See discussions, stats, and author profiles for this publication at: https://www.researchgate.

net/publication/234115857

The “Tustea Puzzle”: Hadrosaurid (Dinosauria, Ornithopoda) Hatchlings

Associated with Megaloolithidae Eggs in the Maastrichtian of the Hateg Basin
(Romania)

Article in AMEGHINIANA · March 2010

DOI: 10.5710/AMGH.v47i1.9

CITATIONS READS

18 390

1 author:

Dan Grigorescu
University of Bucharest
134 PUBLICATIONS 4,193 CITATIONS

SEE PROFILE

Some of the authors of this publication are also working on these related projects:

Biomineralization through SEM analysis on dinosaurs and birds eggshells. View project

Hateg Country dinosaurs geopark View project

All content following this page was uploaded by Dan Grigorescu on 21 May 2014.

The user has requested enhancement of the downloaded file.

AMEGHINIANA (Rev. Asoc. Paleontol. Argent.) - 47 (1): 000-000. Buenos Aires, 30-03-2010 ISSN 0002-7014

The “Tustea Puzzle”: hadrosaurid (Dinosauria,

Ornithopoda) hatchlings associated with Megaloolithidae
eggs in the Maastrichtian of the Hateg Basin (Romania)

Dan GRIGORESCU1

Abstract. Since 1991 when the coinciding occurrence of megaloolithid eggs and hatchlings of the hadrosaurid
Telmatosaurus transsylvanicus was suggested by few limb bone remains found close to the first discovered eggs
in the Hateg Basin, tens of eggs grouped in clutches together with neonate bones were unearthed within the
place that become “Tustea nesting site”. In the meantime the original vertical escarpment near the village of
Tustea was leveled and converted into a horizontal platform. By their external characters and eggshell mi-
crostructure all the eggs belong to the Megaloolithus oogenus, closer to M. siruguei; similarly, all the baby re-
mains whose state of preservation allows their taxonomic assignment belong to Telmatosaurus transsylvanicus,
one of the common dinosaur species from the rich Maastrichtian faunal assemblage of the Hateg Basin
(Romania), and the basalmost hadrosaurid. Most of the baby remains consist of isolated bones and teeth, but
partial skeletons with articulated elements were also found. The preliminary ontogenetic study reveals a
rapid growth of the hatchlings, a fact also supported by the discovery of most of the skeletal remains outside
the egg clutches, but in their close vicinity. In few cases bones were found within the egg clutches and even
inside the egg. It is worth to mention that no sauropod remains were found within the nesting horizon from
Tustea.

Resumen. EL “ENIGMA DE TUSTEA”: PICHONES DE HADROSÁURIDOS (DINOSAURIA, ORNITHOPODA) ASOCIADOS CON

HUEVOS MEGALOOLITHIDAE EN EL MAASTRICHTIANO DE LA CUENCA HATEG (RUMANIA). Desde 1991, cuando la co-
incidencia de la aparición de huevos megaloolítidos y pichones del hadrosáurido Telmatosaurus transsyl-
vanicus fue sugerida sobre la base de unos pocos huesos apendiculares encontrados en proximidad a los
primeros huevos descubiertos en la Cuenca Hateg, decenas de agrupaciones de huevos junto con huesos de
neonatos fueron desenterrados del sitio que pasó a llamarse “sitio de nidificación Tustea”. Mientras tanto, el
escarpado vertical original próximo a la Villa de Tustea fue nivelado y convertido en una plataforma hori-
zontal. Por sus caracteres externos y microestructura de cáscara todos los huevos pertenecen al género
Megaloolithus, cercano a M. siruguei; del mismo modo, todos los restos de pichones cuyo estado de preser-
vación permite su asignación taxonómica pertenecen a Telmatosaurus transsylvanicus, una de las especies co-
munes de dinosaurios de la rica asociación faunística del Maastrichtiano de la Cuenca Hateg (Rumania), y el
hadrosáurido más basal. La mayoría de los restos de pichones consiste en huesos y dientes aislados, aunque
esqueletos parciales con elementos articulados fueron también encontrados. El estudio ontogenético prelimi-
nar revela un rápido crecimiento de los pichones, un hecho también soportado por el descubrimiento de una
mayoría de restos esqueletales fuera de las agrupaciones de huevos, pero en su proximidad. En unos pocos
casos fueron encontrados dentro de las agrupaciones de huevos, y aún en el interior de los huevos. Es im-
portante destacar que no se han hallado restos de saurópodos dentro del horizonte de nidificación de Tustea.

Key words. Dinosaur eggs. Megaloolithidae. Hatchlings. Hadrosaurids. Telmatosaurus. Maastrichtian. Hateg Basin.
Romania.
Palabras clave. Huevos de dinosaurios. Megaloolithidae. Pichones. Hadrosáuridos. Telmatosaurus. Maastrichtiano.
Cuenca Hateg. Rumania.

Introduction by Franz Nopcsa. The eggs disposed in clutches,

The first dinosaur eggs and egg remains from
Romania were found in 1988, ninety years after the
first dinosaur bones were recorded in the Uppermost
Cretaceous continental deposits of the Hateg Basin
1University of Bucharest, Faculty of Geology and Geophysics,
Laboratory of Paleontology, 1 Blvd. Nicolae Balcescu, 010041
Bucharest, Romania, dangrig@geo.edu.ro
©Asociación Paleontológica Argentina
were found in a vertical escarpment, near the village
of Tustea, in the north-western part of the basin
(figure 1.2). The egg and eggshell characters corre-
spond to the Megaloolithidae, the most widespread
oofamily during the Late Cretaceous (mostly in the
Campanian and Maastrichtian), with a large number
of oospecies found in South America, Europe and
Asia.
In southern France, where no less than eight
megaloolithid oospecies were described (Garcia and
AMGHB2-0002-7014/10$00.00+.50
2 D. Grigorescu

Figure 1. 1, Location of the Hateg Basin in Romania; 2, Geological map of the western part of the Hateg Basin (F, the Tustea nesting site);
3, Lithostratigraphic column (in meters) from Tustea and the facies interpretation / 1, Ubicación de la Cuenca Hateg en Rumania; 2, Mapa
geológico de la parte oeste de la Cuenca Hateg (F, el sitio de nidificación Tustea); 3, Columna litoestratigráfica (en metros) de Tustea y la interpretación
de las facies. Ch, channel deposits; Cs, crevasse deposits, Fr, reddish fines (paleosoil with calcrete concretions).

Vianey-Liaud, 2001) this type of egg was tentatively
assigned to the titanosaurid sauropod Hypselosaurus
Matheron, 1869 based on some occasional co-occur-
rences of the eggs with adult bone remains. This re-
ferral not consistently proved since nowhere in
France or elsewhere in Europe embryo or hatchling
remains were found closely linked with mega-
loolithid eggs. The assignment remained questiona-
ble, until 1998, when the discovery from Auca
Mahuevo in Argentina of numerous megaloolithid
eggs containing titanosaurid sauropod embryonic re-
mains was announced (Chiappe et al., 1998).
The close similarity of the eggs from Tustea with
the French ones, in both macroscopic and microscop-
ic aspects, made us to presume at the beginning that
the Romanian eggs were laid by Magyarosaurus
Huene, 1932 the only sauropod genus known at that
time in the dinosaur assemblage of the Hateg Basin,
without excluding the possibility that other di-
nosaurs, including the hadrosaur Telmatosaurus
Nopcsa, 1900 might have laid the eggs (Grigorescu et
al., 1990). Soon after that a few hatchling bones, un-
doubtedly belonging to the hadrosaur Telmatosaurus
transsylvanicus Nopcsa, 1900 were found closely as-
sociated to the first discovered egg clutches in
Tustea.
The first announcement on the connection of
megaloolithid eggs and hadrosaurid embryonic and
AMEGHINIANA 47 (1), 2010
hatchling skeletal remains from Tustea was made in
few successive articles (Weishampel et al., 1991;
Grigorescu, 1993; Grigorescu et al., 1994). The fact
was (and still is) regarded with natural suspicion.
This is well expressed in the Summary of the
“Dinosaur eggs and babies” book, edited by
Carpenter, Hirsch and Horner in 1994, that includes
one of the above-quoted articles: “This conflict (sug-
gested by the Tustea case) between identity of the
embryonic remains and the shell microstructure
could be a simple misidentification of the eggs as
hadrosaur, or it could reveal that the diversity of
eggshell type in the Hadrosauria is much more di-
verse than hitherto realized” (Carpenter et al., 1994,
pp. 367, 369).
The number of neonatal and embryo remains in-
creased after the original vertical outcrop from
Tustea was converted into a horizontal platform by
the help of a bulldozer. The searches made on this
platform revealed more than a dozen of new egg
clutches close to which and in few cases inside
whom hatchling bones were found. All the bones
whose stage of preservation allow the taxonomic as-
signment belong to Telmatosaurus, while no
sauropods, babies or adults, were found in the nest-
ing site as well as in the entire geological section
from Tustea.
The case of Tustea highlights once again the prob-
 Tustea Puzzle: hadrosaur hatchlings along megaloolithid eggs
lem of identity of the egg-laying dinosaur taxa based
on egg morphology and eggshell microstructure.
The purpose of this paper is to review the evi-
dences of this intriguing situation that can be named
“the Tustea puzzle”.
Stratigraphical setting
The sequence that include the Tustea nesting lev-
el belongs to the middle member of the Densus-Ciula
Formation (D-C F), that outcrops in the north-west-
ern part of the Hateg Basin, a subsiding intermoun-
tain basin, formed in connection with the main
thrusting phase of the Laramian tectogeny within the
Southern Carpathians (figure 1.2). The D-C F repre-
sents continental molasse deposits and includes a
wide range of siliciclastic rocks, from conglomerates
to mudstones. The deposits were accumulated in a
braided-river system, concomitantly to volcanic
eruptions, whose fine and coarser products are inti-
mately associated with the terrigenous elements. The
regional stratigraphic context indicate a Maas-
trichtian age for the lower and middle members of
the D-C F (Grigorescu and Melinte, 2002) that rough-
ly agrees with the conclusion based on the palyno-
logic content of the deposits (Antonescu et al., 1983).
The egg clutches are located in a red, mica rich
mudstone, 4 m. thick that is divided by a thin layer of
greenish grey sand (figure 1.3). The sedimentologic
environment of the mudstone is a well drained flood-
plain, subjected to pedogenetic processes (Gri-
gorescu and Csiki, 2002). All the clutches correspond
to a single incubation horizon, located at less than a
meter below the uneven base of a thick grey-green-
ish, matrix-supported conglomerate bed.
The overall paleontological content of the red
mudstones includes invertebrates, mostly fresh-wa-
ter and terrestrial gastropods and more abundant
and diverse vertebrates. The vertebrates are found as
disarticulated bones or as small fragments in mi-
cropaleontologic samples, attributed to discoglossid
frogs, albanerpetontids, lizards, crocodilians, turtles,
pterosaurs, theropod and ornithopod dinosaurs and
multituberculate mammals. It is worth to mention
that sauropods, either adult or young are absent
from the entire section at Tustea; the only dinosaur
from which neonatal remains were recorded being
the hadrosaur Telmatosaurus.
In regards of the eggshells, except the mega-
loolithid type, characteristic of the egg-laying hori-
zon, only rare and small fragments of geckoid and
possible bird eggshells were found at different levels
of the mudstone bed.
The taphonomic features of the elements indicate
the mixture within the fossil assemblage of au-
3
tochthonous (floodplain dwelling taxa) with al-
lochthonous ones, whose remains, more fragmented,
were brought into the floodplain episodes together
with the sediments during flood events.
Eggs and nests
The first collection of eggs made in 1988 in the
vertical escarpment from Tustea includes 14 egg re-
mains, most of them preserving only parts of the
lower halves, more or less fragmented and crushed
due to the burial. All, except one egg that is almost
complete with the upper part preserved, seem to
have been hatched. The collection increased consis-
tently after the area was leveled and a horizontal
platform was created allowing systematic excava-
tion. On a rather small surface of only 180 m2, 18 egg
clutches including 2 to 10 eggs more or less frag-
mented, were excavated up to date; the total number
of eggs is around 90 (figure 2).
The individual eggs external characters and inter-
nal microstructure of the eggshell indicate they be-
long to the Megaloolithidae oofamily. The eggs are
sub-spherical, the larger diameter ranging between
14 and 16 cm (figure 3.1). The calculated eggs volume
varies between 1000 and 1500 ml. The eggshell thick-
ness ranges from 2.1 to 2.6 mm, although it typically
has 2.3-2.4 mm thickness. The outer surface of the
shell is covered by closely packed, rounded tuber-
cles, variable in size, from 0.4 to 1.1 mm with a mean
diameter, calculated on eggshell areas of different
eggs, of 0.78 mm. The tubercles may coalesce in small
chains, but usually they are separated by narrow
spaces, some of these occupied by the pore openings.
The internal surface of the eggshell displays a hiero-
glyph pattern given by the coalesced bases of the
growth units, encrusted by secondary calcite. Some
eggshells show internal cratering of the mammillary
knobs, in some other cases the cratering is missing.
The eggshell microstructure is discretispherulitic;
in radial section the eggshell is composed of fan-
shaped units with variable widths, often individual-
ized and delimited by well-defined margins (figure
3.2). The fine growth lines, arched upwardly cross
continuously among adjacent bundles. The herring-
bone pattern is sometimes present. The analysis in
cathodoluminescence reveals that the eggshell mi-
crostructure is not deteriorated by diagenetic
changes.
The pore pattern corresponds to the tubocanalicu-
late system, consisting of a complicated network of
sinuous tubes, with branches and anastomosis
among them. The diameter along the pore length
varies between 0.13 and 0.38 mm. The pore geometry
and distribution was studied through seriated thin
AMEGHINIANA 47 (1), 2010
4 D. Grigorescu

Figure 2. Distribution of egg clutches in Tustea nesting horizon. The baby remains are indicated by stars. (The numbers indicate the
chronology of discoveries.) / distribución de las agrupaciones de huevos en el horizonte de nidificación de Tustea. Los restos de pichones son in-
dicados por estrellas. (Los números indican la cronología de los descubrimientos.)

sections across the entire eggshell thickness, the suc-
cessive sections being spaced at approximately 0.2
mm interval. The study revealed a high pore density,
expressed by ca.100 pores/cm² at the surface; the to-
tal area occupied by pores representing 3.7 % of the
total eggs surface. From all the known Mega-
loolithidae species the eggs from Tustea seems closer
to Megaloolithus siruguei Vianey-Liaud, 1994 from
Southern France (in Vianey-Liaud et al., 1994).
More recently, 11 clutches with approximately 40
eggs, in all regards very similar with the Tustea ones
and closely resembling Megalooithus siruguei were
discovered in superposed levels of grayish and red-
dish calcretic paleosoils on the Raul Mare valley,
near the Nalat Vad village, southwards from Tustea
(Codrea et al., 2002).
Nesting pattern and incubation environment
In most of the cases the number of eggs in a clutch
varies from 3 to 7, rarely clutches with more or less
eggs were unearthed; only one egg was found isolat-
ed. The largest clutches include respectively 10 and
13 eggs (nrs. 12 and 20 in figure 2), disposed in two
parallel rows (clutch nr. 12), or randomly (clutch nr
20). Most of the eggs in the two clutches were obvi-
ously hatched, preserving only their bottom halves,
or only large eggshell fragments. Tree eggs in the
clutch nr.12 are almost completely preserved, with
slightly deteriorated upper caps, however the CT
AMEGHINIANA 47 (1), 2010
scans did not indicate embryonic material inside. The
eggs are filled with red mudstone material, identical
with that of the encasing rock.
The eggs within the clutches are more randomly
than regularly disposed. Sometimes the eggs are lin-
early aligned, in straight or slightly curved rows.
More frequently they are grouped in clusters with a
random disposal of the eggs. As Cousin has men-
tioned in France, the random position of eggs in
clutches might represent a regroupement during or
after the hatching of the eggs that originally were dis-
posed in rows (Cousin, 2002). The eggs in the clutch-
es are closely packed; sometime the neighbor eggs
are tangent or overlap by their margin. The slightly
uneven position of the eggs in the clutches suggests
that they were laid either in the existing holes on the
ground or the mothers dug such superficial holes for
laying the eggs. The distance among the neighbor
clusters varies from 0.5 to 3 m.
The clutches seem to represent original nests not
much affected after hatching, except the crushing
and flattening produced by the pressure and com-
paction of the surrounding sediments.
Based on sedimentologic, clay mineralogy and
stable isotope analysis (Bojar et al., 2005) the environ-
ment of incubation was a humid one that confirms
the conclusion drawn a decade ago based on the high
pore density and pore geometry of the eggs
(Grigorescu et al., 1994). The eggs were laid within a
fine-grained substrate of paleosol, formed in the dis-
tal part of a well-drained floodplain, well vegetated
 Tustea Puzzle: hadrosaur hatchlings along megaloolithid eggs 5

Figure 3. Megaloolithid eggs and Telmatosaurus transsylvanicus babies from Tustea. 1, Individual egg (larger diameter 16 cm); 2, Radial
thin section of eggshell in polarized light showing fan-shaped units and weathered mammillary zone; 3, Left femora of a hatchling- (left)
(FGGUB R 1850) and of an adult (right) (BMNH R 4914) (scale bar=2 cm); 4, Right distal femur of a hatchling (FGGUB 248) in lateral,
cranial and ventral views (from left to right) (scale bar=1 cm); 5, Right dentary tooth of a hatchling found isolated (FGGUB R 2089) (scale
bar=1 mm); 6, Right dentary tooth found within an egg matrix together with incompletely ossified embryonic skeletal remains of a near-
term embryo (FGGUB R 2091) (scale bar=0.5 mm); 7, Fragment of a left dentary (FGGUB R 1850) (scale bar=1 mm) / huevos megaloolíti-
dos y pichones de Telmatosaurus transsylvanicus provenientes de Tustea. 1, Huevo individual (diámetro mayor 16 cm); 2, Sección delgada radi-
al de cáscara de huevo vista con luz polarizada mostrando las unidades en abanico y la zona mamilar erosionada; 3, Fémures derechos de un pichón
(izquierda) (FGGUB R 1850) y de un adulto (derecha) (BMNH R 4914) (Barra de escala=2 cm); 4, Porción distal de fémur derecho of un pichón (FG-
GUB 248) en vistas lateral, craneal y ventral (de izquierda a derecha) (Barra de escala=1 cm); 5, Diente dentario derecho aislado de un pichón (FG-
GUB R 2089) (Barra de escala=1 mm); 6, Diente dentario derecho hallado dentro de la matriz de un huevo junto con restos esqueletales incompleta-
mente osificados de un embrión ya maduro (FGGUB R 2091) (Barra de escala=0.5 mm); 7, Fragmento de un dentario izquierdo (FGGUB R 1850)
(Barra de escala=1 mm) .

by small swampy plants and bioturbated. The incu-
bation took place concomitantly to the evolution of
pedogenetic processes on the ground, which explains
the location of the nesting level in a calcrete rich pa-
leosol. The calcrete nodules are frequently found as-
sociated with the bottom portions of the eggs.
The rarity of completely crushed eggs within the
clutches and generally, the small quantity of isolated
eggshell fragments in the nest indicate that the babies
did not remain for a long time in the nest, a fact that
is sustained by the position of the hatchling remains
in the nesting horizon (see below).
Hatchlings and embryos
Two distal parts of femora, two proximal and one
distal tibiae, the distal half of a fibula were the only
AMEGHINIANA 47 (1), 2010
6 D. Grigorescu

Figure 4. Skeletal remains of a hatchling with bones in almost articulated position (FGGUB R 2088) / restos esqueletales de pichones con
huesos en posición casi articulada. pg, pelvic girdle; fe, femur; ti, tibia; ve, vertebra.

neonate remains found in the original, vertical es-
carpment from Tustea within the level which provid-
ed the first egg clutches, at about half meter laterally
from one of these clutches.
Tens of new neonatal and possible embryonic
bones were found within the Tustea egg horizon af-
ter the vertical section was converted into a horizon-
tal platform. The distribution of the baby remains
within the incubation horizon is shown by small
stars in figure 2. Most of the remains were found, as
in the case of the first discovery, outside the egg
clutches, but very close to them, usually within one
meter of the eggs; in a single case the bones were
found at more than three meters from the nearest
clutch. Rarely baby bones were found within egg
clutches and in a single case a dentary tooth was re-
covered from within the matrix of a broken egg, to-
gether with very small, porous, apparently repre-
senting the stage of replacement of the embryonic
cartilage by bone tissue. Most of the remains consist
in isolated bones and teeth, more or less completely
preserved. They include elements from all skeletal
regions: skull, vertebrae, ribs, girdles and limbs.
Beside the numerous isolated elements, three par-
AMEGHINIANA 47 (1), 2010
tial skeletons of grown hatchlings, with closely asso-
ciated bones from the same skeleton region (figure 4).
The diagnostic characters shown by all the elements
are definitely not of a sauropod but of the
hadrosaurid Telmatosaurus transsylvanicus (Weisham-
pel et al., 1991; Grigorescu et al., 1994), all the skeletal
remains, including dentition and individual teeth
supporting this assignment.
Telmatosaurus was the first dinosaur from the
Hateg assemblage described by Nopcsa (1900) under
the name of Limnosaurus. Telmatosaurus was re-
viewed by Weishampel et al. (1993) and interpreted
as the basalmost hadrosaur, which lacked several
apomorphic features of the other, more derived
members of Hadrosauridae. The study includes ref-
erences to the first hatchling remains, femora and tib-
iae, from Tustea that were compared with adult
bones from different collections. As in adults of
Telmatosaurus, the femur (figure 3.4) has well formed
distal condyles that start to contact each other cra-
nially, enclosing the tunnel-like intercondylar
groove; also, as in adults, the caudal intercondylar
groove is very deep and includes a small condylid on
the lateral condyle. Similar to the adult condition, the
 Tustea Puzzle: hadrosaur hatchlings along megaloolithid eggs 7
Table 1. Measurements of selected Telmatosaurus limb bones. * estimated measurements; ** dimensions as preserved / medidas de hue-
sos apendiculares selectos de Telmatosaurus. * medidas estimadas; ** dimensiones de la parte preservada. L, length; PW, DW, MW, prox-
imal, distal, medial width. 1, additional measurements: deltopectoral crest length, in humerus; distance of 4th trochanter from proximal
end, in femur.

Element Collection no. L PW DW MW 1 Ontogenetic status

Humerus

left FGGUB R.1980.1 28.58 8.34 6.22 3.38 11.16 Hatchling

left FGGUB R.1852.1 23.04* 6.38 5.04 2.99 - Newborn?

Ulna

right FGGUB R.1851.2 36.11 6.12 5.25 2.74 - Hatchling

left FGGUB R.1980.2 27.51 5.08 4.55 2.2 - Hatchling

left FGGUB R.1850.2 14.42** 5.57 - 2.32 - Hatchling

right FGGUB R.1852.2 11.27** 4.74 - 2.13 - Newborn?

Femur

right FGGUB R.248 50.27 - 11.56 6.07 - Hatchling

left FGGUB R.1850.1 48.11 12.31 11.22 6.93 24.98 Hatchling

left FGGUB R.1980.3 46.12* - 9.48 5.2 25.39 Hatchling

right FGGUB R.1980.4 43.63* - 9.39 5.45 23.73 Hatchling

right FGGUB R.1981.1 43.3 - - 4.92 24.39 Hatchling

right FGGUB R.1852.3 35.38 - 6 4 - Newborn?

Tibia

right FGGUB R.1853 63.4 17.73 16.65 5.9 - Large baby

left FGGUB R.246+250 41.5* 10.77 8.92 3.85 - Hatchling

right FGGUB R.1982.1 41.05* - 12.03 3.68 - Hatchling

left FGGUB R.1851.1 38.72 10.53 10.37 3.81 - Hatchling

right FGGUB R.1979 35.78 10.9 8.97 3.73 - Hatchling

left FGGUB R.1852.4 33.6* - 7.38 3.06 - Newborn?

fourth trochanter is positioned towards the middle of The size and stage of preservation differ slightly
the shaft, it is blade shaped, prominent, well-extend- among the skeletal remains, even in the case of ele-
ed from the shaft and slightly pendent at its tip. ments found associated in the same place, indicating
AMEGHINIANA 47 (1), 2010
8 D. Grigorescu
minor differences in age of the hatchlings or near-
term embryos. For example, in the case of the first
discovered remains, which include two distal femo-
ra, two proximal and one distal tibiae, the surface
texture, the preservation of articular surfaces and the
size of the bone fragments indicate that they come
from distinct individuals, with slightly different
stages of ontogenetic development. The best pre-
served femur (Faculty of Geology and Geophysics of
the University of Bucharest -FGGUB R.248) (figure
3.4), a proximal (FGGUB R.250) and the distal tibia
(FGGUB R.246), both, apparently from the same in-
dividual represent hatchling elements, while the oth-
er femur (FGGUB R.249) and tibia (FGGUB R.245)
seem to come from near-term embryos or early
hatchlings.
Generally, the bone remains which display a very
porous surface texture and, in the case of the appen-
dicular elements, lack ossified articular ends are in-
terpreted as near-term embryos or early hatchlings,
while the limb bones of older hatchlings are indicat-
ed by well developed articular morphology, as well
as by the presence of an external cortical layer that
gives a shiny appearance to the bone surface.
The hatchling limb bones that are more numerous
have allowed a preliminary ontogenetic study based
on comparative analysis of hatchling limb bones and
adult bones (Grigorescu and Csiki, 2006). Besides this
close similarity in shape and proportions between
hatchlings, adults and sub-adults of Telmatosaurus
transsylvanicus (figure 3.3) the study highlighted
some allometric changes that occurred during
growth in the appendicular skeleton of Tematosaurus.
This study also suggested a high growth rate during
early post-natal development. The measurements of
limb bones coming from different hatchling or possi-
ble newborn individuals are shown in table 1.
The dentition of hatchlings and embryos also re-
vealed close similarity with adult Telmatosaurus
transsylvanicus. Few individual dentary teeth were
found isolated (figure 3.5), one of these within an egg
matrix (figure 3.6). The apex of the teeth is pointed as
in the adults, but strait or very slightly recurved dis-
tally; the margin of the teeth are denticulate, the den-
ticles are irregular in size. The enameled lingual face
of each tooth display a strong median carina, some
teeth presents also a lateral, slightly bent ridge. In all
the cases, teeth do not show signs of wear.
A dentary fragment (14.4 mm long) was found to-
gether with forelimb remains, vertebrae and ribs, ap-
parently of the same individual early hatchling (figu-
re 3.7). Like in all hadrosaurids, the dentition is orga-
nized into dental batteries; the fragment preserves
seven erupted teeth and the apex of an unerupted
tooth; the teeth are disposed in three rows. The teeth
are leaf-shaped (6.5 mm high and 3.0 mm width),
AMEGHINIANA 47 (1), 2010
covered by a thick layer of enamel and crossed me-
dially by a crest, less prominent or even absent in the
lowermost part of the crown. The two sides around
the medial crest are either flat or banded towards the
crest. Except the lower part of the crown, the teeth
margins are denticulate with 8 to 10 slightly rounded
denticles not supported by marginal ridges, as is the
case in adults.
Conclusions
The taxonomic identification of the hatchlings as-
sociated with megaloolithid eggs within the incuba-
tion horizon from Tustea as belonging to the
hadrosaurid Telmatosaurus transsylvanicus has been
confirmed by D.Weishampel (Weishampel et al.,
1991, 1993), J. Horner and Ph. Currie (personal com-
munication, 1991). All those colleagues who could
not visit the site have questioned the association of
eggs and babies in the same level, suggesting that,
possibly, the egg clutches and baby bones represent
different stratigraphic levels. To answer to this ques-
tion we mention that the “marker bed” we used as a
stratigraphic control in the Tustea outcrop is a pale-
osol level, with a constant position towards the base
of the thick conglomerate bed that dominates the
Tustea geological section (see figure 1.3). This
“marker bed” includes the egg clutches and baby re-
mains and preserves its identity throughout the en-
tire outcrop. Besides this stratigraphic aspect, the
discoveries of the last years have increased the num-
ber of hatchling remains found inside the egg clutch-
es, while teeth and skeletal fragments were also
identified within the egg matrix. All these facts sup-
port the idea that the megaloolithid eggs from
Tustea were laid by the hadrosaurid Telmatosaurus
transsylvanicus. We are not prepared to give a con-
sistent explanation to this convergence in egg mor-
phology and eggshell microstructure between
sauropods and this “most basal hadrosaurid di-
nosaur” (Weishampel et al., 1993) that is
Telmatosaurus transsylvanicus.
Acknowledgements
I thank T. Niculescu for preparing the egg in which embryon-
ic remains, including the tooth were found. I also thank
R.Dimitrescu and M. Dumbrava for the drawings of figs. 1 and 3
(E-G). I am grateful to Z. Csiki for discussions and assistance in or-
ganizing the text and illustrations of the present paper. Finally, I
am indebted to R Coria and L.Chiappe for the opportunity to vis-
it the famous Auca Mahuevo nesting site and for their special hos-
pitality during the 3rd Symposium on Dinosaur Eggs and Babies
from Plaza Huincul in Argentina. I am grateful to L. Salgado for
the Spanish translation of the Abstract (Resumen) and the help in
organizing the final version of the paper.
 Tustea Puzzle: hadrosaur hatchlings along megaloolithid eggs 9
References Grigorescu, D. and Csiki, Z. 2006. Ontogenetic development of
Telmatosaurus transsylvanicus (Ornitischia: Hadrosauria) from
the Maastrichtian of the Hateg Basin, Romania–evidence from
Antonescu, E., Lupu, D. and Lupu, M. 1983. Correlation pali-
the limb bones. Hantkeniana 5: 20-26.
nologique du Cretace terminal du sud-est des Monts
Huene, F., 1932. Die fossile Reptile-Ordnung Saurischia ihre
Metaliferi et des Depressions de Hateg et de Rusca Montana.
Entwicklung und Geschichte. Monographien zur Geologie und
Anuarul Institutului de Geologie si Geofizica al Romaniei 59: 71-77.
Palaeontologie 1, 4, 1–361.
Bojar, A.V., Grigorescu, D., Ottner, F. and Csiki, Z. 2005.
Matheron, M.P., 1869. Notice sur les Reptiles fossils des dépots
Palaeoenvironmental interpretation of dinosaur and mammal
fluvio-lacustres Crétacés du Bassin a lignite de Fuveau.
– bearing continental Maastrichtian deposits, Hateg Basin,
Memoires de l‘Academie Impériale des Sciences, Belle-Lettres et Arts
Romania. Geological Quarterly 49: 205-222.
de Marseille, 39 pp. 4 tab.
Carpenter, K., Hirsch, K.F. and Horner, J.R (ed.). 1994. Dinosaur
Nopcsa, F. 1900. Dinosaurierreste aus Siebenburgen I. Schädel
Eggs and Babies. Cambridge University Press, 372 pp.
von Limnosaurus transsylvanicus nov. gen et nov. spec.
Chiappe, L.M., Coria, R.A., Dingus, L., Jackson, F., Chinsamy, A.
Denkschriften der königlichen Akademie der Wissens-
and Fox, M. 1998. Sauropod dinosaur embryos from the Late
chaften,.Mathematisch-Naturwissenschaftlichen Klasse 68: 555-
Cretaceous of Patagonia. Nature 396: 258-.261.
591.
Codrea, V., Smith, T., Dica, P., Folie, A., Garcia, G., Godefroit, P.
Vianey-Liaud, M., Mallan, P., Buscail, O. and Montgelard, C.
and Van Itterbeeck, J. 2002. Dinosaur egg nests, mammals and
1994. Review of French dinosaur eggshells: morphology,
other vertebrates from a new Maastrichtian site of the Hateg
structure, mineral and organic composition. In K. Carpenter,
Basin (Romania). C. R. Palevol 1: 173-180.
K.F. Hirsch and J.R. Horner (eds.), Dinosaur Eggs and Babies.
Cousin, R. 2002.Organisation des pontes de dinosauriens de la
Cambridge University Press, New York, pp.151-183.
famille des Megaloolithidae Zhao,1979. Bulletin trimestriel de la
Weishampel, D., Grigorescu, D. and Norman, D. 1991. The
Societe geologique de Normandie et des Amis du Museum du Havre.
Dinosaurs of Transylvania. National Geographic Research
Editions du Museum d’Histoire Naturelle du Havre, 177 pp.
and Exploration, Washington DC, 7: 196215.
Garcia, G.and Vianey-Liaud, M. 2001. Nouvelles donnees sur les
Weishampel, D., Norman, D. and Grigorescu, D. 1993.
coquilles d’oeufs de dinosares Megaloolithida du Sud de la
Telmatosaurus transsylvanicus from the Late Cretaceous of
France: systematique et variabilite intraspecifique. C. R.
Romania: the most basal hadrosaurid dinosaur. Paleontology
Academie des Sciences, Paris, Sciences de la Terre et des planetes
36: 361385.
332: 185-191.
Grigorescu, D. 1993. The Latest Cretaceous Dinosaur eggs and
embryos from the Hateg Basin–Romania. Revue de Paleobiologie
7: 9599.
Grigorescu, D., Weishampel, D., Norman, D. and Seclaman, M.
1990. Dinosaur eggs from Romania. Nature 346: 417.
Grigorescu, D., Weishampel, D., Norman, D., Seclaman, M., Rusu,
M., Baltres, A. and Teodorescu, V. 1994. Late Maastrichtian di-
nosaur eggs from the Hateg Basin (Romania). In K. Carpenter,
K.F. Hirsch and J.R. Horner (eds.), Dinosaur Eggs and Babies.
Cambridge University Press, New York, pp. 7587.
Grigorescu, D. and Melinte, M. 2002. Stratigraphy of the Upper
Cretaceous marine sediments from the North-Western Hateg
Basin. Acta Palaeontologica Romaniae 3: 153-160.
Grigorescu, D. and Csiki, Z. (eds.). 2002. Abstracts volume and
Excursions Field Guide, 7th European Workshop on Vertebrate Recibido:
Paleontology, Sibiu, Ars Docendi, Bucuresti, 86 pp. Aceptado:

AMEGHINIANA 47 (1), 2010

View publication stats