ArticlePDF Available

A Second Specimen of Citipati Osmolskae Associated With a Nest of Eggs from Ukhaa Tolgod, Omnogov Aimag, Mongolia


Abstract and Figures

Adult dinosaurs preserved attending their nests in brooding positions are among the rarest vertebrate fossils. By far the most common occurrences are members of the dinosaur group Oviraptorosauria. The first finds of these were specimens recovered from the Djadokhta Formation at the Mongolian locality of Ukhaa Tolgod and the Chinese locality of Bayan Mandahu. Since the initial discovery of these specimens, a few more occurrences of nesting oviraptors have been found at other Asian localities. Here we report on a second nesting oviraptorid specimen (IGM 100/1004) sitting in a brooding position atop a nest of eggs from Ukhaa Tolgod, Omnogov, Mongolia. This is a large specimen of the ubiquitous Ukhaa Tolgod taxon Citipati osmolskae. It is approximately 11% larger based on humeral length than the original Ukhaa Tolgod nesting Citipati osmolskae specimen (IGM 100/979), yet eggshell structure and egg arrangement are identical. No evidence for colonial breeding of these animals has been recovered. Reexamination of another "nesting" oviraptorosaur, the holotype of Oviraptor philoceratops (AMNH FARB 6517) indicates that in addition to the numerous partial eggs associated with the original skeleton that originally led to its referral as a protoceratopsian predator, there are the remains of a tiny theropod. This hind limb can be provisionally assigned to Oviraptoridae. It is thus at least possible that some of the eggs associated with the holotype had hatched and the perinates had not left the nest.
Content may be subject to copyright.
Copyright © American Museum of Natural History 2018 ISSN 0003-0082
Number 3899, 44 pp. April 26, 2018
A Second Specimen of Citipati Osmolskae
Associated with a Nest of Eggs from Ukhaa Tolgod,
Omnogov Aimag, Mongolia
Adult dinosaurs preserved attending their nests in brooding positions are among the rarest
vertebrate fossils. By far the most common occurrences are members of the dinosaur group
Oviraptorosauria. e rst nds of these were specimens recovered from the Djadokhta Forma-
tion at the Mongolian locality of Ukhaa Tolgod and the Chinese locality of Bayan Mandahu.
Since the initial discovery of these specimens, a few more occurrences of nesting oviraptors
have been found at other Asian localities.
Here we report on a second nesting oviraptorid specimen (IGM 100/1004) sitting in a
brooding position atop a nest of eggs from Ukhaa Tolgod, Omnogov, Mongolia. is is a large
specimen of the ubiquitous Ukhaa Tolgod taxon Citipati osmolskae. It is approximately 11%
larger based on humeral length than the original Ukhaa Tolgod nesting Citipati osmolskae
specimen (IGM 100/979), yet eggshell structure and egg arrangement are identical. No evidence
for colonial breeding of these animals has been recovered.
Reexamination of another “nesting” oviraptorosaur, the holotype of Oviraptor philoceratops
(AMNH FARB 6517) indicates that in addition to the numerous partial eggs associated with
the original skeleton that originally led to its referral as a protoceratopsian predator, there are
the remains of a tiny theropod. is hind limb can be provisionally assigned to Oviraptoridae.
It is thus at least possible that some of the eggs associated with the holotype had hatched and
the perinates had not le the nest.
1 Division of Paleontology, American Museum of Natural History.
2 Richard Gilder Graduate School, American Museum of Natural History
3 Center for Functional Anatomy and Evolution, Johns Hopkins University.
4 Department of Biological Science, Florida State University, Tallahassee.
Adult dinosaur remains denitively associated with nests of eggs are among the rarest
vertebrate fossils (Varricchio et al., 2008). Interestingly, the rst such combination to be
found was the holotype of Oviraptor philoceratops (AMNH FARB 6517) (Osborn, 1924) (g.
1). is association was incorrectly interpreted over 90 years ago as a case in which O. philo-
ceratops was preying upon the eggs of Protoceratopos andrewsi. In 1994 it was discovered that
the egg type associated with the holotype was an oviraptorid (Norell et al., 1994). AMNH
FARB 6517 (the holotype of O. philoceratops) was therefore a parent rather than a predator
(Norell et al., 1994, 1995, 2001). Although fossil remains of nesting dinosaurs have become
more commonplace (Dong and Currie, 1996; Fanti et al., 2012), the most complete speci-
mens are found in the bright red, unstructured sandstone deposits of Ukhaa Tolgod, Omno-
gov Aimag, Mongolia (Dashzeveg et al., 1995; Dingus et al., 2008). In 1995 our research
group reported on the rst of these specimens, IGM 100/979 (Norell et al., 1995), which was
excavated during the 1993 expedition (gs. 2, 3). Later, aer more detailed examination, this
specimen was referred to the ubiquitous Ukhaa Tolgod taxon Citipati osmolskae (Clark et al.,
1999, 2001). Here we report on a second dinosaur nest attended by an adult oviraptorid
(IGM 100/1004) from Ukhaa Tolgod.
IGM 100/1004 (g. 4) was discovered during the 1995 installment of the American
Museum of Natural History–Mongolian Academy of Sciences Paleontological Expedition.
IGM 100/1004 was found on the face of the Camel’s Humps amphitheater (g. 5), at the
southern terminus of the Death Row sublocality. Dinosaur nests are very common at Ukhaa
Tolgod, and there is strong geological evidence that they were preserved as sequential event
horizons caused by rapidly collapsing sand dunes (Dingus et al., 2008). is Camel’s Hump
fossiliferous horizon is the result of one such catastrophic event that preserved several dino-
saurian taxa in life positions. ese include Pinacosaurus grangeri (IGM 100/3186, IGM
100/1014) (Hill et al., 2003; 2015) and Shuvuuia deserti (IGM 100/977) (Chiappe et al., 1998;
Schweitzer et al., 1999). e specimen was excavated over a number of days (g. 6). Some of
this excavation was lmed and photographed, appearing as part of a magazine story (Web-
ster, 1996) and documentary on the 1995 expedition (Truitt, 1996). Because of the steepness
of the exposure the specimen had to be carefully rigged to lower it down the escarpment.
Because of the excellent preservation of specimens at this locality and the great deal of expo-
sure, the absence of closely packed nests makes it unlikely that it was a colonial or group
nesting site for Citipati osmolskae.
Some of these fossils, including dinosaur nests such as the one described here, have been exca-
vated in accordance with Mongolian law by professionals and are part of the Mongolian Academy
of Sciences Institute of Paleontology collection (g. 7). At Ukhaa Tolgod we have excavated several
of these occurrences; sadly, others have been illegally poached (M.A.N., personal obs.) (g. 8).
IGM 100/1004 was purposely le incompletely prepared from the matrix so as to preserve the
relationship between the skeleton and the underlying nest (gs. 4, 8, 10–12). Because the Citipati
osmolskae type specimen (IGM 100/978) is remarkably preserved (Clark et al., 2001; Clark et al.,
2002), it is more important to keep this specimen in context than to remove the individual bones
and eggs from the matrix. In addition to the photographs and illustrations found herein, 3D surface
scanning of the specimen was conducted using a Space Spider scanner (Artec, Luxemborg). An .stl
(stereolithography) le of the entire block is available from the senior author.
e specimen is an incomplete skeleton of a large adult Citipati osmolskae (table 1) sitting
atop a nest of eggs (gs. 4, 9–11). Much of the skeleton including the skull, tail, and parts of
the hind limbs had eroded prior to its discovery in 1995. However, because IGM 100/1004 was
most likely buried alive (Dingus et al., 2008), these elements were probably present at the time
of burial. e skeleton is the largest Citipati osmolskae specimen yet reported. Based on the
length of the humerus it is 11% larger than the other Ukhaa Tolgod nester (IGM100/979) and
about 6% larger than the Citipati osmolskae holotype (IGM 100/978) (table 1).
FIGURE 1. AMNH FARB 6517 the type specimen of Oviraptor philoceratops found associated with a nest of
eggs. From Osborn (1924).
FIGURE 2. IGM 100/979. e nesting Citipati osmolskae as it was rst found at Ukhaa Tolgod in 1993. Le
Amy Davidson, right Louis Chiappe.
In what appears to be the stereotypical nesting posture for the taxon, the forelimbs extend
from the torso, so that the humeri lie near perpendicular to the body and the distal limb ele-
ments (radii, ulnae, and both manus) lie nearly parallel to the nest, with the palmar surfaces
of the manus directed toward the torso. e neck is arched back beside the torso, suggesting
that the head, which is missing, was nestled next to the body. is posture may reect the
stereotypical resting position of derived theropods (including modern birds) (Xu and Norell,
2004). A similar position of the head and neck is inferred for a nesting specimen of the ovi-
raptorid, Nemegtomaia barsboldi (Fanti et al., 2012).
e referral of IGM 100/1004 to Citipati osmolskae is based on a number of characters
unique to selective subsets of oviraptorids and a combination of characters present in the
holotype IGM 100/978 (Clark et al., 2001). ese include: (1) fusion of the greater and lesser
trochanters into a trochanteric crest on the femur (Balano and Norell, 2012); (2) elongate
cervical vertebrae that are at least twice as long as wide (Clark et al., 2001), the longest relative
FIGURE 3. IGM 100/979 in dorsal view aer preparation.
10 cm
10 cm
FIGURE 4. IGM 100/1004. An adult Citipati osmolskae collected in 1995 from the Death Row sublocality at
Ukhaa Tolgod, Omnogov Aimag, Mongolia, in dorsal view (opposite page and above).
length-to-width ratio for any oviraptorid; (3) ischia that form a symphysis distally (Clark et al.,
2001); and (4) a U-shaped furcula with an elongate hypocleidium (Nesbitt et al., 2009).
Axial Skeleton
C V: Eleven cervical vertebrae are present in IGM 100/1004, excluding
the atlas and axis, which are not preserved. is corresponds to the 12 or 13 cervical vertebrae
typically found in oviraptorosaurs (Osmólska et al., 2004). Because the vertebral centra remain
encased in matrix, only the neural arches are visible in dorsal view. In dorsal view (g. 13), the
vertebrae display the characteristic X shape seen in other maniraptorans (Makovicky and Sues,
1998), however as in the Citipati osmolskae holotype (IGM 100/978) the cervicals are more
elongate. e anterior vertebrae are heavily weathered, but low neural spines can be discerned
on the more posterior ones. e spines are centered on the neural arches as they are in other
oviraptorosaurs. e postzygapophyses do not diverge signicantly from the midline, even in
the more posterior vertebrae, thereby diering from the morphology present in Conchoraptor
gracilis and Khaan mckennai (g. 14) (Balano and Norell, 2012). e condition in Oviraptor
FIGURE 5. Death Row sub locality of the Camels Humps. Arrow signies where IGM 100/1004 was
FIGURE 6. Michael Novacek (le) and Mark Norell (right) excavating IGM 100/1004. Courtesy of Louis
philoceratops is dicult to determine, but it appears to be similar to the Citipati osmolskae
holotype (IGM 100/978) and IGM 100/979. e posterior cervical ribs are fused to the verte-
brae as they are in IGM 100/978, Avimimus portentosus, Anzu wyliei, Heyuannia huangi, Apa-
toraptor pennatus (Funston and Currie, 2016) and Khaan mckennai (Balano and Norell, 2012;
Lamanna et al., 2014).
D V: Ten trunk vertebrae are present in IGM 100/1004, including the
cervicodorsal vertebra (g. 4, and accompanying .stl le available as an online supplement
at, which is recognized by its expanded, fan-shaped trans-
verse processes (Osmólska et al., 2004). e Citipati osmolskae holotype (IGM 100/978) pre-
serves only seven trunk vertebrae as two were inadvertently destroyed during collection of
the specimen. e neural spines become taller in the more posterior regions of the trunk
series where they are approximately as elongate dorsoventrally as anteroposteriorly as in the
type specimen. e neural spines, however, are not preserved in the last ve vertebrae of this
series in IGM 100/1004. Similar to other oviraptorids, the transverse processes of the trunk
vertebrae are as wide as long, are square in dorsal view, and extend horizontally from the
neural arch. Ten dorsal ribs are preserved in IGM 100/1004, which are wide and attened to
FIGURE 7. A dinosaur nest being excavated at Ukhaa Tolgod in July 2013. Le to right: Suzann Goldberg,
Maraal Bayra, and Jian-Ye Chen.
a greater degree than in the holotype. Uncinate processes are preserved on the right side of
this specimen (g. 15). Similar processes also are present in a variety of derived theropod
taxa including IGM 100/978 and the oviraptorids Conchoraptor gracilis, Heyuannia huangi,
Apatoraptor pennatus, and Caudiptery (Ji et al., 1998; Clark et al., 2001; Lü, 2002; Funston
and Currie, 2016; Codd et al., 2007). e uncinate processes span two ribs, have expanded
heads at their anterior contact, and taper posteriorly as is typical of derived theropod dino-
saurs (Codd et al., 2007). ey articulate just proximal to the angle of the rib. e second
uncinate process (associated with trunk vertebrae 3 and 4) is the largest, and the fourth
process is the smallest. Overall the uncinate processes are relatively larger than in Conchorap-
tor gracilis, and more comparable in size to those of the Citipati osmolskae holotype (g. 16)
but slightly more gracile than in IGM 100/979.
S V: e sacral vertebrae are too heavily weathered to discern much mor-
phology. At least four are present, but we suspect there were ve as seen in the holotype. e
sacral ribs expand where they contact the ilia.
C V: No caudal vertebrae or chevrons are preserved, whose morphology
some contend can be used to determine the sex of the individual (Persons et al., 2015).
FIGURE 8. An unsuccessful poaching attempt that resulted in the destruction of an oviraptorid nest, summer
of 2014. Fragments of bone suggest that an adult may have been associated. Apparently the nest fell out of a
jacket as it was being ipped.
Forelimb and Pectoral Girdle
S: e scapula and coracoid are fused into a single element in IGM
100/1004 as in most oviraptorids outside of Caudipteryx, Conchoraptor gracilis, and Jiangxis-
aurus ganzhouensis (Lamanna et al., 2014). Although matrix and the surrounding bones largely
obscure the region of the coracoid, the scapulocoracoid appears to form a gentle arc as it does
in most other derived theropods. is arc is not as extreme as seen in more advanced mani-
raptorans, such as Velociraptor mongoliensis, where the scapulocoracoid shows an L-shaped
TABLE 1. IGM 100/1004 measurements (mm).
Scapula (right): 292.83
Humerus (right): 230.11
Radius (le): 212.44 212.44
Ulna (le): 211.46 211.46
Manus (total length): 284.26
MC II (le): 103.52
Phalanx I-1 (le): 92.92
Phalanx I-2 (le): 84.05
Phalanx II-1 (le): 68.3 68.3
Phalanx II-2 (le): 75.56 75.56
Phalanx II-3 (le) (two pieces): 53.0 (distal), 13.5 (proximal) 53.0 (distal)
13.5 (proximal)
Phalanx III-1 (le) 44.5
Phalanx III-2 (le) 48.2
Phalanx III-3 (le) 53.65
Phalanx III-4 (le) 57.8
Ilium anteroposterior length (le) ~253.0
Femur (le) ~402.4
Tibia (right, two pieces) 183 (distal)
270 (proximal)
MT I (right) 42.43
Phalanx I-1 (right) 31.92
Phalanx I-2 (right) 35.7
Phalanx II-1 (right) 53.15
Phalanx II-2 (right) 36.98
Phalanx IV-1 (right) 38.55
Phalanx IV-2 (right) 35.66
Phalanx IV-3 (right) 31.61
Phalanx IV-4 (right) 28.98
Phalanx IV-5 (right) 49.7
conguration contributing to a reorientation of the glenoid (Norell and Makovicky, 1997,
1999). e glenoid fossa, formed equally by these two elements, faces laterally. e pectoral
girdle is preserved in articulation; therefore, the scapular blade can be seen extending posteri-
orly, perpendicular to the rib shas. e scapular blade expands only weakly at its distal end.
is condition is dicult to compare to IGM 100/978, which has a heavily weathered distal
scapula. e acromion process of the scapula extends anteriorly and is in line with the dorsal
surface of the scapular blade, as in other oviraptorids except Ajancingenia yanshini, in which
this extension is more laterally directed.
F: e articulated furcula (g. 17) lies ush with the dorsal edge of the acromion
and scapular blade on the more medial side of the scapula as in Velociraptor mongoliensis
(Norell et al., 1997). e morphology of the furcula resembles that of IGM 100/978 in being
U-shaped with tapering epicleidial processes (Nesbitt et al., 2009). A swelling is present along
the ramus between the symphysis and the epicleidial process, which also resembles that of IGM
100/978. Too little of this element is preserved in IGM 100/979 to establish a similar swelling.
In IGM 100/1004 and 100/978, the hypocleidium is elongate and tapers distally. IGM 100/1004
does not possess a midline keel on the anterior surface of the furcula as is present in Oviraptor
philoceratops, but the lateral processes are more expansive anteroposteriorly than mediolater-
ally. e hypocleidium articulates with the sternum as has been suggested for O. philoceratops
and Heyuannia huangi (Barsbold, 1983; Lü, 2002).
S: Only the posterior surface of the right side of the sternum is visible (fig.
18), thus whether this element is paired, as in most oviraptorids (and other basal paravi-
ans), or fused, as in Ajancingenia yanshini, cannot be determined. The visible surface is
FIGURE 9. IGM 100/1004 in right lateral view. Anterior is to the right.
featureless. The anterior margin of the sternum has a sigmoidal rim resembling both IGM
100/978 and 100/979. The lateral margin bears two processes—a distally tapering cranial
process and a larger, more rounded caudal (xiphoid) process. The caudal process has a
distal cranial extension that is not present in other specimens of Citipati osmolskae (IGM
100/978, IGM 100/979) where this feature can be observed. The sternum is emarginated
between these processes for the costal articulations. Three sternal ribs are preserved near
this articulation in IGM 100/1004—the same number found in IGM 100/978 and 100/979
(fig. 18).
H: Both humeri are preserved in articulation with the radii and ulnae and are
similar in overall morphology to other oviraptorids (g. 19). ey have a sigmoidal shape with
a large deltopectoral crest (107 mm) spanning almost 50% of the total length of the element
(g. 4). e lateral margin of the crest is rugose and likely served as an attachment site for the
deltoid musculature. It is not present in Oviraptor philoceratops (g. 20). e proximal articular
surface is highly eroded on both sides but appears to have been mediolaterally elongate as in
all oviraptorids. e distal articulation is discernible only on the le side of IGM 100/1004 (g.
21). is region is wider than the humeral sha and has an anterior articulation with the radius
and ulna.
R: The radius and ulna (fig. 21) are approximately the same length and slightly
shorter than the humerus (table 1), yet the radius extends slightly further distally than the
ulna. The proximal articulation with the humerus is largely obscured by the overlying
humerus and ulna, but the distal articular surface is mediolaterally compressed and spatu-
late in form (fig. 22).
FIGURE 10. IGM 100/1004 in le lateral view. Anterior is to the le.
U: e ulna is best preserved on the le side of IGM 100/1004. Similar to other ovi-
raptorids (and most maniraptorans) other than Heyuannia huangi and Gigantoraptor erlianen-
sis, the sha is bowed posteriorly (Gauthier, 1986). Proximally, a small olecranon process is
present (~12 mm tall). As in other oviraptorids except for Apatoraptor pennatus (Funston and
Currie 2016), there is no indication of feather quill knobs (sensu Turner et al. 2007). A large
foramen found on the lateral side of the olecranon process appears to be the result of weather-
ing, preparation, or a large insect cavity. e last mentioned are found in many other Gobi
Desert specimens, on the sternal plates, ilium, and pubis of Velociraptor mongoliensis (IGM
100/985) (Norell and Makovicky, 1997: gs. 3, 9, 14; Fanti et al., 2012; Clark et at., 2001: g.
2). Just distal to the proximal end, the ulnar sha is mediolaterally compressed, however, the
distal end is compressed anteroposteriorly and expanded mediolaterally.
M: Only the left manus of IGM 100/1004 is preserved. It is 284.3 mm long and
makes up approximately 40% of the total forelimb length. Similar to other oviraptorids,
digits II and III are approximately the same length and both exceed the length of digit I.
FIGURE 11. IGM 100/1004 in anterior view.
Digit I (fig. 23) is large and robust relative to the rest of the digits that have a large curved
ungual. It is similar in relative size to the same digit in IGM 100/978, Khaan mckennai,
Conchoraptor gracilis, and Machairasaurus leptonychus (Longrich et al., 2010; Balanoff and
Norell, 2012), but does not achieve the level of robustness seen in Ajancingenia yanshini
(Barsbold, 1981).
C: e carpals and metacarpals are preserved on the le side of the specimen. e
semilunate carpal can be distinguished and covers the proximal ends of MC II and III as in
other oviraptorids. An additional small carpal, likely the radiale, is present at the distal end of
the radius, just proximal to the semilunate carpal. is attribution is in accord with Zanno and
Sampson (2005).
M: MC I is missing from the le manus, but MCs II and III are approximately
the same length as in other oviraptorids with the exception of Hagryphus giganteus (Zanno and
Sampson, 2005). e ulna sits on top of the proximal end of MC III, so that its length cannot
be measured with certainty. e metacarpals do not fuse proximally. Both preserved metacar-
pals have distal ginglymoid articulations. MC III appears to be mediolaterally compressed
towards the proximal end as in all known oviraptorids.
FIGURE 12. IGM 100/1004 in posterior view.
M P: IGM 100/1004 retains the plesiomorphic 2-3-4 phalangeal formula of
maniraptors. e ventral surfaces of the phalanges are relatively straight in lateral view. Phalanx
I-1 is similar to that of Khaan mckennai and other specimens of Citipati osmolskae, in that it is
robust compared with the other two digits and has a ginglymoid distal articulation. e collateral
ligament pits are deep, tear shaped and situated dorsally. In phalanx I-2, the ungual (g. 23) is
highly curved along its ventral margin. is element lacks an upturned dorsal lip on its dorso-
ventrally elongate articulation surface, diering from Chirostenotes pergracilis, Elmisaurus rarus,
Hagryphus giganteus, and Machairasaurus leptonychus (Osmólska, 1981; Currie and Russell, 1988;
Zanno and Sampson, 2005; Longrich et al., 2010). e large exor tubercle is separated from this
surface by a small space. A deep groove runs along the medial surface.
Digits II and III are approximately the same size and equally robust. Phalanx II-1 has
a large dorsal lip on its proximal articulation surface that is not present in IGM 100/978
and symmetrical extensor tubercles at its distal articulation (fig. 24). The collateral liga-
ment pits are deep, round and centrally positioned. Phalanx II-2 is broken and very little
morphology can be discerned on this element (fig. 21). The ungual of digit II (phalanx
II-3) is broken into two pieces, exaggerating its ventral curvature. Phalanx III-1 is short
with a relatively shallow, circular collateral ligament pit that is centrally placed. Phalanx
III-2 is subequal in length to III-1. It has an unusually tall dorsal lip (fig. 24) on the proxi-
mal articular surface, which is expressed to a greater degree than in IGM 100/978. The
collateral ligament pits are shallow and not easily discerned. Phalanx III-3 is slightly longer
and straighter than III-2, but similar in morphology to the more proximal phalanx. Pha-
lanx III-4 (ungual) differs little from the unguals of the other two digits. It has a dorso-
ventrally elongate articular surface with a large, anteriorly placed flexor tubercle, and a
dorsal lip as in the holotype and in Oviraptor philoceratops. The lateral surface similarly
bears a deep groove.
FIGURE 13. Close up of cervical vertebrae of IGM 100/1004.
FIGURE 14. Close up of cer-
vical vertebrae of: A. the Citi-
pati osmolskae holotype (IGM
100/978); B. Conchoraptor
gracilis (IGM 100/1203); and
C, the Oviraptor philocera-
tops holotype (AMNH FARB
6517). Note how long the ver-
tebrae of Citipati are relative
to their widths compared to
the other taxa.
Hind Limb and Pelvis
I: e three pelvic bones are not completely fused, as is the condition in all ovirap-
torids excluding Avimimus portentosus (Kurzanov 1983). Only the lateral surface of the le
ilium is visible in IGM 100/1004. It is dorsoventrally concave (g. 25). e dorsal margin of
the midportion and posterior edge of the element is missing, but enough remains to show
that the proportions of the preacetabular and postacetabular processes are roughly equal.
Although this is the condition typically found in oviraptorids, these proportions can vary
more widely within the more inclusive clade Oviraptorosauria (Osmólska et al., 2004). e
preacetabular margin is hooked, but it does not extend ventral to the acetabulum as it does
in Caudipteryx zoui and caenagnathids (Ji et al., 1998; Osmólska et al., 2004; Lamanna et al.,
2014). As in other oviraptorids, the cuppedicus fossa is evident as a at shelf on the ventral
surface of the preacetabular process. e brevis fossa is not visible. e pubic peduncle
extends ventrally indicating that the pubis also extended ventrally. e ischial peduncle is
not preserved.
P: e proximal region of the right pubis is the only region remaining in IGM 100/1004.
is portion shows the typical oviraptorid condition in that it is anteriorly concave and projects
vertically, and so is nearly perpendicular to the long axis of the ilium (Osmólska et al., 2004).
I: Almost the entire le ischium is visible in IGM 100/1004, although its proximal
articulation with the ilium is obscured by matrix. e distal portion of the right ischium is also
2 cm
2 cm
FIGURE 15. Ribs and uncinate processes on the right side of IGM 100/1004.
1 cm
1 cm
1 cm
1 cm
FIGURE 16. e uncinate processes of: A. Conchoraptor gracilis (IGM 100/1203) and B. Citipati osmolskae
(IGM 100/978).
preserved. Only the internal surfaces of these elements are observable, but it can be discerned
that the ischia contact each other along their distal margin as they do in IGM 100/978 (g 26).
Similar to other oviraptorids, the obturator process is situated at approximately midsha. Alter-
ation from weathering has given the process a more rounded appearance than the triangular
shape present in IGM 100/978. A tubercle, which has not been described for other oviraptorids,
is present on the internal surface of both ischia near the level of obturator process. e poste-
rior edge of the ischium is straight distally, but concave posteriorly.
F: Both femora are preserved. However, only the distal end of the right femur is
present. Much of the proximal end of the le femur is obscured as it is still articulated with
the ilium. e sha of the femur is long and straight. e lesser and greater trochanters
appear to be fused into a single trochanteric crest as is found in IGM 100/978 and many
other oviraptorids like Gigantoraptor erlianensis, but not Khaan mckennai or Conchoraptor
gracilis (Balano and Norell, 2012). Due to heavy weathering, however, the presence of this
feature cannot be denitely conrmed. As in other oviraptorids, a small ridge runs along the
1 cm
FIGURE 17. Right oblique view of the pectoral region of IGM 100/1004.
1 cm
1 cm
FIGURE 18. e right sternal region of IGM 100/1004.
sha of the femur proximolaterally to distomedially from the lesser trochanter and ends just
above the medial condyle.
T: Both tibiae are preserved, although the right element is broken into two pieces. e
proximal end is not exposed on either side as the le side is covered by matrix and the right is
covered by its own distal region (g. 27). A small portion of the quadrangular-shaped bular crest
is exposed and extends from the lateral surface and appears again just below the level of the
proximal head (g. 27). e tibia is roughly circular in cross section at midsha. Distally, the
astragalus is not fused to the tibia and they contact along a strong horizontal suture. (g. 28).
F: Both bulae are present, however much the morphology is distorted. e proximal
and distal ends cannot be delineated on the le side and are not preserved on the right side.
A large tubercle is present approximately 1/3 down the length of the sha. e bula tapers
distally and is attenuated, but still appears to nearly reach the tarsals. e proximal bula is
distinctly bowed laterally.
P: e le foot is almost completely obscured under the torso. e right foot is partially
preserved from the midpoint of the metatarsals, but most of the phalanges are obscured by the
overlying tibia and bula.
T: Only the posterior surfaces of both astragali are visible (g. 29). e articular
surface is simple and smooth and similar to other oviraptorosaurs. No distal tarsals are visible
in IGM 100/1004.
M: The metatarsals are visible on the right side of the specimen. MT II,
III, and IV are eroded at their midpoints and only the unfused distal portions remain
intact. The distal portion of the metatarsus is unfused. MT I is complete. As in all ovirap-
torids, it is reduced to a small pyramidal bone that articulates with the distal end of MT
II. The distal articulation of MT I is gingylmoid. MT II and III are flattened in a dorso-
plantar direction. Although only partially preserved, MT III is similar to that of IGM
100/978 and does not appear to significantly taper proximally on its dorsal surface. MT
IV is more slender than MT III and laterally diverges from the other metatarsals. A deep,
oblong ligament pit is visible on its lateral surface. The distal articulation of the MT IV is
ovoid, enabling a large range of motion (Currie and Russell, 1988). MT V is not
1 cm
1 cm
FIGURE 19. Detail of the anterior right humerus of IGM 100/1004.
P: e phalanges are not all exposed (g. 27). e incomplete phalangeal for-
mula for the foot is 2-?-?-5. e ungual of digit I has a curved ventral margin and a deep
lateral groove as in the holotype IGM 100/978. Only the medial surfaces of the phalanges of
digit II are visible. e collateral ligament pits are well developed and dorsally positioned.
Digit III is not visible. e phalanges of digit IV are dorsoventrally attened, possibly a result
of postmortem distortion. Deep, elongate collateral ligament pits are present in this digit.
e ungual of digit IV has a deep lateral groove and highly curved ventral margin. Pro-
nounced dorsal lips are found on all of the pedal unguals.
IGM 100/1004 lies over the remains of 12 exposed partial to nearly complete eggs
arranged in a ring. Presumably if preparation were continued, more eggs would be discov-
2 cm
FIGURE 20. e humerus of the Oviraptor philoceratops holotype (AMNH FARB 6517).
5 cm
5 cm
FIGURE 21. e le arm and manus of IGM 100/1004.
ered. No traces of developing embryos are present. One egg is represented only by a small
mass of eggshell fragments beneath the right pectoral girdle. e eggs are grouped into ve
pairs within the ring. One egg and the mass of fragments lack partners. Assuming these two
eggs were also paired with corresponding eggs, there would have been at least 14 eggs in the
clutch. e eggs appear to have shied vertically with respect to one another around the ring,
but no observable eggs directly overlie one another. is suggests that only a single layer of
eggs is preserved. is stands in contrast to the clutch of IGM 100/979, which has stacked
eggs exposed in one portion of the clutch (Norell et al., 1995; Clark et al., 1999). is dier-
ence might represent an artifact of incomplete preparation, a taphonomic dierence between
the two specimens, individual variation in the arrangement of the clutch, or clutches cap-
tured at dierent stages of laying by the female. Nevertheless, in some of the video and
images taken during the excavation of the specimen there appears to be several eggs below
the level of the primary layer of eggs, especially just posterior to the right pes. ese are
hidden in the supporting eld jacket, which could not be removed without compromising
the integrity of the specimen.
1 cm
1 cm
mc-3 mp1-1
FIGURE 22. e le wrist of IGM 100/1004.
Only one egg of IGM 100/1004 is nearly completely exposed. It partially underlies the right
pes and is separated from it by matrix. e egg measures approximately 181 mm long. is
elongate egg appears slightly asymmetric, with the blunt pole pointing inward to the center of
the clutch. e other pole is more crushed; therefore, it remains unclear whether the egg would
have been asymmetric in life. e average width of all adequately exposed eggs (n = 6) is 66.8
mm. is provides an elongation index (egg length:width) of 2.7. All eggs show a ne lineartu-
berculate external ornamentation similar to “variant 2” of Mikhailov (1991: g. 8). ere are
eight to 11 ridges per centimeter, except for the poles, which are smooth. Eggs of this sort are
very common at the Ukhaa Tolgod locality and they oen compose entire or partial clutches
(g. 30), and on occasion are found as unassociated pairs (g. 31). As mentioned above, even
though eggs and nests of the oviraptorid type (as well as other dinosaur taxa) are ubiquitous
at this locality, the nests are not found in direct proximity, which suggests that the animals were
not communal or associative nesters.
e near-completely exposed egg described above was sampled for microstructural exami-
nation under a scanning electron microscope (SEM) (Zeiss EVO 60 Variable Pressure, Zeiss
Inc., Jena, Germany) and a petrographic microscope (Leitz Laborlux 11 POL S; Leitz Inc.,
Wetzlar, Germany). Radial thin sections were ground until transparent. Eggshell thickness and
microstructural dimensions were measured with soware (ImageJ, NIH, Bethesda, Maryland)
from both thin-section photomicrographs (g. 32A) and SEM images (g. 32B). e eggshell
measures 0.71–1.3 mm thick. e equator and blunt pole of the egg have a similar range of
thickness. e eggshell is composed of two structural calcite layers separated by an abrupt,
1 cm
mp 1-1
mp 1-1
mp 1-2
mp 1-2
FIGURE 23. e ungual of IGM 100/1004.
straight boundary. e inner is the mammillary layer (ML) composed of mammillary cones,
each with a radiating crystal fabric; and the outer is the continuous or cryptoprismatic (Jin et
al., 2007) layer (CL) that contains a squamatic structure typical of many nonavian theropod
and avian eggs. e surface is diagenetically eroded in many places. e ML averages 0.19 and
0.26 mm thick at the blunt pole and equator, respectively. e CL measures 0.53–0.72 mm thick
at the blunt pole and 0.50–1.0 mm thick at the equator. ese ranges reect dierences among
measured individual fragments and whether thickness is measured below the raised ornamen-
tation or in the “valleys” between ridges. e CL:ML ratio ranges, on average, from 2.2 in the
“valleys” at the equator to 3.7 beneath the ornamentation at the equator and blunt pole. No
crystal splaying (Jin et al., 2007) is evident along the ML-CL boundary. Accretion lines are
visible in thin section throughout the CL and their undulations mirror those of the eggshell
surface. ey are also visible under SEM in the outer third of the eggshell (g. 32B). e pore
canals are straight, narrow tubes (angusticanaliculate pore system) that vary slightly in diam-
eter along their width (g. 32B).
e above macro- and microstructural characters allow assignment to the oofamily Elon-
gatoolithidae, as is the case for all other eggs associated with oviraptorosaur skeletal remains
(Norell et al., 1994, 1995; Dong and Currie, 1996; Sato et al., 2005; Cheng et al., 2008; Weisham-
pel et al., 2008; Fanti et al., 2012; Pu et al., 2017; Wang et al., 2016). e eggs of IGM 100/1004
are nearly identical in size, shape, and microstructure to other conrmed Citipati osmolskae
FIGURE 24. Phalanges on digits 2 and 3 in IGM 100/1004.
1 cm
1 cm
eggs (IGM 100/971 [Norell et al., 1994, 2001] and IGM 100/979 [Norell et al., 1995; Clark et
al., 1999]). Citipati osmolskae eggs blur the distinction between the oogenera Elongatoolithus
and Macroolithus, as their length overlaps the range for Macroolithus eggs reported by Mikhailov
(1994), but they correspond more closely in surface ornamentation and microstructure to
Given the overlapping nature and susceptibility to intraspecic variation and/or tapho-
nomic alteration of some characters used to distinguish Elongatoolithus oospecies (e.g., eggshell
thickness, surface ornamentation), caution is warranted when attempting to assign a given egg
to any oospecies. We do not attempt to make a denitive ootaxonomic characterization of
Citipati osmolskae eggs, as this would likely require an extensive review of Elongatoolithidae,
which is outside the scope of this paper. Nevertheless, we oer detailed comparisons to existing
oospecies below.
As stated by Mikhailov (2014), Citipati osmolskae eggs are most similar to Elongatoolithus
frustrabilis (Mikhailov, 1994), also from the Djadokhta Formation of Mongolia. ey are also
similar in size and eggshell thickness to those of E. sigillarius, but lack the short transverse
ridges and nodes along the equator of that oospecies (Mikhailov, 1994). Citipati osmolskae eggs
similarly overlap the eggshell thickness range of eggs of E. subtitectorius, but this oospecies is
known solely from fragments, hindering further comparisons (Mikhailov, 1994). Citipati
osmolskae eggs, at about 180–190 mm long, exceed the size range given for E. frustrabilis by
Mikhailov (1994) (140–170 mm). However, they closely resemble this oospecies in other char-
acteristic features, presenting partially overlapping total eggshell thicknesses, CL:ML thickness
FIGURE 25. e le ilium of IGM 100/1004.
5 cm
1 cm
1 cm
ratios, and ridge densities of the lineartuberculate ornamentation. Similar eggs other than those
described by Mikhailov (1994) are known from the Ukhaa Tolgod (IGM 100/1125 [Grellet-
Tinner et al., 2006]) and Bayn Dzak (AMNH FARB 6633, AMNH FARB 6509 [Carpenter et
al., 1994]) localities.
As noted by Clark et al. (1999), Citipati osmolskae eggs are longer than those of both Ovi-
raptor philoceratops from the Djadokhta Formation of Mongolia (Osborn, 1924) and the nest-
ing oviraptorid from the Bayan Mandahu redbeds of Inner Mongolia, China (Dong and Currie,
1996). Citipati osmolskae eggs may also be longer than the estimated 140 to 160 mm long eggs
of a nesting specimen of Nemegtomaia barsboldi from the geologically younger Nemegt Forma-
tion, but these eggs are too incompletely preserved to make more condent macrostructural
or microstructural comparisons (Fanti et al., 2012).
Partial eggs containing oviraptorid embryos from the Bugin Tsav locality of the Nemegt
Formation of Mongolia are most similar to Elongatoolithus andrewsi or E. elongatus, though
the lack of complete eggs makes such assignments tentative (Weishampel et al., 2008). As in
Citipati osmolskae eggs, these eggs exhibit a straight ML-CL contact. eir CL:ML thickness
ratio is 2:3, within the range for that of C. osmolskae eggs. Weishampel et al. (2008) note that
the Bugin Tsav eggs dier from a Citipati osmolskae egg (IGM 100/971) in possessing more
variability in mammillary layer thickness within an egg.
Previous authors describe associations of Macroolithus yaotunensis (or similar) eggs with
oviraptorosaur skeletal remains from the Upper Cretaceous Nanxiong Formation in Jiangxi
Province, China. ese include eggs inside the pelvis of a female oviraptorid (Sato et al.,
2005) and eggs containing oviraptorid embryos (Cheng et al., 2008; Wang et al., 2016). ese
eggs all dier from Citipati osmolskae eggs by their undulating boundary between the ML
and CL and coarser lineartuberculate to ramotuberculate ornamentation. Other Macroolithus
oospecies described by Mikhailov (1994) dier from Citipati osmolskae eggs as well in pos-
sessing thicker eggshell overall and coarser lineartuberculate ornamentation with six to eight
ridges per centimeter.
Wang et al. (2016) describe a Citipati osmolskae egg (IGM 100/971) as most similar to those
of Elongatoolithus elongatus, but according to Mikhailov (1994), eggs of this oospecies are
substantially shorter (at 115–131 mm) than the complete Citipati osmolskae eggs of IGM
100/979 and IGM 100/1004. us, apart from being slightly more elongate, Citipati osmolskae
eggs are most similar to E. frustrabilis among currently described Elongatoolithus.
Midsha diaphyseal samples of a femur, dorsal rib, and bula of IGM 100/1004 were
sampled for petrographic histological analysis. e histological make-up of each of these
elements was characterized. Given that the specimen is very likely a reproductively active
adult, as is IGM 100/979, which was also found on a nest, the medullar cavity was examined
for the presence of medullar bone indicative of female oviposition and hence, sex (Schweitzer
FIGURE 26. e ischiac symphysis of IGM 100/1004.
2 cm
2 cm
FIGURE 27. e le
leg of IGM 100/1004.
et al., 2005). Counts of lines of arrested growth (LAGS) and annuli were made in each ele-
ment and back-calculated to infer age (Erickson et al., 2007). In a previous study, spacing
between the LAGs was used to develop a percentage of adult size to age growth curve from
which the developmental stage of the specimen was inferred (Erickson et al, 2007). It was
determined that the animal was somatically mature. Here additional histological details sup-
porting that interpretation are presented.
FIGURE 28. e right astragalus of IGM 100/1004.
1 cm
1 cm
The histological characterization (fig. 33A) shows that the diaphysis of the femur is
composed almost entirely of woven bone with plexiform vascularization (Francillon- Vieil-
lott et al., 1990). Negligible Haversian remodeling is present near the endosteal border of
the element and sporadically within the cortex (Francillon-Vieillott et al., 1990). Osteo-
clastic erosion spanning the entire endosteal border with a thin veneer of lamellar endos-
teal bone is present. The endosteal border of the cortex shows a thick veneer of lamellar
bone and lacks medullar bone as found in the femur of IGM 100/979 (Varricchio et al.,
2008). Nine definitive LAGs are present, two of which are within the EFS (external funda-
mental system, sensu Cormack, 1987). The spacing between the growth lines diminishes
5 cm
egg 1
egg 1egg 1
egg 1
egg 2
egg 2
pp4-3 pp4-5
pp4-1 pp4-2
pp4-2 pp4-4
FIGURE 29. e right pes of IGM 100/1004.
toward the outer cortex becoming attenuated in the EFS. Back-calculation to the center of
the medullar cavity based on the average width of the innermost three growth zones, is
continued to where the addition of an average zone width cannot be encompassed without
exceeding the center point and the remaining bone assumed to approximate the hatchling
bone radius, suggests four growth lines had been resorbed prior to death. The diminishing
growth-line spacing culminating in an EFS indicates the animal had reached advanced
somatic maturity. This coupled with the growth curve generated from it showing the speci-
men reached a somatic growth plateau suggests the animal died at or near full adult size
(Erickson et al., 2007). The idea that the animal was a somatic adult is also reinforced by
the fusion of the neural arches onto the centra in the dorsal vertebrae and the partial
fusion of the astragali to the distal tibiae.
e dorsal rib section is poorly preserved histologically (g. 33B). Much of the original
osseous matrix is eaced by fungal intrusion. Furthermore, the element is highly fragmented
where it was sampled for histological analysis. What can be gleaned is that the medullar region
is expansive, with only thin remnants of the cortex. e bulk of the primary matrix appears to
be woven bered bone with longitudinal vascularization. Very large osteoclast erosion rooms
5 cm
FIGURE 30. IGM 100/3505, a clutch of oviraptorid eggs from Ukhaa Tolgod.
with moderate Haversian inlling make up most of the inner cortex. e outermost cortex
shows up to six LAGS, one or two of which compose what appears to be an EFS. As with the
femur, these results suggest IGM 100/1004 is somatically mature. Estimation of the absolute
age of the specimen from this element was not possible owing to the sample’s fragmented
nature, which had made identication of its center uncertain.
e bula (g. 33C) is primarily composed of woven bone with longitudinal vasculariza-
tion that grades into parallel-bered matrix and then EFS structuring. Nevertheless, one of the
outermost growth zones is partially composed of woven bone with reticular vascularization.
Haversian remodeling is prevalent through the inner two-thirds of the cortex and sporadically
in the outermost zones. Ten prevalent growth lines are present. e inner three are annuli
FIGURE 31. IGM 100/1125, an unassociated pair of oviraptorid eggs from Ukhaa Tolgod.
whereas the remainders are LAGS. e zone spacing diminishes toward the periosteal surface
of the element. ree to four LAGs are encompassed in the EFS. Back-calculated age, based on
the mean width of the inner three growth zones, suggests IGM 100/1004 was approximately
13 years of age at the time of its demise (the same estimate derived from the femur; see above).
Again the histological indices suggest the animal was somatically mature.
As explained above, fossilized nests of dinosaurs with attending adults are incredibly rare.
Yet in addition to IGM 100/1004, four other oviraptorids have been discovered associated with
nests of eggs. First was Oviraptor philoceratops, AMNH FARB 6517 (Osborn, 1924). is speci-
men is quite poorly preserved (see Clark et al., 2002), and its misinterpreted association with
a group of eggs led to the original misdirected nomenclatural moniker. But an additional ele-
ment of this specimen has never been reported. Associated with the AMNH FARB 6517 skel-
eton are the remains of a juvenile oviraptorid (now numbered AMNH FARB 33092) that is
presumably from the same taxon (g. 34). e tibia of this animal is just 58.7 mm long and
likely represents a perinate (i.e., an embryo or hatchling) within the nest. ese bones will be
described in detail in another paper. Notably there are also several known multi-individual
associations of oviraptorosaur specimens both published (Funston et al., 2016) and unpub-
lished. e unpublished specimens include Khaan mckennai (IGM 100/3616) from Ukhaa
Tolgod, an undescribed oviraptorid from Udan Sayr (IGM MAE 16-08), and a Conchoraptor
gracilis specimen (IGM 199/1275) from Khulsan. All of these show at least one and usually
more adult or near-adult specimens accompanied by groups of juveniles. Collectively these
nds suggest oviraptorids were social animals throughout their lives.
Comparisons with other nesting specimens (Dong and Currie, 1996; Clark et al., 1999;
Fanti et al., 2012) also show a remarkable conservation in the posture of these animals sitting
on the nest in a stereotypical fashion. e bulkiest part of the torso sits in the middle of the
nest in a hollow in the center of a ring of tiered eggs. e forelimbs are extended away from
the body at the perimeter of the nest much in the same fashion as extant Aves. While it is easy
FIGURE 32. A. Radial thin section of IGM 100/1004 eggshell under plain polarized light. Scale bar equals 0.5
mm. B. Radial view of IGM 100/1004 eggshell under SEM. Scale bar equals 0.1 mm. e double-headed arrow
indicates a pore. Accretion lines are visible to either side of the pore in the upper third of the eggshell (marked
with triangles).
to homologize this brooding position among these taxa, some caution must be exercised.
Except for extant Aves and a putative occurrence of the dromaeosaurid Deinonychus anthir-
ropus (Grellet-Tinner and Makovicky, 2006), oviraptorids are unique among theropods in
showing multiple occurrences of individuals brooding their eggs. Some other associations of
theropods with eggs have been reported such as Troodon formosus (Varricchio et al., 2002), a
troodontid (IGM 100/1129) (Erickson et al., 2007), and a non-ornithuromorph bird (Varric-
chio and Barta, 2015). While not denitive without a more extensive taxonomic sample a
preliminary hypothesis of homology regarding brooding position can be made.
Discoveries of theropod dinosaur nests are increasingly common. However, those pre-
served with attending adults remain incredibly rare. Curiously, the overwhelming preponder-
ance of such specimens are oviraptorids (Erickson et al., 2007). e same holds true for most
of the maniraptoran group assemblages that have been discovered (e.g., Funston et al., 2016)—
these include unpublished specimens collected by American Museum of Natural History–Mon-
golian Academy of Sciences Expeditions and a specimen that was looted from the Gobi Desert
and repatriated to Mongolia. Taken together, these occurrences suggest that oviraptorid dino-
saurs were very social animals and that this sociality extended to their early days in the nest.
is nesting behavior, as is still seen in living birds, most likely has its origin at some level near
the base of Maniraptora. is once again pushes what was thought to be an “avian” character-
istic back into the evolutionary history of nonavian dinosaurs (e.g., Norell and Xu, 2005; Bala-
no et al., 2013; Nesbitt et al., 2009).
FIGURE 33. Histological sections of IGM/1004. Specimens are viewed using polarized petrographic light
microscopy under oil- immersion. White circles denote growth lines.
Great thanks to the 1995 American Museum of Natural History–Mongolian Academy of
Sciences eld crew. We thank D. Lawver and D.J. Simon for helpful comments on the manu-
script. B. Goldo and D. Ebel provided access to equipment and assistance with sectioning of
eggshell in the AMNH Department of Earth and Planetary Sciences. M. Eklund helped with
petrographic microscopy of the eggshell. H. Towbin assisted with scanning electron micros-
copy in the AMNH Microscopy and Imaging Facility. Many thanks are due Mick Ellison for
the gures and Lynn Merrill for support and scanning of the specimen. We thank Louis Psi-
hoyos for permission to use the image in gure 6. Marilyn Fox prepared the specimen. Greg
Funston and an anonymous reviewer greatly improved the manuscript. e 1995 expedition
was in part supported by the National Geographic Society. e histological analyses were sup-
ported by a National Science Foundation grant (EAR 0207744 to GE and MN). D.E.B. is sup-
1 cm
1 cm
FIGURE 34. e hind limb of a small oviraptorid (AMNH FARB 33092) associated with the Oviraptor philo-
ceratops holotype (AMNH FARB 6517).
ported by a Richard Gilder Graduate School Graduate Fellowship and M.A.N. by the Macaulay
Family endowment.
Balano, A.M., and M.A. Norell. 2012. Osteology of Khaan mckennai (Oviraptorosauria: eropoda).
Bulletin of the American Museum of Natural History 372: 1–77.
Balano, A.M., G.S. Bever, T. Rowe, and M.A. Norell. 2013. Complex patterns of endocranial expansion
near the origin of avian ight. Nature 501: 93–96.
Barsbold, R. 1981. [Toothless carnivorous dinosaurs of Mongolia]. Transactions of the Joint Soviet Mon-
golian Paleontological Expedition 15: 28–39. [in Russian]
Barsbold, R. 1983. O ptich’ikh chertakh v stroyenii khishchnykh dinozavrov. [“Avian” features in the
morphology of predatory dinosaurs]. Transactions of the Joint Soviet Mongolian Paleontological
Expedition 24: 96–103. [in Russian]
Carpenter, K., K.F. Hirsch, and J.R. Horner. 1994. Introduction. In K. Carpenter, K.F. Hirsch, and
J.R. Horner (editors), Dinosaur eggs and babies: 1–11. Cambridge: Cambridge University
Cheng, Y., J.I. Qiang, X.C. Wu, and H.-Y. Shan. 2008. Oviraptorosaurian eggs (Dinosauria) with embry-
onic skeletons discovered for the rst time in China. Acta Geologica Sinica (English Edition) 82:
Chiappe, L.M., M.A. Norell, and J.M. Clark. 1998. e skull of a relative of the stem-group bird
Mononykus. Nature 392: 275–278.
Clark, J.M., M.A. Norell, and L.M. Chiappe. 1999. An oviraptorid skeleton from the Late Cretaceous of
Ukhaa Tolgod, Mongolia, preserved in an avianlike brooding position over an oviraptorid nest.
American Museum Novitates 3265: 1–36.
Clark, J.M., M.A. Norell, and R. Barsbold. 2001. Two new oviraptorids (eropoda: Oviraptorosauria)
Upper Cretaceous Djadoktha Formation, Ukhaa Tolgod, Mongolia. Journal of Vertebrate Paleontol-
ogy 21 (2): 209–213.
Clark, J.M., M.A. Norell, and T. Rowe. 2002. Cranial anatomy of Citipati osmolskae (eropoda, Ovirap-
torosauria), and a reinterpretation of the Oviraptor philoceratops holotype. American Museum Novi-
tates 3364: 1–24.
Codd, J., P. Manning, M.A. Norell, and S. Perry. 2007. Avian-like breathing mechanics in maniraptoran
dinosaurs. Proceedings of the Royal Society of London, Series B, Biological Sciences 275 (1631):
Cormack, D. 1987. Hams histology. New York: Lippincott, 732 pp.
Currie, P.J., and D.A. Russell. 1988. Osteology and relationships of Chirostenotes pergracilis (Saurischia,
eropoda) from the Judith River (Oldman) Formation of Alberta, Canada. Canadian Journal of
Earth Sciences 25: 972–986.
Dashzeveg D., et al. 1995. Extraordinary preservation in a new vertebrate assemblage from the Late
Cretaceous of Mongolia. Nature 374: 446–449.
Dingus, L., et al. 2008. e geology of Ukhaa Tolgod (Djadokhta Formation, Upper Cretaceous, Nemegt
Basin, Mongolia). American Museum Novitates 3616: 1–40.
Dong, Z.-M., and P.J. Currie. 1996. On the discovery of an oviraptorid skeleton on a nest of eggs at Bayan
Mandahu, Inner Mongolia, People’s Republic of China. Canadian Journal of Earth Sciences 33:
Erickson, G., K.C. Rogers, D. Varricchio M.A. Norell, and X. Xu. 2007. Growth patterns in brooding
dinosaurs reveals the timing of sexual maturity in non-avian dinosaurs and genesis of the avian
condition. Biology Letters 3: 558–561.
Fanti, F., P.J. Currie, and D. Badamgarav. 2012. New specimens of Nemegtomaia from the Baruungoyot
and Nemegt Formations (Late Cretaceous) of Mongolia. PLoS One 7: e31330.
Francillon-Vieillot, H., et al. 1990. A microstructure and mineralization of vetebrate skeletal tissues. In
J.G. Carter (editor), Skeletal biomineralization: patterns, processes and evolutionary trends: 471–
530. New York: Van Nostrand Reinhold.
Funston, G.F., et al. 2016. e rst oviraptorosaur (Dinosauria: eropoda) bonebed: evidence of gre-
garious behaviour in a maniraptoran theropod. Scientic Reports 6 (1): 35782.
Funston, G.F., and P.J. Currie. 2016. A new caenagnathid (Dinosauria:Oviraoptorosauria) from the
Horseshoe Canyon Formation of Alberta, Canada, and a reevaluation of the relationships of Cae-
nagnathidae. Journal of Vertebrate Paleontology 36 (4): e1160910. [doi:
Gauthier, J.A. 1986. Saurischian monophyly and the origin of birds. Memoirs of the California Academy
of Sciences 8: 1–15.
Grellet-Tinner, G., L. Chiappe, M.A. Norell, and D. Bottjer. 2006. Dinosaur eggs and nesting behaviors:
a paleobiological investigation. Palaeogeography, Palaeoclimatology, Palaeoecology 232: 294–321.
Grellet-Tinner, G. and P.J. Makovicky. 2006. A possible egg of the dromaeosaur Deinonychus antirrhopus:
phylogenetic and biological implications. Canadian Journal of Earth Sciences 43 (6): 705–719.
Hill, R.V., L.M. Witmer, and M.A. Norell. 2003. A new specimen of Pinacosaurus grangeri (Dinosauria:
Ornithischia) from the Late Cretaceous of Mongolia: ontogeny and phylogeny of ankylosaurs.
American Museum Novitates 3395: 1–29.
Hill, R.V., M. D’Emic, G.S. Bever, and M.A. Norell. 2015. A complex hyobranchial apparatus in a Cre-
taceous dinosaur and the antiquity of the paraglossalia in avians. Zoological Journal of the Linnean
Society 175 (4): 892–909.
Ji, Q., P.J. Currie, M.A. Norell, and S.-A Ji. 1998. Two feathered theropods from the Upper Jurassic/Lower
Cretaceous strata of northeastern China. Nature 393: 753–761.
Jin, X., Y. Azuma, F.D. Jackson, and D.J. Varricchio. 2007. Giant dinosaur eggs from the Tiantai basin,
Zhejiang Province, China. Canadian Journal of Earth Sciences 44: 81–88.
Kurzanov, S.M. 1983. New data on the pelvic structure of Avimimus. Paleontological Journal 1983 (4):
Lamanna, M.C., H.-D. Sues, E.R. Schachner, and T.R. Lyson. 2014. A new large-bodied oviraptorosaurian
theropod dinosaur from the latest Cretaceous of western North America. PLoS One: e9202.
Longrich, N.R., P.J. Currie, and D. Zhi-Ming . 2010. A new oviraptorid (Dinosauria: eropoda) from
the Upper Cretaceous of Bayan Mandahu, Inner Mongolia. Palaeontology 53 (5): 945–960.
Lü, J. 2002. A new oviraptorosaurid (eropoda: Oviraptorosauria) from the Late Cretaceous of south-
ern China. Journal of Vertebrate Paleontology 22: 871–875.
Makovicky, P.J., and H.-D. Sues. 1998. Anatomy and phylogenetic relationships of the theropod dinosaur
Microvenator celer from the Lower Cretaceous of Montana. American Museum Novitates 3240:
Mikhailov, K.E. 1991. Classication of fossil eggshells of amniotic vertebrates. Acta Palaeontologica
Polonica 36: 193–238.
Mikhailov, K.E. 1994. eropod and protoceratopsian dinosaur eggs from the Cretaceous of Mongolia
and Kazakhstan. Paleontological Journal 28: 101–120.
Mikhailov, K.E. 2014. Eggshell structure, parataxonomy and phylogenetic analysis: some notes on articles
published from 2002 to 2011. Historical Biology 26: 144–154.
Nesbitt, S.J., A.H. Turner, M. Spaulding, J.L. Conrad, and M.A. Norell. 2009. e theropod furcula.
Journal of Morphology 270: 856–879.
Norell, M.A., and P. Makovicky. 1997. Important features of the dromaeosaur skeleton: information from
a new specimen. American Museum Novitates 3215: 1–28.
Norell, M.A., and P. Makovicky. 1999. Important features of the dromaeosaur skeleton II: information from
newly collected specimens of Velociraptor mongoliensis. American Museum Novitates 3282: 1–45.
Norell, M.A., and X. Xu. 2005. Feathered dinosaurs. Annual Reviews of Earth and Planetary Sciences
33: 277–299.
Norell, M.A., et al. 1994. A theropod dinosaur embryo and the anities of the Flaming Clis dinosaur
eggs. Science 266: 779–782.
Norell, M.A., J.M. Clark, L.M. Chiappe, and D. Dashzeveg. 1995. A nesting dinosaur. Nature 378: 774–
Norell, M.A., P.J. Makovicky, and J.M. Clark. 1997. A Velociraptor wishbone. Nature 389: 447.
Norell, M.A., J.M. Clark, and L.M. Chiappe. 2001. An embryonic oviraptorid (Dinosauria, eropoda)
from the Upper Cretaceous of Mongolia. American Museum Novitates 3315: 1–17.
Osborn, H.F. 1924. ree new eropoda, Protoceratops zone, central Mongolia. American Museum
Novitates 144: 1–12.
Osmólska, H. 1981. Coossied tarsometatarsi in theropod dinosaurs and their bearing on the problem
of bird origins. Palaeontologica Polonica 42: 79–95.
Osmólska, H., P.J. Currie, and R. Barsbold. 2004. Oviraptorosauria. In D.B. Weishampel, P. Dodson,
and H. Osmólska (editors), e Dinosauria, 2nd ed.: 165–183. Berkeley: University of California
Persons, W., G. Funston, P.J. Currie, and M.A. Norell. 2015. A possible instance of sexual dimorphism
in the tails of two oviraptorosaur dinosaurs. Nature Scientic Reports 5: 9472. [doi: 10.1038/
Pu, H., et al. 2017. Perinate and eggs of a giant caenagnathid dinosaur from the Late Cretaceous of
central China. Nature Communications 8: 14952.
Sato, T., Y. Cheng, X. Wu, D.K. Zelenitsky, and Y. Hsiao. 2005. A pair of shelled eggs inside a female
dinosaur. Science 308: 375–375.
Schweitzer, M.H., et al. 1999. Beta-keratin specic immunological reactivity in feather-like structures of
the Cretaceous alvarezsaurid, Shuvuuia deserti. Journal of Experimental Zoology B, Molecular and
Developmental Evolution 285:146–157.
Schweitzer, M.H., J.L. Wittmeyer, and J.R. Horner. 2005. Gender-specic reproductive tissue in ratites
and Tyrannosaurus rex. Science 308: 1456–1460.
Truitt, L. 1996. Dinosaur hunters—secrets of the Gobi Desert. National Geographic Films.
Turner, A.H., P.J. Makovicky, and M.A. Norell. 2007. Feather quill knobs in the dinosaur Velociraptor.
Science 317: 1721.
Varricchio, D.J., and D.E. Barta. 2015. Revisiting Sabath’s “larger avian eggs” from the Gobi Cretaceous.
Acta Palaeontologica Polonica 60 (1): 11–25.
Varricchio, D.J , J.R. Horner, and F.D. Jackson. 2002. Embryos and eggs for the Cretaceous theropod
dinosaur Troodon formosus. Journal of Vertebrate Paleontology 22 (3): 564–576.
Varricchio, D.J., et al. 2008. Avian parental care had dinosaur origins. Science 322: 1826–1828.
Wang, S., S. Zhang, C. Sullivan, and X. Xu. 2016. Elongatoolithid eggs containing oviraptorid (erop-
oda, Oviraptorosauria) embryos from the Upper Cretaceous of Southern China. BMC Evolutionary
Biology 16: 1–21.
Webster, D. 1996. Dinosaurs of the Gobi: unearthing a treasure trove. National Geographic 190 (1):
Weishampel, D.B., et al. 2008. New oviraptorid embryos from Bugin-Tsav, Nemegt Formation (Upper
Cretaceous), Mongolia, with insights into their habitat and growth. Journal of Vertebrate Paleontol-
ogy 28: 1110–1119.
Xu, X., and M.A. Norell. 2004. A new troodontid dinosaur with avian-like sleeping-posture from China.
Nature 431: 838–841.
Zanno, L.E., and S.D. Sampson 2005. A new oviraptorosaur (eropoda, Maniraptora) from the Late
Cretaceous (Campanian) of Utah. Journal of Vertebrate Paleontology 25 (4): 897–904.
A A
ac acromion
CL continuous layer
cv cervical vertebra
dpc deltopectoral crest
Dv-1 dorsal vertebra 1
Dv-2 dorsal vertebra 2
egg egg
t bular tubercle
ht humeral tubercle
hyp hypocleidium
lac le astragalo-calcaneum
lf le femur
l le bula
le humerus
lateral humeral rugosity
li le ischium
lil le ilium
lr le radius
ls le scapula
lt le tibia
lu le ulna
mc-2 metacarpal 2
mc-3 metacarpal 3
ML mammillary layer
mp1-1 manual digit1-phalanx 1
mp1-2 manual digit1-phalanx 2
mp2-1 manual digit2-phalanx 1
mp2-2 manual digit2-phalanx 2
mp2-3 manual digit2-phalanx 3
mp3-1 manual digit3-phalanx 1
mp3-2 manual digit3-phalanx 2
mp3-3 manual digit3-phalanx 3
mp3-4 manual digit3-phalanx 4
pp4-1 pedal digit 4- phalanx 1
pp4-2 pedal digit 4- phalanx 2
pp4-3 pedal digit 4- phalanx 3
pp4-4 pedal digit 4- phalanx 4
pp4-5 pedal digit 4- phalanx 5
rac right astragalo-calcaneum
rc right coracoid
rf right femur
r right bula
rh right humerus
ri right ischium
rmt-2 right metatarsal 2
rmt-3 right metatarsal 3
rmt-4 right metatarsal 4
rp right pubis
rr right radius
rs right scapula
rt right tibia
ru right ulna
sl semilunate
stp sternal plate
str sternal rib
unc uncinate process
I A
IGM Institute of Paleontology, Mongolian Academy of Sciences
AMNH FARB American Museum of Natural History, Fossil amphibians, reptiles and birds
... In keeping with the goal of developing a conceptual framework comparable with present-day analogues, vertebrate concentrations in the Upper Cretaceous Gobi beds can be grouped into two distinct assemblages: individuals (or groups of individuals) presenting evidence of direct, permanent burial, and others showing clear reworking and postmortem transportation (Rogers and Kidwell 2007). Clear examples of the first typologies include brooding dinosaurs (Osborn 1924a;Norell et al. 1995;Dong and Currie 1996;Fanti et al. 2012;Norell et al. 2018); undisturbed but isolated nests of eggssometimes in nesting grounds (Andrews 1923;Sabath 1991;Mikhailov et al. 1994); massive ankylosaurs trapped in upright positions (Lefeld 1971;Jerzykiewicz et al. 1993;Currie et al. 2011;Mallon et al. 2018;Jerzykiewicz 2018); dinosaurs mired in mud Lee et al. 2018); smaller dinosaurs in life positions trapped in "escaping" or even "fighting" positions (Figs. 3, 4); and the thousands of footprints documented in the area (Ishigaki 1999;Currie et al. 2003;Ishigaki et al. 2009). The second type includes large bonebeds such as the Saurolophus-dominated Dragon's Tomb (Efremov 1958;Rozhdestvensky 1965;Bell et al. 2018); the isolated skeletons of large sauropods ; and countless other individual skeletons that litter the surface of several key localities (Fig. 5). ...
... Regardless of the rarity of embryos, the Djadokhta depositional environment was alkaline enough to preserve the eggshell, and both eggs and nests were buried before hatching. In some cases, even the brooding mothers were buried on top of the nests (Osborn 1924a;Norell et al. 1995;Dong and Currie 1996;Clark et al. 1999;Norell et al. 2018). ...
Three distinct but overlapping dinosaur-dominated faunas characterize the Upper Cretaceous Djadokhta, Baruungoyot and Nemegt formations of the Nemegt Basin of Mongolia. Documented faunal differences cannot be explained easily by temporal succession, but can be understood in the light of physical processes controlling life, death, and burial of taxa. The stratigraphy of the Gobi Desert region records tectonically driven geometries, clearly documenting preservational processes different than those acting in most other dinosaur-dominated beds worldwide. Small, asymmetric tectonic grabens were filled with Upper Cretaceous, dinosaur bearing deposits showing asymmetric distributions of facies, here termed Lithobiotopes. The water-lain fluvial and alluvial plain facies of the Nemegt Lithobiotope supported and preserved a fauna dominated by gigantic dinosaurs, but had a preservational bias against smaller animals. The Nemegt passed laterally into interdune facies of the Baruungoyot Lithobiotope, which represented a hostile environment for large species, but preserved smaller animals. This in turn passed laterally into the aeolianite facies of the Djadokhta Lithobiotope, which is characterized by remains of small dinosaurs and a rich fauna of other animals. The Nemegt Gobi Basin can be visualized as an oasis with a central pond supplied with water from ephemeral channels and surrounded by a semi-arid alluvial plain and dune fields.
... 3). Partial skeletons of multiple species of similarly small adult oviraptors have been found positioned on traces of their own nests from the Late Cretaceous of Mongolia and China (Osborn, 1924;Norell et al., 1995;Dong and Currie, 1996;Clark et al., 1999;Fanti et al., 2012;Bi and Xu, 2017;Norell et al., 2018). These and other oviraptors left nesting traces defined by eggs in 2-4 stacked near-circular concentric rings ranging 0.6-.8 m in diameter with circular interior openings ranging 0.2-.3 m in diameter centered within the rings and often slightly to significantly raised (Yang et al., 2019). ...
... Without the benefit of footprints, the oviraptor nesting posture has been described as "crouching" (Fanti et al., 2012), "sitting" (Norell et al., 2018) and "oviposition" (Yang et al., 2019). With the abdominal region directly on the center of the nest, with the arms extended laterally and flexed 90° caudally at the elbow along the circumference of the nest, a prone posture is illustrated by Clark et al. (1999, fig. ...
Full-text available
After description of the first known footprint attributed to Tyrannosaurus rex by Lockley and Hunt in 1994, the lead author (TC) began a systematic search for more tracks of this giant theropod in the Raton Basin. During parts of eight field seasons in the mid and late 1990s and early 2000s, two finds were made and interpreted to be tracks of large tyrannosaurids very similar in size to the one that made the original footprint. The material, which is preserved as convex hyporelief, consists of a tridactyl right pes print lacking a hallux impression coming from a locality near Ludlow, Colorado, and a track array, here interpreted as a convex hyporelief left pes print with a single toe impression, likely that of a hallux, and a pair of parallel convex hyporelief forearm prints with partial hand impressions from Cimarron, New Mexico. Near the track array an "L"-shaped impression in convex hyporelief is interpreted to have been the impression of carrion or a prey item. Like the original material of Tyrannosauripus pillmorei, these traces are present in the Upper Cretaceous lower coal zone of the Raton Formation. The track array is hypothesized to have been made by a prone adult T. rex, rising from a quadrupedal position. In standing up, the dinosaur stepped forward and made the left pes print, while at nearly the same time it made the forearm prints as it pushed down with its forearms and wrists to facilitate its rise to a standing, walking, or running posture.
... For example, the scapula and coracoid are tightly sutured together, but not fused, in a juvenile specimen of Velociraptor mongoliensis (MPC-D100/54), and a fused scapulocoracoid is seen in an adult specimen (IGM 100/986) (Hone et al., 2012;Norell and Makovicky, 1999). Complete fusion of the scapulocoracoid, leaving no visible suture, is the usual condition in adult individuals (e.g., Citipati osmolskae IGM 100/1004, Anzu wyliei CM 78 001) (Lamanna et al., 2014;Norell et al., 2018). Among non-ornithothoracine avialans, all known jinguofortisids, and all known confuciusornithiforms except a single subadult Eoconfuciusornis zhengi specimen (IVPP V 11977), possess a fused scapulocoracoid (Wang et al., 2018a;Wu et al., 2021). ...
Full-text available
The morphology of the pectoral girdle, the skeletal structure connecting the wing to the body, is a key determinant of flight capability, but in some respects is poorly known among stem birds. Here, the pectoral girdles of the Early Cretaceous birds Sapeornis and Piscivorenantiornis are reconstructed for the first time based on computed tomography and three-dimensional visualization, revealing key morphological details that are important for our understanding of early flight evolution. Sapeornis exhibits a double articulation system (widely present in non-enantiornithine pennaraptoran theropods including crown birds) which involves, alongside the main scapula-coracoid joint, a small subsidiary joint, though variation exists with respect to the shape and size of the main and subsidiary articular contacts in non-enantiornithine pennaraptorans. This double articulation system contrasts with Piscivorenantiornis in which a spatially restricted scapula-coracoid joint formed by a single set of opposing articular surfaces, a feature also present in other members of Enantiornithines, a major clade of stem birds known only from the Cretaceous. The unique single articulation system may reflect correspondingly unique flight behavior in enantiornithine birds, but this hypothesis requires further investigation from a functional perspective. Our renderings indicate that both Sapeornis and Piscivorenantiornis had a partially closed triosseal canal (a passage for muscle tendon that plays a key role in raising the wing), and our study suggests that this type of triosseal canal occurred in all known non-euornithine birds except Archaeopteryx , representing a transitional stage in flight apparatus evolution before the appearance of a fully closed bony triosseal canal as in modern birds. Our study reveals additional lineage-specific variations in pectoral girdle anatomy, as well as significant modification of the pectoral girdle along the line to crown birds. These modifications produced diverse pectoral girdle morphologies among Mesozoic birds, which allowed a commensurate range of capability levels and styles to emerge during the early evolution of flight.
... Characteristics of the aforementioned egg and eggshell, are indicative of the Elongatoolithidae (Zhao, 1975). This oofamily is common throughout the Cretaceous of Asia, and previous discoveries of embryos or adults associated with elongatoolithid eggs confirm their identity as oviraptorosaur (e.g., Norell et al., 1994;Norell et al., 1995;Dong and Currie, 1996;Weishampel et al., 2008;Wang et al., 2016;Pu et al., 2017;Norell et al., 2018). ...
Full-text available
Despite the discovery of many dinosaur eggs and nests over the past 100 years, articulated in-ovo embryos are remarkably rare. Here we report an exceptionally preserved, articulated oviraptorid embryo inside an elongatoolithid egg, from the Late Cretaceous Hekou Formation of southern China. The head lies ventral to the body, with the feet on either side, and the back curled along the blunt pole of the egg, in a posture previously unrecognized in a non-avian dinosaur, but reminiscent of a late-stage modern bird embryo. Comparison to other late-stage oviraptorid embryos suggests that prehatch oviraptorids developed avian-like postures late in incubation, which in modern birds are related to coordinated embryonic movements associated with tucking — a behavior controlled by the central nervous system, critical for hatching success. We propose that such pre-hatching behavior, previously considered unique to birds, may have originated among non-avian theropods, which can be further investigated with additional discoveries of embryo fossils.
... Similarly, the Two Medicine and Frontier formations in Montana, USA, were influenced by volcanism (Dyman et al., 2008;Foreman et al., 2008), and eggshell from these formations are as dark as Korean and Japanese eggshells (SC, personal observation). In contrast, Upper Cretaceous deposits from southern Mongolian are characterized by the absence of volcanic rocks (Eberth et al., 2009;Eberth, 2018) and no signs of deep burial (Graf et al., 2018), and potentially for these reasons, eggshells are light in color (Fanti et al., 2012;Varricchio and Barta, 2015;Pei et al., 2017;Norell et al., 2018;Montanari, 2018). ...
Raman spectroscopic analyses of thermally altered organic materials can be used to assess the paleothermometry of the sedimentary deposits. Although this technique has been widely applied to diverse microfossils, macroscopic vertebrate fossils have been neglected. In this paper, we show that fossil eggshells can be used for this purpose by demonstrating the paleothermometric potential of diverse amniotic eggshells from the Wido Volcanics (Upper Cretaceous, South Korea) that contain thermally altered organic material. We estimate the maximum paleotemperature recorded in the eggshells using Raman spectroscopic data and the spectrum deconvolution technique, which was invented and developed by organic geochemists. The results show that the same type of eggshells record different paleotemperature gradients depending on their spatial distribution in the fossil locality, whereas different types of eggshells from similar locations show similar paleotemperature gradients (except for one specimen with a peculiar microstructure). These findings are further supported by Fourier transform infrared spectroscopic results, which focus on thermally altered organic materials, and electron backscatter diffraction data, which focus on calcite twinning. This study suggests that fossil eggshells are useful and reliable materials for paleothermometry because thermally altered organic materials and calcites of eggshells provide independent opportunities to assess paleothermometry. We propose that fossil eggshells may be useful for evaluating the thermal evolution of sedimentary basins.
Since their discovery in the 1920s, some asymmetric, elongated dinosaur eggs from the Upper Cretaceous of Mongolia have been interpreted as ceratopsian eggs. However, recent views support a maniraptoran affinity mainly based on macroscopic features. Technical advancements in palaeontology provide a novel approach to diagnose maniraptoran eggs, and the discovery of soft ceratopsian eggs makes the debate a timely issue worth revisiting. Here, we analysed Protoceratopsidovum eggshell from southern Mongolia with electron backscatter diffraction and the results were compared with East Asian and North American maniraptoran and ornithischian eggs. The microstructure and crystallography of Protoceratopsidovum confirms the maniraptoran hypothesis, given that it shows diagnostic features of the clade absent from ornithischian (e.g. hadrosaur) eggs. Additional characters, such as egg shape, ornamentation, egg pairing and clutch structure also support a maniraptoran affinity. Protoceratopsidovum has both elongatoolithid (oviraptorosaur egg) and prismatoolithid (troodontid egg) features, and it may bridge the morphological gap between the two established oofamilies. Considering the similarity between Protoceratopsidovum and potential dromaeosaur eggs, at least some Protoceratopsidovum may be deinonychosaur eggs. Fossil localities that yield both deinonychosaurs and Protoceratopsidovum may yield conclusive evidence for the true affinity of this ootaxon. Future discovery of the egg‐laying taxon of Protoceratopsidovum will not only improve our understanding of maniraptoran reproductive biology but also make Protoceratopsidovum a reliable indicator of the palaeobiogeography of that maniraptoran clade.
Full-text available
Khulsanurus magnificus gen. et sp. nov., is described based on a partial skeleton, including cervical and caudal vertebrae, scapulocoracoids, humerus, and pubis from the Late Cretaceous (Campanian) Barungoyot Formation at Khulsan locality in Gobi Desert, Mongolia. The new taxon differs from other alvarezsaurids by a combination of characters that include cervicals lacking pleurocoels, carotid processes, and epipophyses, dorsoventrally thick and subtriangular in cross-section transverse process of the anterior caudals, prominent infrapostzygapophyseal fossa on the transverse process of anterior caudals, short and mostly anteriorly directed prezygapophyses of anterior caudals, neural arch of anterior caudal lacking the interzygapophyseal ridges and having a prominent dorsal depression around the anterior end of the neural spine. The new taxon retains a number of plesiomorphic traits: slightly convex posterior centrum condyle of anterior caudals, prominent anterior, and posterior bumps at the neurocentral junction in anterior caudals, neural arch of anterior caudals extending for the entire length of the centrum, and pubic foot and apron. Khulsanurus shares with Shuvuuia the deltopectoral crest of humerus that is continuous with the humeral head. The phylogenetic analysis placed Khulsanurus in the Parvicursorinae in a polytomy with Mononykus, Shuvuuia, Albinykus, and Xixianykus.
Pelecanimimus polyodon was discovered in 1993 in the Spanish Barremian fossil site of Las Hoyas, being the first ornithomimosaur described from Europe. So far, there has been no detailed description of the holotype of Pelecanimimus, which is composed of the anterior-half of an articulated skeleton. Here we report a new, detailed, revised and more accurate osteological description of its postcranial skeleton, comparing this new data to information about Ornithomimosauria from the last three decades. This osteological and phylogenetic analysis of Pelecanimimus shows several ornithomimosaur synapomorphies and a unique combination of characters that emend its original diagnosis. Pelecanimimus diverged early in Ornithomimosauria and reveals an enlargement trend of the manus, shared with derived ornithomimosaurians, due to a long metacarpal I and elongated distal phalanges. This evolutionary novelty, and other synapomorphies, has led to the definition of a new clade, Macrocheiriformes, including Pelecanimimus and more derived ornithomimosaurs. Pelecanimimus has the only ossified sternal plates among ornithomimosaurs and the first evidence of uncinate processes in a nonmaniraptoran theropod, indicating a convergent appearance of these structures in Coelurosauria. The character combination in an early-diverging ornithomimosaur like Pelecanimimus found in this analysis provides a key step in the evolution of the manus and pectoral girdle in Ornithomimosauria
Full-text available
The abundance of dinosaur eggs in Upper Cretaceous strata of Henan Province, China led to the collection and export of countless such fossils. One of these specimens, recently repatriated to China, is a partial clutch of large dinosaur eggs (Macroelongatoolithus) with a closely associated small theropod skeleton. Here we identify the specimen as an embryo and eggs of a new, large caenagnathid oviraptorosaur, Beibeilong sinensis. This specimen is the first known association between skeletal remains and eggs of caenagnathids. Caenagnathids and oviraptorids share similarities in their eggs and clutches, although the eggs of Beibeilong are significantly larger than those of oviraptorids and indicate an adult body size comparable to a gigantic caenagnathid. An abundance of Macroelongatoolithus eggs reported from Asia and North America contrasts with the dearth of giant caenagnathid skeletal remains. Regardless, the large caenagnathid-Macroelongatoolithus association revealed here suggests these dinosaurs were relatively common during the early Late Cretaceous.
Full-text available
A monodominant bonebed of Avimimus from the Nemegt Formation of Mongolia is the first oviraptorosaur bonebed described and the only recorded maniraptoran bonebed from the Late Cretaceous. Cranial elements recovered from the bonebed provide insights on the anatomy of the facial region, which was formerly unknown in Avimimus. Both adult and subadult material was recovered from the bonebed, but small juveniles are underrepresented. The taphonomic and sedimentological evidence suggests that the Avimimus bonebed represents a perimortem gregarious assemblage. The near absence of juveniles in the bonebed may be evidence of a transient age-segregated herd or ‘flock’, but the behaviour responsible for this assemblage is unclear. Regardless, the Avimimus bonebed is the first evidence of gregarious behaviour in oviraptorosaurs, and highlights a potential trend of increasing gregariousness in dinosaurs towards the end of the Mesozoic.
Full-text available
Our understanding of caenagnathids has benefited from recent discoveries, including nearly complete skeletons from the Hell Creek Formation of Montana. However, their phylogenetic relationships remain unclear. A new specimen from the Horseshoe Canyon Formation of Alberta has implications for the phylogeny and paleobiology of these creatures. The partial skeleton is articulated and includes a mandible, a full cervical and dorsal series of vertebrae, a right pectoral girdle and arm, a sternum, gastralia, a partial ilium, and a partial hind limb. The mandible is edentulous and the articular ridge is intermediate in form between Caenagnathus collinsi and Chirostenotes pergracilis. The neck is long and composed of at least 11 well-pneumatized cervical vertebrae with fused cervical ribs. The dorsal ribs have finger-like uncinate processes dissimilar in shape to those of other oviraptorosaurs. The pectoral girdle is large and typically maniraptoran, except that the glenoid of the scapulocoracoid faces laterally instead of posteroventrally. The arm is well muscled and can be interpreted to have been a pennibrachium, as indicated by ulnar papillae on the ulna. The manus is characterized by a short first metacarpal but an elongate phalanx I-1 and oviraptorid-like phalangeal proportions in the second digit. These and other features indicate that the specimen represents a new taxon, Apatoraptor pennatus, gen. et sp. nov. Phylogenetic analysis resolves the complicated relationships of Caenagnathidae and allows the evolution of display features to be traced throughout Oviraptorosauria. SUPPLEMENTAL DATA—Supplemental materials are available for this article for free at Citation for this article: Funston, G. F., and P. J. Currie. 2016. A new caenagnathid (Dinosauria: Oviraptorosauria) from the Horseshoe Canyon Formation of Alberta, Canada, and a reevaluation of the relationships of Caenagnathidae. Journal of Vertebrate Paleontology. DOI: 10.1080/02724634.2016.1160910.
Full-text available
Oviraptorids, like many other dinosaurs, clearly had a complex pattern of skeletal growth involving numerous morphological changes. However, many ontogenetic skeletal changes in oviraptorids were previously unclear due to the lack of well preserved specimens that represent very young developmental stages. Here we report three elongatoolithid dinosaur eggs from the Upper Cretaceous Nanxiong Formation of Nankang District, Ganzhou City, Jiangxi Province, China that contain in ovo embryonic skeletons. The eggs themselves show diagnostic features of the oofamily Elongatoolithidae, whereas the embryos are identified as taxonomically indeterminate oviraptorids. The three new specimens display pathological eggshell features, including double-layered and multilayered cones in the columnar layer, which probably result from high levels of pathogenic trace elements in the environment. Nevertheless, the skeletons of the preserved embryos exhibit no structural or histological abnormalities. Comparisons between the new embryos and other oviraptorid specimens reveal 20 osteological features that appear to change substantially during ontogeny in oviraptorids. For example, the dorsoventral height of the skull increases more rapidly than the anteroposterior length during oviraptorid ontogeny, and the initially paired nasals fuse at an early stage, presumably facilitating growth of a crest. The new specimens represent the first known oviraptorid embryos associated with pathological eggshells. The absence of structural and histological abnormalities indicates the environmental factor that led to the eggshell pathologies did not affect the skeletal development of the oviraptorids themselves. As in tyrannosaurids, but in contrast to the situation in other maniraptorans, the oviraptorid skull becomes proportionally dorsoventrally deeper during ontogeny. Although oviraptorids and therizinosauroids occupy broadly the same grade of maniraptoran evolution, the embryonic ossification patterns of the vertebral column and furcular hypocleidium appear to differ significantly between the two clades. The limb proportions of juvenile oviraptorids indicate that they were bipedal, like adults. Oviraptorids may have differed greatly from therizinosauroids in their growth trajectories and locomotor modes during early post-hatching ontogeny, essentially occupying a different ecological niche.
Full-text available
The first parataxonomic description of a morphological group of elongated eggs of protoceratopsian and theropod dinosaurs from the Cretaceous of Mongolia and Kazakhstan (four genera and twelve species of the families Prismatoolithidae and Elongatoolithidae) is given. The history of the finds, their taphonomy, paleoecology, stratigraphy and geographic distribution are also presented. -Journal summary
The abundance of dinosaur eggs in Upper Cretaceous strata of Henan Province, China led to the collection and export of countless such fossils. One of these specimens, recently repatriated to China, is a partial clutch of large dinosaur eggs (Macroelongatoolithus) with a closely associated small theropod skeleton. Here we identify the specimen as an embryo and eggs of a new, large caenagnathid oviraptorosaur, Beibeilong sinensis. This specimen is the first known association between skeletal remains and eggs of caenagnathids. Caenagnathids and oviraptorids share similarities in their eggs and clutches, although the eggs of Beibeilong are significantly larger than those of oviraptorids and indicate an adult body size comparable to a gigantic caenagnathid. An abundance of Macroelongatoolithus eggs reported from Asia and North America contrasts with the dearth of giant caenagnathid skeletal remains. Regardless, the large caenagnathid-Macroelongatoolithus association revealed here suggests these dinosaurs were relatively common during the early Late Cretaceous.
Elongate and asymmetric eggs of the oospecies Prismatoolithus levis occur regularly in the Upper Cretaceous Two Medicine Formation of western Montana. These eggs had previously been assigned to the ornithischian Orodromeus makelai, for both juvenile and adult remains are typically associated with these eggs. Reexamination of the embryos shows them to exhibit at least 24 apomorphies of the clades Dinosauria, Theropoda and Paraves. The embryos also display a pneumatic quadrate, closely placed basal tubera, a high tooth count, a metatarsal II much narrower than IV and a strongly constricted metatarsal III, all possible synapomorphies of the Troodontidae. Presence of large basal tubera and a broadly rounded anterior border of the maxillary fenestra permit assignment to Troodon formosus. Most but not all bones appear ossified, suggesting a developmental level comparable to stages 35–38 of avian embryos and a time approaching hatching. Embryos show a consistent level of development from one egg to another indicating synchronous hatching of the clutch. Embryonic Troodon exhibit long distal segments and radically different hindlimb proportions in comparison to adults. Orodromeus and other small vertebrate remains associated with Troodon egg horizons may represent prey of the adults during egg-laying and brooding. Troodon eggs show several aspects either shared or convergent with some birds, and further demonstrate the close relationship of Troodontidae and Aves. These features include: asymmetric egg form, non-branching angusticanaliculate pores, distinct structural differentiation of the mammillary and overlying prismatic layer, barrel-shaped mammillary cones with a blocky calcite cleavage, and prismatic structure visible throughout the second structural layer.
Background: Oviraptorids, like many other dinosaurs, clearly had a complex pattern of skeletal growth involving numerous morphological changes. However, many ontogenetic skeletal changes in oviraptorids were previously unclear due to the lack of well preserved specimens that represent very young developmental stages.