knobs rim front outer edge.
Notes: The only centrosaurine ceratopsid known from Asia, probably represents an incursion from
North America. Shared its habitat with a variety of hadrosaurs. Prey of Zhuchengtyrannus.
Panzarin
ZP
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84
Ceratopsines
Chasmosaurines
Anchiceratops ornatus
See Page 268
Notes: It is not certain that the holotype skull belongs with the skeleton that it was assigned to.
Torosaurus latus (and/or utahensis)
8.5 m (28 ft) TL, 10 tonnes See page 266
Fossil Remains: Majority of skulls and skeletons, juvenile to adult.
Anatomical Characteristics: Snout shallow, nasal horn small, brow horns long and straight. Frill greatly
elongated with two large openings, rimmed by small hornlets. Skeleton robust.
Distribution: Saskatchewan, Montana, Wyoming, Dakotas; Frenchman, Laramie, Denver, Hell Creek.
Habitat: Well-watered forests to dry woodlands.
Habits: Low-level browser. Frill used mainly for display, brow horns for defense against predators and
in conflicts with own species.
Notes: Resurrected genus and species from Triceratops horridus, where it was previously considered
the fully mature form. Discovery of mislabeled juvenile remains were the cause; however, these
remains are from the more westerly Torosaurus utahensis (shown here), which may actually be a
different genus. In the possible event that T. latus is refound to be a form of Triceratops, T. utahensis
will likely require a new genus name. Classification as a chasmosaurine is not certain.
Hunt & Lehman
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85
Kosmoceratops richardsoni
Anatomical Characteristics: Brow horns short, curve down perpendicular to the skull. Nasal horn short
and thin. Frill is wider than long, slightly concave, with ten short downward curving horns lining the top,
small hornlets along sides. Frill openings small.
Notes: Shortness of frill and small frill openings separate this genus from closely related Chasmosaurus.
Main enemy Teratophoneus. Not present in Albertas Dinosaur Park Formation.
Sampson et al.
ZP
Sampson et al.
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86
Coahuilaceratops (or Chasmosaurus) magnacuerna

6.4 m (21 ft) TL, 4.7 tonnes
Anatomical Characteristics: Brow horns very large, curve forward.
Distribution: Northern Mexico; Cerro del Pueblo.
Notes: Shared its habitat with Hypacrosaurus (=Velafrons) coahuilensis.
Mojoceratops (or Chasmosaurus) perifania
4.3 m (14 ft) TL, 1.5 tonnes
Fossil Remains: Partial skulls and minority of skeleton.
Anatomical Characteristics: Brow horns long and arch sideways. Nasal horn short. Frill subhorizontal,
back rim very broad and forms a shallow U, corners bore large hornlets, small hornlets along rims.
Notes: Split from Chasmosaurus russelli, may still be the same genus or species, perhaps as the fully
mature or male form.
Mercuriceratops (or Chasmosaurus) gemini
Fossil Remains: Minority of skulls.
Anatomical Characteristics: Frill subhorizontal, squamosal region of frill flares out into small wing-like
processes rimmed with small hornlets.
Distribution: Alberta, Montana; Judith River, Dinosaur Park.
Habitat: Well-watered forests and floodplains.
Notes: Unique frill ornaments likely enhanced display of frill.
Utahceratops (or Pentaceratops) gettyi
Fossil Remains: Complete skull and majority of skeletons.
Anatomical Characteristics: Nasal horn short and placed high on snout, brow horns very short and
point to the sides. Frill subhorizontal, broadest at base, large vertical frill openings placed high up,
strongly indented, large hornlets at top decrease in size down sides.
Notes: May be a species of Pentaceratops or Chasmosaurus. Shared its habitat with Kosmoceratops.
Main enemy Teratophoneus.
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87
Sampson et al.
ZP
Sampson et al.
Page
88
Vagaceratops (or Chasmosaurus) irvinensis

See Page 270
Notes: New genus for Chasmosaurus irvinensis.
Judiceratops (or Chasmosaurus) tigris
4.5 m (15 ft) TL, 1.2 tonnes
Anatomical Characteristics: Frill rounded with a thick midline bar, hornlets large at base of frill and
decrease in size toward rear. Brow horns moderately long and splay forward and outward slightly, teardrop shaped in cross section.
Distribution: Montana; Judith River.
Notes: The oldest known chasmosaurine.
Pentaceratops (or Chasmosaurus) aquilonius
Habitat: Well-watered forest and floodplain with coastal marshes and cool winters.
Notes: Validity questionable, probably belongs in a species of Chasmosaurus.
Bravoceratops (or Pentaceratops) polyphemus
Anatomical Characteristics: Snout narrow, frill long and tall with large openings and a fan-shaped
median bar. Brow horns moderately large, nasal horn small.
Distribution: Texas; Javelina.
Habitat: Seasonally dry open woodlands.
Notes: Possibly ancestral to Torosaurus? utahensis, as features are intermediate of advanced
chasmosaurines such as Torosaurus and less advanced forms such as Coahuilaceratops (or
Pentaceratops), the latter of which appears to be close relative. Main enemy Tyrannosaurus.
Wick and Lehman
Page
89
Titanoceratops (or Pentaceratops) ouranos

Anatomical Characteristics: Nasal horn short, brow horns long and curve forward. Frill probably
elongated, tilted upward. Belly broad, and hind legs bowed.
Distribution: New Mexico; upper Fruitland or lower Kirtland.
Notes: Split from Chasmosaurus (Pentaceratops) sternbergi. Other remains placed in the former genus
could belong here, but this is not certain.
Longrich
Regaliceratops peterhewsi
Fossil Remains: Majority of skull.
Anatomical Characteristics: Beak not strongly hooked. Nasal horn straight, fairly slender, and relatively
short. Brow horns very short and slender, angled forward. Frill relatively short, subhorizontal, with a
rounded rear margin. Frill openings small. Flattened subtriangular hornlets rim the edge of the frill,
growing largest at the top. A single more rounded hornlet occupies the center of the frills margin,
placed slightly forward and overlapping the others.
Age: Late Cretaceous, Middle Maastrichtian.
Distribution: Alberta; St. Mary River.
Habits: Small fairly delicate horns better suited for display and combat within species than in defense
against predators.
Notes: One of the most distinct chasmosaurines yet discovered. Shared its habitat with
Montanoceratops and Pachyrhinosaurus.
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90
Brown & Henderson
ZP
Brown & Henderson
Ornithopods
Hypsilophodonts
Notes: This is likely a waste-basket group containing a number of superficially similar forms.
Kulindadromeus zabaikalicus
1.5 m (4.5 ft) TL,
Fossil Remains: Majority of skulls and skeletons, juvenile to adult, filament and scale impressions.
Anatomical Characteristics: Snout subtriangular, jaws tipped with a short blunt beak. Eyes fairly large.
Remainder of body fairly standard for small hypsilophodontid-type ornithischian. Short (up to 3 cm)
filaments cover body, neck, upper arm, leg down to shin, and at least most of head; these fall into three
categories: standard hair-like filaments cover majority; upper arm and thigh covered in groups of
hexagonal non-overlapping scale-like structures each of which bears 6-7 filaments projecting downward
from it (up to 1.5 cm); leg below knee down to mid shin covered in bundles of ribbon-like structures (up
to 2 cm) each which is composed of many much smaller filaments. Small scales are present on exposed
portions of limbs and tail; overlapping hexagonal scales on lower shins and non-overlapping rounded
scales on the remainder of the limbs. At least five overlapping rows of rectangular scales cover tail,
largest at the top and decreasing with size ventrally, each scale possesses a forward facing prong on its
leading edge, scales at end of tail are smaller, rounded, and non-overlapping
Age: Mid to Late Jurassic, late Bajocian to early Tithonian.
Distribution: East Russia; Ukureyskaya.
Habitat:
Habits: Low-level browsing herbivore, probably eating small animals on occasion. Well suited for
running and possibly burrowing. Specimens found in association suggest some sort of herding behavior.
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91
Notes: Filamental covering used for insulation, longer ribbon-like structures may have been for display.
Overlapping rows of scales on tail may have helped keep the tail keep rigid and/or acted as low grade
armor for protection against small predators. If the filaments are homologous to the feathers of
theropods, it is likely that the origin of filamental coverings can be extended to the base of dinosauria,
and possibly even further to the common ancestor of dinosaurs and pterosaurs. Kulindadromeus likely
falls outside of hypsilophodonts, but is placed here until the taxonomy of the group is sorted out.
Kulindapteryx, Daurosaurus, and the alleged ornithomimosaur Lepidocheirosaurus are invalid
synonyms of this species.
Godefroit et al.
ZP
Hexinlusaurus multidens
See Page 274
Notes: New genus for Agilisaurus? unnamed species. Slightly more derived than Agilisaurus.
Gideonmantellia amosanjuanae
Fossil Remains: Partial skeleton, juvenile.
Notes: Closest relatives appear to be in North America. Prey of Camarillasaurus.
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92
Haya griva
2 m (6.5 ft) TL, 20 kg (45 lb)
Fossil Remains: Complete skull and majority of skeletons, juvenile and adult. Gastroliths.
Anatomical Characteristics: Skull subtriangular with pointed beak. Legs robust.
Distribution: Mongolia; Javkhlant.
Habitat: Swallowed gastroliths to help digest plant matter. Probably not as agile as similar species.
Notes: Shared its habitat with Yamaceratops.
Buchholz
ZP
Koreanosaurus boseongensis
2.5 m (8 ft) TL, 45 kg (100 lb)
Anatomical Characteristics: Robustly constructed. Arms fairly long. Legs relatively short.
Age: Late Cretaceous, Santonian or Campanian.
Distribution: South Korea; Seonso Conglomerate.
Habits: Like its closest relatives, likely dug burrows using robust arms.
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93
Yueosaurus tiantaiensis
2 m (6.5 ft) TL, 20 kg (45 lb)
Anatomical Characteristics: Standard for hypsilophodonts, but skull is unknown.
Age: Early Cretaceous, probably Albian.
Distribution: Eastern China; Liangtoutang.
Notes: Among the most basal known ornithopods from Asia.
Zheng et al.
Albertadromeus syntarsus
2 m (6.5 ft) TL, 20 kg (45 lb)
Anatomical Characteristics: Lower leg long and slender.
Distribution: Alberta; Oldman.
Habits: Well adapted for running.
Notes: Closely related to Orodromeus and Zephyrosaurus.
Thescelosaurus assiniboiensis
Anatomical Characteristics: Hips relatively small.
Distribution: Saskatchewan; Frenchman.
Notes: Somewhat intermediate between Parksosaurus and Thescelosaurus neglectus; and considerably
smaller than both.
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94
Iguanodontians
Tenontosaurs
Rhabdodonts
Mochlodon (or Rhabdodon) vorosi
Distribution: Austria; Gosau.
Habitat: Forested island.
Notes: Likely includes some remains assigned M. suessi, which is invalid.
Elasmaria
Notes: A unique radiation of Patagonian, Antarctic, and Australian ornithopods more derived than the
hypsilophodonts they were formerly categorized with. May be especially close to dryosaurs.
Leaellynasaura amicagraphica
See Page 276
Anatomical Characteristics: Eyes large, tail exceptionally long and flexible.
Notes: If a dense covering of fibers were present, flexible tail could have been used for insulation while
at rest; potential other uses include a balancing aid or means of visual communication to other
members of the group.
Buchholz
Fulgurotherium australe
See Page 276
Quantassaurus intrepidus
See Page 276
Notohypsilophodon comodorensis
See Page 277
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95
Anabisetia saldiviai
See Page 277
Macrogryphosaurus gondwanicus
See Page 277
Gasparinisaura cincosaltensis
See Page 278
Talenkauen santacrucensis
See Page 278
Trinisaura santamartaensis
Distribution: Western Antarctica; Snow Hill Island.
Habitat: Polar forests with warm, day-light dominated summers and cold, dark winters.
Notes: The first ornithopod described from Antarctica. Shared its habitat with Antarctopelta.
Morrosaurus antarcticus
Distribution: Western Antarctica; Snow Hill Island.
Habitat: Polar forests with warm, day-light dominated summers and cold, dark winters.
Habits: Well adapted to running.
Notes: Larger than Trinisaura, with which it shared its habitat.
Dryosaurs
Eousdryosaurus nanohallucis
1.6 m (5 ft) TL, 13 kg (30 lb)
Anatomical Characteristics: Hallux reduced.
Age: Late Jurassic, late Kimmeridgian.
Distribution: Portugal, Alcobaca
Habits: Well adapted for running.
Notes: A basal dryosaurid.
Dysalotosaurus (or Dryosaurus) lettowvorbecki
See Page 283
Notes: Resurrected genus for Dryosaurus lettowvorbecki.
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96
Ankylopollexia
Camptosaurs
Uteodon (or Camptosaurus) aphanoecetes
See Page 285
Notes: New genus for Camptosaurus aphanoecetes.
Ankylopollexia Miscellanea
Kukufeldia tilgatensis
Distribution: Southeast England; Weald Clay.
Notes: Originally placed in Iguanodon anglicus. Shared its habitat with Baryonyx.
Barilium dawsoni
See Page 286
Notes: New genus for Iguanodon dawsonii.
Bolong yixianensis
Anatomical Characteristics: Jaws relatively long, teeth robust.
Age: Early Cretaceous, Barremian or Aptian.
Notes: Main enemy Yutyrannus. The most primitive iguanodont known from Asia.
paleofile.com
Cumnoria prestwichii
See Page 285
Notes: New genus for Camptosaurus prestwichii. Found to be a more derived iguanodont than
Camptosaurus, though not all researchers agree.
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97
Sellacoxa (or Barilium) pauli

Age: Early Cretaceous, Early Valanginian.
Notes: Originally placed in Barilium dawsonii. May still be a species of Barilium.
Ratchasimasaurus suranareae
Distribution: Thailand; Khok Kruat.
Notes: Shows a combination of primitive and advanced characteristics. Potential synonymous with
Siamodon.
Iguanodontoids
Iguanodonts
Delapparentia turolensis
Notes: Main enemy Camarillasaurus.
Dollodon bampingi
See Page 289
Notes: Correct spelling of Dollodon bambingi.
Hippodraco scuttodens
4.5 m (15 ft) TL, 400 kg (900 lb)
Anatomical Characteristics: Skull subrectangular.
Age: Early Cretaceous, Late Barremian or Early Aptian.
Habitat: Short wet season, otherwise with floodplain prairies, open woodlands, and riverine forests.
Habits: Middle-level browser.
Notes: Shared its habitat with Iguanacolossus, Eolambia, and Animantarx.
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98
McDonald et al.
Hypselospinus fittoni
See Page 288
Notes: New genus for Iguanodon fittoni. Likely includes Huxleysaurus hollingtoniensis.
Iguanacolossus fortis
Habitat: Short wet season, otherwise with floodplain prairies, open woodlands, and riverine forests.
Habits: Middle- and high-level browser.
Notes: The largest iguanodont known from North America. Shared its habitat with Hippodraco,
Eolambia, and Animantarx.
McDonald et al.
Proa valdearinnoensis
Anatomical Characteristics: Lower jaw fairly deep, lower beak pointed.
Distribution: Spain; Escucha.
Habits: Beak well suited for cracking hard objects, such as nuts.
Notes: The latest known Early Cretaceous iguanodontoid from Europe.
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99
Darwinsaurus evolutionis
See Page 288
Notes: New genus and species name for Unnamed genus and species. May include remains assigned
to Mantellodon carpenteri.
Siamodon nimngami
Notes: Potentially synonymous with Ratchasimasaurus.
Iguanodon galvensis
Fossil Remains: Majority of skulls and skeletons, hatchling to adult.
Anatomical Characteristics: In hatchlings: snouts of young very short and sloping, thumb spike reduced.
Notes: The presence of hatchlings and adults in close association suggest iguanodonts engaged in some
form of parental care early in their youngs lives, similar to hadrosaurs. Hatchling shown below.
Verd et al.
ZP
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100
Morelladon beltrani
Anatomical Characteristics: Elongated vertebral spines formed a modest sail/ridge over the body.
Age: Early Cretaceous, Late Barremian.
Distribution: Spain; Arcillas de Morella.
Notes: Shared its habitat with Mantellisaurus and Iguanodon, to which it was related. Sail may have
functioned in display, though not as well developed as Ouranosaurus.
Hadrosaurs
Baso-Hadrosaurs
Sirindhorna khoratensis
Fossil Remains: Majority of skull.
Anatomical Characteristics: Beak narrow and rounded, lower jaw fairly gracile.
Habits: Probably a generalist browser.
Notes: One of the most basal hadrosauroids known. Shared its habitat with Psittacosaurus, Siamodon,
Ratchasimasaurus, spinosaurs, and titanosaurs.
Shibata et al.
Koshisaurus katsuyama
Anatomical Characteristics: Possesses an antorbital fossa on the maxilla.
Distribution: Main Island Japan; Kitadani.
Notes: The most basal known hadrosaur. Shared its habitat with Fukuisaurus and Fukuititan. Main
enemy Fukuiraptor.
Zhuchengosaurus maximus
See Page 294
Notes: Probably synonymous with Shantungosaurus.
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101
Batyrosaurus rozhdestvenskyi
Age: Late Cretaceous, Santonian to Campanian.
Distribution: Kazakhstan; Bostobinskaya.
Notes: Remains assigned to Arstanosaurus akkurganensis are probably a juvenile of this genus.
Huehuecanauhtlus tiquichensis
Anatomical Characteristics: Vertebral spines over hips form shallow ridge.
Distribution: Western Mexico; Barranca Los Bonetes.
Notes: One of the best known southerly hadrosaurs. May be an euhadrosaur.
Ramrez-Velasco et al.
Jeyawati rugoculus
Anatomical Characteristics: Snout elongated and slightly curved down, beak narrow. Wrinkled bone on
upper eye socket likely contained excessive blood vessels.
Distribution: New Mexico; Moreno Hill.
Habitat: Coastal swamps and marshes.
Habits: Low-level browser and grazer. May have had a display structure above or behind eyes that
could be flushed red with excess blood.
Notes: Classification as a baso-hadrosaur uncertain, may be a derived iguanodontoid. Shared its habitat
with Nothronychus and Zuniceratops.
Nanyangosaurus zhugeii
5 m (16 ft) TL, 600 kg (1,200 lb)
Anatomical Characteristics: Hand relatively long.
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102
Distribution: Eastern China; Sangping.

Notes: The earliest confirmed hadrosaur.
Xuwulong yueluni
4 m (13 ft) TL, 300 kg (650 lb)
Anatomical Characteristics: Snout narrow, beak rounded, lower jaw fairly robust.
Distribution: Northwest China; Xinminpu.
Habits: Middle- and low-level browser.
Notes: Main enemy Xiongguanlong. Shared its habitat with Beishanlong and Suzhousaurus.
paleofile.com
ZP
Lapampasaurus cholinoi
Fossil Remains: Minority of skeletons.
Age: Late Cretaceous, Late Campanian or Early Maastrichtian.
Distribution: Southern Argentina; lower Allen.
Notes: Yet to be described in detail, could be synonymous with Willinakaqe.
Yunganglong datongensis
Distribution: Eastern China; Zhumapu.
Notes: Probably includes remains assigned to Bactrosaurus.
Gongpoquansaurus mazongshanensis
Anatomical Characteristics: Skull subrectangular, hand short.
Distribution: Central China; Xinminbo Group.
Notes: New genus for Probactrosaurus mazongshanensis as has been found to be less derived than
Probactrosaurus. Some remains may belong to Equijubus.
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103
Zhanghenglong yangchengensis
5 m (16 ft) TL, 600 kg (1,200 lb)
Age: Late Cretaceous, Middle Santonian.
Habits: Likely a mid-level browser.
Notes: An advanced non-hadrosaurid hadrosauroid.
Probactrosaurus gobiensis
See Page 292
Notes: Probably a moderately derived hadrosaur, close to euhadrosaurs, rather than an iguanodont.
Zuoyunlong huangi
Age: Early Late Cretaceous, Cenomanian.
Distribution: Northeast China; Zhumapu.
Notes: Suggests that basal hadrosaurs first arrived in North America right around the Early-Late
Cretaceous boundary. Closest relative appears to be the slightly earlier Probactrosaurus.
Plesiohadros djadokhtaensis
Anatomical Characteristics: Skull subtriangular, beak rounded.
Distribution: Mongolia; Djadokhta.
Habits: Likely a generalist browser and grazer.
Notes: Yet to be described in detail.
Datonglong tianzhenensis
Anatomical Characteristics: Lower jaw robust.
Age: Late Cretaceous.
Distribution: Northern China; Huiquanpu.
Notes: Shared its habitat with Huabeisaurus and Saichania (=Tianzhenosaurus).
Euhadrosaurs
Hadrosaurines
Kundurosaurus nagornyi
Page
104
Anatomical Characteristics: Snout short.

Distribution: Eastern Russia; Udurchukan.
Notes: Appears to be an intermediate between Saurolophus and Edmontosaurus. Shared its habitat
with Amurosaurus.
Godefroit et al.
Latirhinus uitstlani
7.5 m (25 ft) TL, 2.5 tonnes
Anatomical Characteristics: Beak broad, nasal crest well developed.
Distribution: Northeastern Mexico; Cerro del Pueblo.
Habits: Low-level browser.
Notes: Skeletal characteristics intermediate between hadrosaurines and lambeosaurines. Shared its
habitat with Hypacrosaurus and Coahuilaceratops.
Willinakaqe salitralensis
Fossil Remains: Several partial skulls and skeletons, juvenile and adult.
Anatomical Characteristics: Snout short, beak squared off, low ridge between nasal openings.
Age: Late Cretaceous, Late Campanian and/or Early Maastrichtian.
Distribution: Southern Argentina; lower Allen.
Habits: Likely consumed low lying vegetation.
Notes: The southerly hadrosaur known from the most complete remains. Shows strong similarities to
North American Kritosaurus. Shared its habitat with Austroraptor and a variety of sauropods.
Page
105
Valieri et al.
ZP
Edmontosaurus regalis
See Page 297
Anatomical Characteristics: May have possessed a small soft tissue headcrest or comb at the back of
the head.
Notes: Crest may be indicative of sexual dimorphism. Alternatively, the crest may simply be displaced
neck tissue.
Ugrunaaluk (or Edmontosaurus) kuukpikensis
Fossil Remains: Majority of skulls and skeletons, juvenile.
Anatomical Characteristics: In juveniles, snout fairly short and sloping, beak not especially broad.
Distribution: Alaska; Prince Creek.
Habitat: Well-watered forest with humid summers and cool winters.
Habits: Mostly a low-level grazer, but likely adaptable.
Notes: Shared its habitat with Alaskacephale and Pachyrhinosaurus. Main enemy Nanuqsaurus.
Page
106
Mori et al.
Augustynolophus (or Saurolophus) morrisi
Anatomical Characteristics: Beak spoon-shaped. Solid spike-like crest emerges from just before the
orbits, angled slightly backwards. Tip of crest slightly bifurcated, may have acted as an attachment point
for soft tissue.
Distribution: California; Moreno.
Notes: Appears intermediate between Prosaurolophus and Saurolophus.
Prieto-Mrquez et al.
Maiasaura (or Brachylophosaurus) peeblesorum
See Page 302
Habits: Growth rapid, reaching sexual maturity in about three years. Skeletal maturity acquired in eight
years. Known to have fed on rotten wood, though this was likely not a major component of its diet.
Acristavus (or Brachylophosaurus) gagslaroni
Fossil Remains: Majority of skulls and minority of skeleton.
Anatomical Characteristics: Beak narrow, squared off. Lacked cranial crest.
Distribution: Montana, Utah; Two Medicine, Wahweap.
Habitat: Seasonally dry woodlands.
Habits: Middle- and low-level browser.
Notes: Too little is known to determine if the lack of a crest is gender related.
Page
107
Gates
ZP
Probrachylophosaurus (or Brachylophosaurus) bergei

Fossil Remains: Partial skulls and skeleton.
Anatomical Characteristics: Beak narrow and squared off. Modest nasal arch present. Short, flat,
horizontal crest projects just over orbits.
Distribution: Montana; Judith River.
Habits: Low- and mid-level browser.
Notes: Crest and age intermediate between crestless Acristavus and fully crested Brachylophosaurus.
Shared its habitat with a variety of ceratopsians.
Freedman Fowler and Horner
ZP
Rhinorex (or Kritosaurus) condrupus

Fossil Remains: Majority of skull and partial skeleton, skin impressions
Anatomical Characteristics: Snout strongly downturned, shallow crest ridge over large nasal openings.
Skin covered in small, non-overlapping, roughly hexagonal scales interspersed with occasional, larger,
slightly domed scales.
Distribution: Utah; Neslen.
Habitat: Coastal region.
Notes: Potentially synonymous with Kritosaurus (or Gryposaurus) monumentensis.
Page
108
Gates and Scheetz
ZP
Canardia garonnensis
Distribution: France; Marnes dAuzas.
Notes: Possibly a basal lambeosaurine, along with close relative Aralosaurus.
Lambeosaurines
Tsintaosaurus sphinorhinus
See Page 308
Anatomical Characteristics: Crest is actually a fan or leaf shaped structure similar to Olorotitan, rather
than just a forward projecting tube.
Prieto-Mrquez and Wagner

Blasisaurus canudoi
Age: Late Cretaceous, Late Masstrichtian.
Distribution: Spain; Arn.
Notes: Potentially synonymous with Arenysaurus, but too little is known to be certain.
Page
109
Kazaklambia convincens
Fossil Remains: Partial skull and majority of skeleton, juvenile.
Anatomical Characteristics: Standard for juvenile lambeosaurine.
Distribution: Kazakhstan; Dabrazinskaya Svita.
Notes: A basal Hypacrosaurus-type lambeosaurine; originally designated a species of Corythosaurus,
and then Procheneosaurus. Procheneosaurus remains, which consist solely of juveniles, are
assignable to other Hypacrosaurus-type lambeosaurines, namely Hypacrosaurus and kin in North
America and Kazaklambia in Asia. Kazaklambia is the most basal known Hypacrosaurus-type
lambeosaurine and may indicate an Asiatic origin for this group.
Adelolophus hutchisoni
Age: Late Cretaceous, Early Campanian
Distribution: Utah; Wahweap.
Notes: Earliest known North American lambeosaurine hadrosaur. Shared its habitat with
Diabloceratops and Acristavus, main enemy Lythronax.
Magnapaulia laticaudus
See Page 314
Fossil Remains: Partial skull and skeleton, skin impressions.
Anatomical Characteristics: Small non-overlapping roughly hexagonal scales on upper portions of body,
slightly larger osteoderm-like scales scattered on tail region.
Notes: New genus for Hypacrosaurus laticaudus.
Mrquez et al.
Page
110
Special Thanks To:
A Field Guide to Mesozoic Birds and Other Winged Dinosaurs, by Matthew Martyniuk
Antediluvian Salad, by Duane Nash
Archosaur Musings, by Dave Hone
Bite Stuff, by Jamie Headden
Deviant Art and all of its talented artists
Dino Goss, by Matthew Martyniuk
DinoChecker.com
DinoData.org
Dinosaur Art: The Worlds Greatest Paleoart, by Steve White
DinosaurCentral.com
Dinosaur Tracking, by Brian Switek
Laelaps, by Brian Switek
MarkWitton.com and Blog, by Mark Witton
Palaeocritti.com
Paleofile.com
PLOS ONE, PeerJ, and other open access scientific journals
Prehistoric-Wildlife.com
Skeletal Drawing.com, by Scott Hartman
Species New to Science, by Pskhun
Tetrapod Zoology, by Darren Naish
Tetrapod Zoology Podcast, by Darren Naish and John Conway
The Complete Dinosaur, 2nd Edition, by Brett-Surman, Holtz, Farlow, Walters, and others
The Dinosaur Toy Forum and all of its fabulous members
The paleontologists from around the world that make these remarkable discoveries
The Princeton Field Guide to Dinosaurs, by Gregory S. Paul
The Society of Vertebrate Paleontology
The Theropod Database and Blog, by Mickey Mortimer
Theropoda Blog, by Andrae Cau
Wikipedia, the Wikimedia Foundation, and all of its knowledgeable contributors
Page
111

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Dinosaur Field Guide Supplement 2nd Ed

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The Dinosaur Field Guide Supplement

September 2010 December 2015

By, Zachary Perry (ZoPteryx)

This supplement is intended to be a companion for Gregory S. Pauls impressive work

Alcober & Martinez

Sereno & Abraczinskas

Habits: Likely an aquatic or semi-aquatic predator akin to its relatives.

Distribution: Colorado; upper Morrison.

Age: Early Cretaceous, Barremian.

Get Away Trike! Blog

Habits: Big game hunter.

Distribution: Northeast China; Yixian.

Get Away Trike! Blog

Skull modified after Sinkkonen

Age: Early Cretaceous, Barremian.

Age: Late Cretaceous, Campanian.

Habitat: Semidesert with dunes and oases.

Distribution: Central China; Qiupa.

Distribution: Southern China; Nanxiong.

Age: Late Cretaceous, Late Campanian or Early Maastrichtian.

Colbert & Rowe

Sertich & Loewen

Efraasia (or Plateosaurus) minor

Unaysaurus (or Plateosaurus) tolentinoi

Age: Early Cretaceous, Barremian.

Distribution: Mongolia; Baruungoyot.

Distribution: New Mexico; Kirtland.

Zheng, Jin, & Xu

Medusaceratops (or Albertaceratops) lokii

Evans & Ryan

Pachyrhinosaurus (or Centrosaurus) perotorum

Fiorillo and Tykoski

Hunt & Lehman

Coahuilaceratops (or Chasmosaurus) magnacuerna

Vagaceratops (or Chasmosaurus) irvinensis

Wick and Lehman

Titanoceratops (or Pentaceratops) ouranos

Brown & Henderson

Brown & Henderson

Sellacoxa (or Barilium) pauli

Distribution: Eastern China; Sangping.

Anatomical Characteristics: Snout short.

Probrachylophosaurus (or Brachylophosaurus) bergei

Freedman Fowler and Horner

Rhinorex (or Kritosaurus) condrupus

Gates and Scheetz

Prieto-Mrquez and Wagner

Special Thanks To:

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