Dinosaurs Innovated When it Came to Egg Production

Many museums include the fossilised remains of dinosaur eggs amongst their natural history collections and palaeontologists are aware that reptiles were laying eggs on land long before the dinosaurs evolved, but what we know about the evolution of reptile eggs (amniotic eggs in general), is largely based on inference and conjecture.  The problem is, for the first 100 million years or so of amniote evolution, there is very little fossil data related to reptile reproduction to study.  What we do know, is based on Middle Jurassic to Late Cretaceous fossils.  Although, dinosaur eggs are rare, the examples we do have, such as those associated with Asian oviraptorids demonstrate that dinosaur eggs had thick, hardened shells.  However, a new study suggests that it was not always like this and that the three main Sub-orders of the Dinosauria probably evolved thick, tough eggs independently.

Examples of Eggs from Different Archosaurs (Avian and Non-avian Dinosaurs)

Examples of whole or partial fossilised eggs.

Picture Credit: Royal Society Open Science

Studying Some of the World’s Oldest Dinosaur Eggs Reveals New Information

Writing in the on-line, open access journal “Scientific Reports”, a team of scientists, including Robert Reisz (University of Toronto Mississauga) and Koen Stein (Royal Belgian Institute of Natural Sciences, Brussels), have examined some of the oldest examples of dinosaur eggs known and revealed new information about the evolution of dinosaur reproduction.  The researchers examined the eggs and eggshells of three coeval, but geographically widely distributed Early Jurassic basal Sauropodomorph dinosaurs (Sinemurian faunal stage).  These fossils came from Argentina, China and South Africa and include the eggs of Massospondylus and Lufengosaurus.  Their analysis showed that the basal Sauropodomorph eggs all had the basic structure, they had a thin calcareous layer less than 100 microns thick.  This thin shell layer contrasts strongly with the much thicker calcareous shells associated with Late Jurassic and later dinosaur eggs.

At approximately 195 million years old, they are the earliest known eggs in the fossil record, and they were all laid by similar, herbivorous dinosaurs that ranged in size from four to eight metres in length and were the most common and widely spread dinosaurs of their time.  These types of plant-eating dinosaur were the forerunners of the giant Sauropods of the Jurassic, dinosaurs such as Brontosaurus, Diplodocus and Brachiosaurus.

A Massospondylus Nesting Site (Life Reconstruction)

Massospondylus (basal Sauropodomorpha) nesting site. Massospondylus fossil eggs from South Africa were used in the study.

Picture Credit: Julius Csotonyi

Putting the research into context Professor Reisz explained:

“Reptile and mammal precursors appear as skeletons in the fossil record starting 316 million years ago, yet we know nothing of their eggs and eggshells until 120 million years later.  It’s a great mystery that eggs suddenly show up at this point, but not earlier.”

The researchers concluded that these Early Jurassic eggs represented a step in the evolution of dinosaur reproduction, their shells were paper-thin and brittle, proportionately much thinner than the eggs of extant birds.  However, thicker, tougher eggshells in the Dinosauria were to evolve across all three Sub-orders later.  The much thicker eggshells associated with Sauropods, Ornithischian dinosaurs and the Theropoda must have evolved independently.

Professor Reisz added:

“We know that these early eggs had hard shells because during fossilisation they cracked and broke, but the shell pieces retained their original curvature.”

Other authors of the scientific paper include Edina Prondvai and Jean-Marc Baele.  Shell thickness was analysed along with membrane thickness, mineral content and distribution of pores, looking for clues about why these early eggs might have developed hard shells.  The scientists concluded that hard-shelled eggs evolved early in dinosaur evolution, with thickening of the calcareous layer (greater than 150 microns), occurring independently in several groups, but a few million years later other reptiles also developed hard-shelled eggs.  One possibility is that hard and eventually thicker shells may have evolved to shield dinosaur embryos and other reptiles from predators.

Professor Reisz commented:

“The hard shells would protect the embryos from invertebrates that could burrow into the buried egg nests and destroy them.”

Linked to Increased Oxygen in the Atmosphere

Advanced mineralisation of amniote eggshell including those of dinosaurs (≥150 microns in thickness), in general occurred not earlier than the Middle Jurassic and may correspond with a global trend of an increase in atmospheric oxygen.  If there were higher levels of atmospheric oxygen, then this would facilitate more efficient gaseous exchange through the porous eggshell and across the egg membranes.  More efficient diffusion would permit the evolution of thicker eggshells, which in turn would offer greater resistance to damage and more protection from predators.

A Lufengosaurus Embryo

New research into 195 million-year-old baby dinosaurs and their eggs.

Picture Credit: D. Mazierski

Raising Further Questions About Mesozoic Reproduction Strategies

The study raises some intriguing questions that may well lead to further research projects.  For example, palaeontologists are aware that many types of marine reptile evolved viviparity (live birth), whilst the fossil evidence for the terrestrial Dinosauria seems to indicate that they continued to rely on egg laying.  Why didn’t the highly diverse dinosaurs evolve different reproductive strategies over their 160 million years of existence?

Everything Dinosaur acknowledges the assistance of a press release from the University of Toronto in the compilation of this article.

The scientific paper: “Structure and Evolutionary Implications of the Earliest (Sinemurian, Early Jurassic) Dinosaur Eggs and Eggshells” by Koen Stein, Edina Prondvai, Timothy Huang, Jean-Marc Baele, P. Martin Sander and Robert Reisz published in Scientific Reports.

Galleonosaurus dorisae – A New Aussie Dinosaur

A new type of Australian dinosaur has been described from the fossilised remains of five upper jaw bones (maxillae) found at the Flat Rocks locality in the Wonthaggi Formation in the famous Gippsland Basin of Victoria.  Five small-bodied Ornithopods are now known from the state of Victoria.  The new plant-eating dinosaur has been named Galleonosaurus dorisae.  The jaw bones are of different sizes and this has permitted palaeontologists to plot growth changes in these little dinosaurs as they matured.

Writing in the Journal of Palaeontology, the researchers which include Matthew Herne (University of New England, New South Wales) and Alistair Evans (Monash University, Melbourne), used detailed CT-scans of the fossil material to gain fresh insights into the structure and morphology of the cranial anatomy and dentition of small Australian Ornithopods.  The research leading to the establishment of this new genus has also helped to define more clearly other small Ornithopods known from the Gippsland Basin and the Otway Basin located on the opposite side of Port Phillip Bay.

A Life Reconstruction of the Newly Described Ornithopod Galleonosaurus dorisae

A life reconstruction of the newly described Australian Ornithopod Galleonosaurus dorisae.

Picture Credit: James Kuether

“Galleon Lizard”

When the scientists were examining the maxillae, their shape reminded them of the upturned hull of an old-fashioned sailing ship – a galleon.  It was the morphology of the jaw that inspired the genus name “Galleon Lizard”.  The species or trivial name honours Doris Seegets-Villiers for her geological, palynological, and taphonomic work on the Flat Rocks fossil vertebrate locality.

Jaw Fossils and a Tooth with a CT-scan of the Fossil Material

Fossil jaw bones, a single tooth and a CT-scan image of a jaw bone (Galleonosaurus dorisae).

Picture Credit: Herne et al

Niche Partitioning in Ornithopods

The plethora of vertebrate fossils at the Flat Rocks site, suggests that several more dinosaurs await discovery.  However, for the moment, the researchers are confident that Galleonosaurus shared its habitat with at least one other small, light, fast-running Ornithopod – Qantassaurus intrepidus.  The jaws of Qantassaurus are more robust and more powerful.  The researchers were able to confirm that Q. intrepidus is uniquely characterised by a deep, foreshortened dentary (lower jaw).  This suggests that the robust Q. intrepidus and the more gracile jawed G. dorisae fed on different vegetation, they did not compete directly for food, an example of niche partitioning.

Dr Herne described Galleonosaurus:

“We know it would have been a two-legged, quite agile plant-eating dinosaur.  It seems that Galleonosaurus was no doubt closely related to possibly as many as four or five other species that look a little bit similar and were similar sizes, but we can tell they’re different by the anatomy of the jaws and the teeth.”

A Lush Conifer Dominated Rift Valley with an Immense Volcanic Mountain Chain to the East

Extensive research on the Otway Formation material to the west of Port Phillip Bay in conjunction with research on the geology of the Gippsland Basin have permitted scientists to build up a picture of what life was like in this part of Australia during the Early Cretaceous.  The dinosaurs lived in an extensive rift valley that had formed as Australia began to separate from Antarctica. Conifer forests dominated and at such high latitudes, the lush environment would have been subjected to long periods of extensive daylight in the summer, but conversely the winters would have been cold with little daylight each day.  Although the Earth’s climate was much warmer than today during the Early Cretaceous, it is quite possible that these little dinosaurs would have had to endure winter temperatures close to freezing.

Gondwana in the Early Cretaceous (Barremian Faunal Stage)

Around 125 million years ago, although Gondwana was breaking up, Australia was still linked to Antarctica with a large volcanic mountain range to the east.

Picture Credit: Herne et al

A Skeletal Reconstruction of the Skull of Galleonosaurus and the Anatomical Position of Jaw Material

An illustration of the skull of Galleonosaurus dorisae with fossil elements placed in the correct anatomical position.  The lower jaw shown in the image might pertain to G. dorisae based on a reassessment of other known lower jaw elements associated with Q. intrepidus and Atlascopcosaurus loadsi.

Picture Credit: Herne et al

A Phylogenetic Analysis

The scientists conclude that a highly diverse, small-bodied Ornithopod fauna flourished in the periodically disturbed, high-latitude, riverine floodplain environment of the Australian-Antarctic rift valley during the Early Cretaceous (Barremian to Early Albian faunal stage).  A phylogenetic analysis places Galleonosaurus as the earliest member of the Elasmaria, a clade of Gondwanan Ornithopods distantly related to the Hypsilophodonts.

The Five Victorian Ornithopods – Spanning 12 million years

The Lower Cretaceous rocks either side of Port Phillip Bay were laid down at different times during the Cretaceous.  The Gippsland Basin deposits close to the town of Inverloch, were laid down around 125 million years ago, however, the Otway Basin deposits (Eumeralla Formation), represent younger material laid down in the Early Albian (113 million years ago).

1. Leaellynasaura amicagraphica – named in 1989 (Early Albian faunal stage), from the Eumeralla Formation (Otway Basin).
2. Atlascopcosaurus loadsi – also named in 1989 from the Eumeralla Formation.
3. Diluvicursor pickeringi – named in 2018 (Eumeralla Formation).  To read an article about the discovery of this dinosaur: Fast-running Ornithopod from Victoria.
4. Qantassaurus intrepidus named in 1999 from the Wonthaggi Formation (Gippsland Basin) – older strata associated with the Barremian faunal stage of the Early Cretaceous.
5. The newly described Galleonosaurus dorisae (2019), also from the Wonthaggi Formation.

Dr Herne stated:

“The interesting thing about that whole coast line is it gives us a decent age range over quite a long period.”

A spokesperson from Everything Dinosaur commented:

“It is likely that many more small dinosaurs are going to be named and described in the future.  Fossil finds from Victoria will, most likely, lead to further revisions of Gondwanan Ornithopod taxonomy.”

The Diminutive Tyrannosaur Moros intrepidus

A small, but speedy dinosaur is the newest member of the Superfamily Tyrannosauroidea, a distant relative of the most famous dinosaur of all Tyrannosaurus rex.  T. rex et al might have a reputation for being giant, bone-crunching apex predators, but for much of their evolutionary history, the Tyrannosaurs have been rather over-shadowed by other super-sized dinosaur carnivores.  Indeed, it was only in the last few million years of the Cretaceous that these types of Theropod emerged as the apex predators of northern latitudes.  The new dinosaur, named Moros intrepidus, at approximately 78 kilograms (data range 53 to 85 kilograms), around the same bodyweight as a South American Jaguar (Panthera onca), is about ninety times lighter than its famous top-of-the-food-chain relative.

Ironically, contrary to public opinion, M. intrepidus might just be more typical of the Tyrannosauroidea bauplan than its more famous relatives – Gorgosaurus, Albertosaurus and T. rex.

The Newly Described Moros intrepidus from the Late Cretaceous of Central Utah

Moros intrepidus from the Late Cretaceous of central Utah.

Picture Credit: Jorge Gonzalez

Teeth and a Hind Limb from a New Theropod

The fossilised remains of a partial right leg consisting of a femur, a tibia, metatarsal bones and some toe bones from the fourth toe were discovered in sediments representing the lower Mussentuchit Member of the Cedar Mountain Formation located in Emery County (Utah).  These fossils, in conjunction with isolated teeth from the front portion of the upper jaw (premaxilla) found nearby provide the basis for this new taxon.  The deposits represent a terrestrial environment, a large delta and they date from approximately 96 million years ago (Cenomanian faunal stage of the Late Cretaceous).

Moros intrepidus represents the oldest known Cretaceous-aged tyrannosauroid discovered to date in North America. It extends the definitive fossil record for these types of dinosaurs by around 15 million years.

The Temporal Relationships and Phylogeny of the Tyrannosauroidea

Phylogenetic and temporal relationships between tyrannosauroids and an examination of faunal turnover.  The Allosaurs/Megaraptor apex predator niche was gradually taken over by Tyrannosaurs.

Picture Credit: Nature Communications Biology

In the diagram (above), the section on the left (a), shows the fossil record gap between Late Jurassic tyrannosauroids and much larger Late Cretaceous members of the Tyrannosauridae family such as Lythronax (L. argestes).  Section (b) demonstrates the temporal range of these Theropods and the change in bauplan, whilst (c) demonstrates key evolutionary anatomical changes.  The blue and pink coloured shapes in (d) reflect the transition from Allosaur/Megaraptoran dominated ecosystems to Tyrannosaur dominated palaeoenvironments.

A Changing of the Guard When it Comes to Apex Predators

Palaeontologists know that the Tyrannosaur lineage dates back a long way.  For example, basal tyrannosaurids such as Stokesosaurus (S. clevelandi) are known from Upper Jurassic deposits of Utah.  By the Late Cretaceous (Campanian faunal stage), Tyrannosaurs were large and had become the iconic apex predators beloved by dinosaur fans and film directors.  The fossil record for North American Tyrannosaurs was essentially blank, giving palaeontologists a T. rex skull-sized headache when it came to piecing together how these Theropods changed over time.

The discovery of Moros helps to narrow a 70-million-year-gap in the fossil record of tyrant lizards in North America.

Lead-author of the study, published in “Nature Communications” Lindsay Zanno of the North Carolina Museum of Natural Sciences explained:

“When and how quickly Tyrannosaurs went from wallflower to prom king has been vexing palaeontologists for a long time.  The only way to attack this problem was to get out there and find more data on these rare animals.”

A Silhouette of M. intrepidus Showing the Anatomical Position of the Known Fossil Material

Silhouette of M. intrepidus showing known fossil elements.  Key = (g) femur, (h) tibia, (i) fourth metatarsal, (j) second metatarsal, and (k) pedal phalanges of the fourth digit.   Scale bar (c) 1 m, (g–k) 5 mm.  Note the tooth (views d-f) are not to scale.

Picture Credit: Nature Communications Biology

Living in the Shadow of Siats meekerorum

In 2013, two of the authors of the Moros intrepidus paper, Lindsay Zanno and Peter Makovicky (Field Museum, Chicago), published a study on a large allosauroid from similar-aged sediments.  The dinosaur, named Siats meekerorum is estimated to have measured around 12 metres in length, dwarfing the contemporary Moros, which had a hip height of around 1.2 metres.  The researchers conclude that within a palaeoenvironment dominated by giant, allosauroid Theropods, Tyrannosaurs such as M. intrepidus relied on their speed and small size and would have kept out of the way of the larger predators.

A spokesperson from Everything Dinosaur commented:

“During the Cenomanian, tyrannosauroids like Moros intrepidus were secondary predators within an ecosystem dominated by apex predators from a completely different part of the Theropod family tree.  For the greater part of the Tyrannosaur evolutionary history, these types of dinosaurs were marginal predators, living in the shadow of much bigger carnivorous dinosaurs.”

A Life Reconstruction of Siats meekerorum with two Tyrannosauroids shown in the Foreground

Siats meekerorum has nothing to fear from these two Tyrannosaurs.  Moros intrepidus may have scavenged the kills of larger Theropods but these types of tyrannosauroid were very much the secondary predators.

Picture Credit: Julio Laceardo

To read Everything Dinosaur’s article on the discovery of Siats meekerorum: Unravelling the Apex Predators of the Cretaceous Before Tyrannosaurs

Phylogeny Points at Asian Ancestry

A study of the longer limb bones indicates that the individual was around six to seven years of age when it died.  It was likely to have reached its adult size.  A phylogenetic assessment indicates an affinity with Asian Tyrannosaur taxa, in essence, the ancestors of famous North American dinosaurs such as Gorgosaurus and Tyrannosaurus rex migrated into North America from Asia.

Assistant Research Professor Zanno stated:

“T. rex and its famous contemporaries such as Triceratops may be among our most beloved cultural icons, but we owe their existence to their intrepid ancestors who migrated here from Asia at least 30 million years prior.  Moros signals the establishment of the iconic Late Cretaceous ecosystems of North America.”

Views of the Lower Leg Bones from the Right Leg of M. intrepidus

Right tibia (a–f) and right fourth metatarsal (g–l) of M. intrepidus (NCSM 33392).

Picture Credit: Nature Communications Biology

What’s In a Name?

The etymology of this new tyrannosauroid reflects the later faunal turnover that led to the apex predator roles in North America being dominated by Tyrannosaurs.  The genus name is from the Greek “Moros”, the embodiment of impending doom, for the descendants of this fast-running dinosaur were to evolve into some of the largest and most formidable terrestrial predators known to science.  The species name is from the Latin “intrepidus”, a reference to these intrepid dinosaurs making the migration from Asia into North America and their subsequent dispersal.

Size is Not Everything

Although around ninety times lighter than Tyrannosaurus rex, Lindsay warns against underestimating the predatory abilities of Moros.

She added:

“Moros was lightweight and exceptionally fast.  These adaptations, together with advanced sensory capabilities, are the mark of a formidable predator.  It could easily have run down prey, while avoiding confrontation with the top predators of the day.  Although the earliest Cretaceous Tyrannosaurs were small, their predatory specialisations meant that they were primed to take advantage of new opportunities when warming temperatures, rising sea-level and shrinking ranges restructured ecosystems at the beginning of the Late Cretaceous.  We now know it took them less than 15 million years to rise to power.”

The scientific paper: “Diminutive fleet-footed tyrannosauroid narrows the 70-million-year gap in the North American fossil record” by Lindsay E. Zanno, Ryan T. Tucker, Aurore Canoville, Haviv M. Avrahami, Terry A. Gates and Peter J. Makovicky published in Nature Communications Biology.

Mnyamawamtuka moyowamkia – Heart-shaped Tail Bones Help to Flesh Out Titanosaur Evolution

Scientists writing in the on-line, open access journal PLOS One, have published details of a new species of African Titanosaur.  It took several years to carefully remove the fossil material from a high cliff wall overlooking the Mtuka riverbed in south-western Tanzania, but the fossils, representing a not fully mature dinosaur, are providing palaeontologists with important information about how African ecosystems changed over the course of the Cretaceous.  The new Titanosaur has been named Mnyamawamtuka moyowamkia (pronounced Mm-nya-ma-wah-mm-too-ka mm-oh-yo-wa-mm-key-ah).  The name is derived from Kiswahili for “animal of the Mtuka river with a heart-shaped tail”.

The heart-shaped tail element of the name refers to the strange shape of the most intact middle caudal vertebra described in the paper.  It bulges out at the sides (dorsolateral expansion of the posterior articular surface of the centrum), a unique tail bone morphology that resembles the shape of a romantic, love heart.

A Life Reconstruction of a Pair of  Mnyamawamtuka moyowamkia Titanosaurs

Mnyamawamtuka moyowamkia life reconstruction.

Picture Credit: Mark Witton

The first evidence of the dinosaur fossils was noted in 2004 and some fossils were excavated from the cliff face, in what were quite hazardous conditions, with field team members having to be lowered over the cliff on numerous occasions to work on the exposed bones.  Annual excavations took place until 2008, it was important to keep returning to the site as the fossils were in danger of being lost to the river in seasonal floods.

A Line Drawing of the Quarry Site Showing the Extent of the Annual Excavations

Quarry map showing the layout and excavation timeline of the M. moyowamkia fossil material.

Picture Credit: PLOS One

Around 110-100 Million Years Old

The specimen was excavated from the Mtuka Member of the Cretaceous Galula Formation, which was deposited around 110 to 100 million years ago (Aptian to Cenomanian faunal stage of the Cretaceous).  A substantial portion of the postcranial skeleton has been recovered.

Dangerous Work!  The Excavation Site in 2007

The quarry dig site above the Mtuka riverbed in south-western Tanzania.

Picture Credit: Ohio University

Commenting on the importance of this discovery, in relation to the evolution of African Titanosaurs, lead author of the paper, Dr Eric Gorscak, a recent PhD graduate of Ohio University and now an assistant professor at the Midwestern University (Illinois), stated:

“Although Titanosaurs became one of the most successful dinosaur groups before the infamous mass extinction capping the Age of Dinosaurs, their early evolutionary history remains obscure, and Mnyamawamtuka helps tell those beginnings, especially for their African-side of the story.  The wealth of information from the skeleton indicates it was distantly related to other known African Titanosaurs, except for some interesting similarities with another dinosaur, Malawisaurus, from just across the Tanzania–Malawi border.”

Adding to the Diversity of Titanosaurian Sauropods from Africa

The field team responsible for this discovery have also found the fossilised remains of two other Titanosaurs in this part of Africa.  In 2017, Everything Dinosaur reported upon the discovery of Shingopana songwensis.

To read about S. songwensis: A New Species of African Titanosaur is Named

In addition, Rukwatitan bisepultus another Titanosaurian Sauropod dinosaur, was named and described in 2014: A New Species of Titanosaurian Sauropod Rukwatitan bisepultus

The researchers conclude that Mnyamawamtuka moyowamkia was distantly related to both Shingopana songwensis and Rukwatitan bisepultus, fossils of which come from younger Cretaceous sediments, although it did share some anatomical characteristics with Malawisaurus dixeyi from Malawi, that might have been contemporaneous.  This new fossil discovery is helping palaeontologists to better understand the distribution of Titanosaurs between Africa and South America and their evolutionary relationships.

Heart-shaped Tail Bones

One of the middle caudal centra (tail bone from the middle portion of the tail), exhibits a unique dorsolateral expansion of the posterior articular surface of the centrum.  This unique characteristic was found to be present in the most intact middle caudal vertebra described.  This unique shape inspired the dinosaur’s name.

Heart-shaped Tail Bone Centrum (Rear View)

A posterior view of a middle caudal vertebra showing the characteristic heart shape.  The term dle = dorsolateral expansion.

Picture Credit: PLOS One

Tail Bones of Different Titanosaurs Compared

A comparison of caudal vertebrae between three Titanosaurs. Mnyamawamtuka moyowamkia (A), compared with Malawisaurus dixeyi (B) and Lohuecotitan pandafilandi of the Late Cretaceous of Spain (C). Posterior views and lateral views, scale bar = 10 cm.  The term dle = dorsolateral expansion.

Picture Credit: PLOS One

The scientific paper: “A New African Titanosaurian Sauropod Dinosaur from the Middle Cretaceous Galula Formation (Mtuka Member), Rukwa Rift Basin, Southwestern Tanzania” by Eric Gorscak and Patrick M. O’Connor published in PLOS One.