Dinosaurs from the North Pole: Live at Prince Creek

When we think about dinosaurs, we probably imagine them walking through a dense, tropical jungle or wandering in a warm, foggy swamp. But as a matter of fact, some dinosaur species lived in very high latitudes, as the ones found in the Prince Creek formation. This Alaskan geologic formation is one of the most important sources of arctic dinosaurs, as many fossils have been found in it. In this entry, we’ll describe some of these dinosaurs from the North Pole, and we’ll explain some of the difficulties they had to endure in order to survive in the northernmost point of the planet.

ALASKA 75 MILLION YEARS AGO

The Prince Creek formation is situated in the north of Alaska and dates from around 80-60 million years ago, at the end of the Cretaceous, the last period of the Mesozoic. At that time, North America was divided by the Western Interior Seaway; the eastern continent or Appalachia, and the western continent or Laramidia, north of which the Prince Creek formation was deposited.

Map of North America at the end of the Cretaceous period, with the Prince Creek formation marked in red, from the article New Horned Dinosaurs from Utah Provide Evidence for Intracontinental Dinosaur Endemism.

At the end of the Cretaceous period, the Prince Creek formation was further north than it is today. Yet, at that time the Earth was going through a greenhouse effect phase, so the climate was a little warmer than it is today. It is thought that the mean annual temperature at Prince Creek was about 5°C, with summer maximums at about 18-20°C. Still, the difference in temperature between summer and winter would have been quite remarkable (currently, at the same latitude, it’s about 56°C).

Even if temperatures were not as low as the ones of present-day Alaska, the dinosaurs of Prince Creek had to endure long, dark winter months. Yet, the slightly higher temperatures and the proximity to the sea, produced a higher diversity of plant species. Observing the fossilized flora, we know that the landscape was that of a polar woodland, with angiosperm-dominated forests and a large number of fern, moss and fungus species, with some areas of seasonally-flooded grasslands.

Drawing by Julio Lacerda of Prince Creek’s landscape and wildlife.

As for the fauna, palaeontologists were surprised at the great number of big animals found. The fact that dinosaurs were found in such high latitudes is what makes us think that these were endotherm animals that generated their own body heat. Also in Prince Creek, there aren’t any fossils of other ectotherm reptiles like turtles, crocodiles or snakes, which are usually found in other United States deposits of the same period. Currently, dinosaurs are thought to be neither endotherm nor ectotherm, but mesotherm animals, which generated body heat metabolically, but were unable to control its temperature or keep it stable.

TOUGH HERBIVORES

The relatively abundant vegetation, allowed the presence of a great diversity of plant-eating dinosaurs in such high latitudes. While the smaller herbivores had little trouble because of their low energetic requirements, the larger herbivores probably had more difficulties in order to find enough food, especially during the harsh winter months. The dinosaur fossil found at the highest latitude is Ugrunaaluk (literally “ancient grazer” in Inupiaq language from northern Alaska) a hadrosaurid or “duck-billed dinosaur”. This ornithopod measured up to 10 metres long and weighed around 3 tonnes, making it one of the largest animals in Prince Creek.

Reconstruction by James Havens of a herd of Ugrunaaluk kuukpikensis, moving under the polar lights.

Ugrunaaluk were herbivorous animals that lived in groups. Even if many author think that these animals performed long migrations like today’s birds and mammals in order to avoid the lack of food during the winter, some others argue that young Ugrunaaluk (which had a less active metabolism than current endotherms) would had been unable to endure such long journeys. Ugrunaaluk probably moved to areas were the vegetation better tolerated the severity of winter, even if it’s thought that these great herbivores survived during the dark winters feeding on bark, ferns and probably aquatic vegetation during the coldest months.

The other great Prince Creek plant-eater was Pachyrhinosaurus (literally “thick-nosed lizard”) a ceratopsid widely-distributed through the United States, with a large protuberance on its nose which may have been used as a weapon during intraspecific combats, and a pair of laterally-facing horns on the top of its frill. Pachyrhinosaurus was the largest animal of Prince Creek, measuring up to 8 metres long and weighing up to 4 tonnes. It is possible that it used its nasal protuberance to shovel through the snow to reach the plants buried under it, similar to today’s bisons.

Reconstruction of a pair of Pachyrhinosaurus perotorum by James Havens.

All the animals of Prince Creek had arduous lives. Almost all the fossils of both Ugrunaaluk and Pachyrhinosaurus, indicate that these species matured quickly and died young. Observing the growth of the different bones that have been found, it is thought that these dinosaurs rarely lived for over 20 years of age, probably due to the harsh conditions of their habitat but also to the presence of predators.

PREDATORS LARGE AND SMALL

The largest predator of the region was Nanuqsaurus (“polar bear lizard”, from Inupiaq language), a tyrannosaurid. This animal had a highly developed sense of smell which allowed it to detect their prey or animal carcasses in low light during the polar winter. Also, although there is no evidence, it was probably covered in feathers which would have protected it from the cold, as many closely-related theropods presented feathers to some extent.

Reconstruction of Nanuqsaurus hoglundi by Tom Parker.

What’s more surprising about Nanuqsaurus is its size, much smaller than that of its relatives. While other tyrannosaurids from the same time measured between 10 or 12 metres long and weighed up to 9 tonnes, Nanuqsaurus was more of a pygmy tyrannosaur, with an estimated length of 6 metres and a weight of 800 kg. This diminutive size was probably caused by the fact that it lived in an environment where food availability varied through the seasons. Apart from the fact that their prey’s population densities probably weren’t very high, during winter months many herbivores would migrate to other areas.

By contrast, there was another theropod that presented the opposite adaptation. Troodon (“wounding tooth”) was a relatively small dinosaur, about 2.9 metres long and 50 kg of weight. This is a pretty abundant dinosaur in many North American deposits. Troodon was a highly active carnivorous animal, with a good binocular vision and it’s also believed to be one of the most intelligent Mesozoic dinosaurs.

Reconstruction of two Troodon inequalis playing in the snow by Midiaou.

While Nanuqsaurus was smaller by the lack of abundant prey, Troodon specimens found at Prince Creek were characterized by their bigger size, compared with the ones from other deposits. This is what is called the Bergmann’s Rule, according to which the populations of a species that live in colder climates tend to be larger than the populations living in warmer climates, as this way they lose less body heat. Also, the larger eyes of Prince Creek’s Troodon, would give them advantage hunting during the long winter nights.

Image from the article A Diminutive New Tyrannosaur from the Top of the World, in which we can see the size of Nanuqsaurus (A) compared with some other tyrannosaurids (B, C, D and E) and two Troodon specimens (F and G) from different latitudes.

As you can see, dinosaurs not only thrived in warm and tropical environments. Even if their populations weren’t very large and their living conditions were harsher, these dinosaurs were able to adapt and survive in the polar forests of Prince Creek, and many of them surely gazed at the spectacular northern lights of 75 million years ago.

Assembly of the different dinosaur species from the Prince Creek formation by James Kuether.

REFERENCES

The following sources have been consulted during the elaboration of this entry:

• Susana Salazar Jaramillo (2014). Paleoclimate and paleoenvironment of the Prince Creek and Cantwell formations, Alaska: terrestrial evidence of middle Maastrichtian greenhouse event. Thesis.
• Live Science. Polar Dinosaurs Endured Cold Dark Winters.
• The Strange Lives of Polar Dinosaurs.
• North Pole Dinosaurs Lived Short, Hard Lives: Discovery News.
• Gangloff, Fiorillo & Norton (2005). The first Pachycephalosaurine (Dinosauria) from the Paleo-Arctic of Alaska and its Paleogeographic implications. Journal of Paleontology. Vol. 79. Pp: 997-1001.
• Mori, Druckenmiller & Erickson (2016). A New Hadrosaurid from the Prince Creek Formation (Lower Maastrichtian) of Northern Alaska. Acta Palaeontologica Polonica. Vol. 61. Pp: 15-32.
• Fiorillo & Tykoski (2014). A Diminutive New Tyrannosaur from the Top of the World. PLoS ONE 9(3).
• Cover Image by Julio Lacerda.

Synapsids: Before dinosaurs ruled the Earth

Before dinosaurs ruled the Earth, at the end of the Palaeozoic Era, the land was dominated by the synapsids. The synapsids (the amniote line that includes mammals) were a highly successful group which occupied most niches during the late Carboniferous and the Permian periods, but at the end of the Palaeozoic Era most families were extinguished by the Permian-Triassic mass extinction (around 252 million years ago) with only the mammalian line surviving to the present day. In this entry we’ll look at some of the more peculiar synapsid groups, which have led to the evolution of mammals like us.

CHARACTERISTICS AND EVOLUTIONARY TRENDS

The clade Synapsida includes mammals and all other amniotes more closely related to them than to reptiles. The synapsids were the first amniotes to diversify and appeared about 320 million years ago, at the middle of the Carboniferous period. These first synapsids were characterized by the presence of only one temporal fenestra behind each orbit through which the jaw muscles pass. Synapsida literally means “fused arches” referencing to the zygomatic arches (because in the past scientist believed that synapsids had evolved from diapsid reptiles and so their arches were thought to be “fused”).

Drawing of Archaeothyris’s skull, in which we can see some of the characteristics of the synapsids, like the temporal fenestrae and caniniform teeth. Drawing by Gretarsson.

Other characteristics that appeared through their evolution were:

• Differentiation of differently-shaped (heterodont) teeth.
• Lower jaw formed by fewer bones.
• Acquisition of a more erect posture and an endothermic metabolism.

The first groups of more primitive or “reptile-like” synapsids are informally called pelycosaurs, while the latter more advanced forms are called therapsids (clade Therapsida, which in fact derived from pelycosaurs). As we will see, the evolution of synapsids is of the kind of “one group, which includes the next group, which includes the next group”.

Modified evolutionary tree of the amniotes by Kenneth D. Angielczyk (2009).

THE ORIGIN OF SYNAPSIDS, THE PELYCOSAURS

Reconstruction of Cotylorhynchus, a caseasaurian that grew up to 3 metres long. Drawing by Dmitry Bogdanov.

The first synapsids had a sprawling limb posture, low slung bodies and were probably ectothermic. If we look at the skull morphology, the earliest groups of synapsids were the caseasaurians (clade Caseasauria) characterized by their small heads, an overdeveloped snout and their huge bodies (they were probably ectothermic and slow-moving creatures). Yet, if we look at the postcranial skeleton, the earliest synapsids were two groups called the varanopids and the ophiacodontids (Varanopidae and Ophiacodontidae families) which were similar to varanids (through convergent evolution), and while the former were quite small and agile creatures, the latter developed bigger forms with huge heads.

Drawings of the varanopid Varanodon (top) and the ophiacodontid Ophiacodon (bottom). Both drawings by Dmitry Bogdanov.

Just before the appearance of the more advanced therapsids, the last two groups of “pelycosaurs” evolved and occupied most land ecosystems. Both groups shared a tall sail along their backs (similarly to Spinosaurus) formed by tall neural spines. In life, this sail probably was covered in skin and had plenty of blood vessels. Although it’s believed that these two groups were still ectothermic, this sail was probably used to gain or lose heat more easily.

Reconstruction of different species of edaphosaurids of the genus Ianthasaurus, showing their characteristic sail. Drawing by Dmitry Bogdanov.

The first of these groups is the Edaphosauridae family. Unlike most basal synapsids, the edaphosaurids were herbivorous and, along with the caseasaurians, they were among the first large amniotes to adopt a vegetarian lifestyle. The sails of edaphosaurids were covered with spiny tubercles, of which their function is still debated.

Skeleton of Edaphosaurus from the Field Museum of Chicago, where the tubercles on its spines are shown. Image by Andrew Y. Huang (2011).

The other group, the Sphenacodontidae family, were the sister group of the therapsids, inside the clade Sphenacodontia. While all other pelycosaur clades had their teeth loosely set in the jaw, the sphenacodontians had their teeth set in deep sockets. Most sphenacodontids were carnivorous, with strong jaws and well-developed caniniform teeth. Some species became the top predators on land before the apparition of the therapsids.

Reconstruction of the sphenacodontid Dimetrodon, by Dmitry Bogdanov.

THE FIRST THERAPSIDS

Skeleton of Biarmosuchus, a basal therapsid in which we can see its more erect posture. Image by Ghedoghedo.

The therapsids (clade Therapsida, “beast arches”) appeared around 275 million years ago and replaced the pelycosaurs as the dominant land animals in the middle Permian. Early therapsids already had a more erect posture, unlike the sprawling limbs of the pelycosaurs. Also, their temporal fenestrae were larger, which made their jaws more powerful.

Reconstruction of Estemmenosuchus, a dinocephalian from which fossil skin imprints have been found and so it’s known that it was covered in smooth, glandular skin without scales. Drawing by Mojcaj.

The therapsids diversified greatly and developed some extraordinary adaptations. The dinocephalians (clade Dinocephalia, “terrible head”) developed bony head knobs which are believed to be involved in some kind of head-butting behaviour. Another group, the anomodonts (clade Anomodontia, “abnormal teeth”) were characterized by having no teeth except for a pair of upper canines (which were probably covered by a beak). The anomodonts were the sister group of theriodontians.

Reconstruction of Placerias, an anomodont which could weigh up to one tonne. Drawing by Dmitry Bogdanov.

THERIODONTIA AND THE FIRST SABER-TEETH

Theriodontians (clade Theriodontia “beast teeth”) became the most successful group of synapsids. The three main groups probably looked pretty mammal-like, with fully-erect posture, a secondary bony palate which allowed them to breathe while swallowing or holding a prey and heterodont teeth (incisiviform, caniniform and molariform teeth). The most primitive theriodontian group were the gorgonopsians (clade Gorgonopsia, Gorgonopsidae family). All members of this group were carnivorous and active predators, as revealed by their sabre-toothed teeth. Although most of them were of a modest size, the larger ones reached up to 3 metres long and had canines of up to 15 cm long.

Reconstruction of Inostrancevia, the largest gorgonopsid genus, preying upon Scutosaurus, a parareptilian. Drawing by Dmitry Bogdanov.

A second group, the therocephalians (clade Therocephalia, “beast head”), were pretty more advanced than the gorgonopsians, although they didn’t reach their cousins’ size. Their feet resembled those of early mammals, they presented small pits on their bones which probably supported whiskers on fleshy lips, and most evidence suggests that they were already endotherms.

Reconstruction of a pair of Pristerognathus, a therocephalian genus in which we can see some more mammalian characteristics. Drawing by Dmitry Bogdanov.

Both gorgonopsians and therocephalians disappeared at the end of the Permian. The only therapsid group that survived through the Mesozoic period and that coexisted with the dinosaurs were the cynodonts

SMALL CYNODONTS

The cynodonts (clade Cynodontia “dog teeth”) appeared at the late Permian and diversified greatly along with the archosaurs. Although it is not really proven, most paleoartists represent cynodonts covered in fur, as evidence suggests an endothermic metabolism. Some characteristics of the cynodonts were:

• Lower jaw formed only by the dentary bone, while the other jaw bones became the ossicles of the middle ear (the articular, the quadrate and the angular bones evolved into the malleus, the incus and the stapes).
• Complex teeth: incisors to hold, canines to pierce, and premolars and molars to chew.
• Only two sets of teeth (diphyodonts), instead of constantly-renewing teeth (polyphyodonts like most reptiles).
• Large brain cavities. Some fossil burrows of different cynodonts have been found, revealing complex social behaviours.

Reconstruction of Thrinaxodon, a burrowing cynodont with whiskers and hair. Image by Nobu Tamura.

Even if they competed with archosaurs, some early forms became quite large. For example, some carnivores, like Cynognathus, had a large head and measured 1 metre long, while Trucidocynodon was about the size of a leopard. Yet, the evolutionary trend would make the cynodonts smaller, like the Brasiliodontidae family which, like most cynodonts, lived in the shadow of dinosaurs and other bigger reptiles. Brasiliodontids are thought to be the sister group of the Mammaliaformes (mammals and their most recent relatives).

Reconstruction of Brasilitherium, one of the most advanced non-mammalian cynodonts, which was only 12 cm in length. Drawing by Smokeybjb.

Finally, mammals appeared at the end of the Triassic period around 225 million years ago. The first mammaliaforms were probably, insectivorous, nocturnal shrew-like animals. It is thought that this nocturnal lifestyle is what actually propelled the development of fur coats, because in therapsids endothermy appeared before fur did. These mammaliaforms probably had mammary glands to feed their young when they had no teeth, but they probably had no nipples like current monotremes.

Live reconstruction of Megazostrodon, a small mammaliaform which represents very well the transition from cynodonts to modern mammals. Image by Udo Schröter.

After the extinction of most archosaurs at the end of the Cretaceous period, the surviving synapsids took over the empty ecological niches. Mammals have ruled the world since then, conquering the land, the sea and even the air, but it wouldn’t have been possible without all the different adaptations acquired by early synapsids throughout their evolution. Thanks to them, humans and all other mammals are currently the dominant animals on the planet.

REFERENCES

The following sources have been used during the elaboration of this entry:

• http://tolweb.org/accessory/Synapsid_Classification_&_Apomorphies?acc_id=466
• http://www.tandfonline.com/doi/full/10.1080/14772019.2011.631042
• http://journals.plos.org/plosone/article?id=10.1371/journal.pone.0094518
• http://www.eurekalert.org/pub_releases/2015-10/sovp-ans102915.php
• http://www.tandfonline.com/doi/abs/10.1080/02724634.2011.627074
• http://dibgd.deviantart.com/gallery/
• Cover image by Mauricio Antón.

The evolution of amphibians: the conquest of the land

Amphibians were the first group of vertebrates to develop limbs and to be able to leave the water to conquer the land. Even if they are seen as simple and primitive animals by most people, amphibians show a wide diversity of survival strategies which have allowed them to occupy most terrestrial and fresh-water habitats. On this entry we’ll explain some of the aspects related to their evolution, explaining how our ancestors managed to get out of the water.

ORIGIN OF THE AMPHIBIANS

Current amphibians, together with reptiles, birds and mammals are found within the superclass Tetrapoda (“four limbs”), the vertebrate group that abandoned the sea to conquer the land. These first tetrapods were amphibians and they evolved around 395 million years ago during the Devonian period from lobe-finned fish named sarcopterygians (class Sarcopterygii, “flesh fins”) within which we find the coelacanth and the current lungfish.

Specimen of coelacanth (Latimeria chalumnae) a sarcopterygian fish, photo by smerikal.

This group of fish is characterized by its fins which, instead of being formed by rays like in most bony fish, they have a bony base that allowed the subsequent evolution of the limbs of the first amphibians. Within the sarcopterygians, the nearest relatives of the tetrapods are the osteolepiformes (order Osteolepiformes) a group of tetrapodomorph fish that got extinct about 299 million years ago.

Restoration of Eusthenopteron, an extinct osteolepiform, by Nobu Tamura.

ADAPTATIONS TO LIVE ON LAND

The conquest of land was not done from one day to the other; it was possible with the combination of multiple adaptations. Some of the most important characteristics that allowed the first amphibians to leave the water were:

• Evolution of lungs, which are homologous to the gas bladder that allows fish to control its buoyancy. Lungs appeared as an additional way to get oxygen from the air. In fact, there is actually a sarcopterygian family the members of which have lungs to get oxygen from the air, for they live in waters poor on oxygen.

• Dissection of Protopterus dolloi a sarcopteryigian fish with lungs.

• Development of the choanaes, or internal nostrils. While fish present a pair of external nostrils at each side of its snout through which water passes on while swimming, the ancestors of the tetrapods only had one external nostril at each side connected to the internal nostrils, the choanae, which communicated with the mouth. This allowed them to get air through their noses using lung ventilation and this way to smell outside of water.
• Apparition of the quiridium-like limb. The quiridium is the tetrapod’s most basic characteristic. This limb is known for having the differentiated parts: the stylopodium (one bone, the humerus or the femur), the zeugopodium (two bones, the radius or tibia and ulna or fibula) and the autopodium (fingers, hands, toes and feet). While the stylopodium and zeugopodium derived from the sarcopterygian’s fins, the autopodium is a newly-evolved structure exclusive from tetrapods.

Simplified drawing of the structure of the quiridium, by Francisco Collantes.

In short, the relatives of the osteolepiformes developed the tetrapod’s typical characteristics before ever leaving water, because they probably lived in brackish, shallow waters, poor in oxygen and that dried out quickly and often.

THE FIRST AMPHIBIANS

Probably the creature known as Tiktaalik is the closest animal to the mid-point between the osteolepiformes and the amphibians. The first recorded amphibians were labyrinthodonts meaning that their teeth had layers of dentin and enamel forming a structure similar to a maze.

Cross-section of a labyrinthodont tooth, form "On the Genesis of Species", by St. George Mivart.

There were four main groups of primitive amphibians, each characterized by: a group that includes the first animals that were able to get out of water, a second group which contains the ancestors of the amniotes (reptiles, birds and mammals) and two more groups, both candidates to be the ancestors of modern amphibians.

Order Ichthyostegalia

Ichthyostegalians were the first tetrapods to be able to leave the water. They appeared at the late Devonian period and they were big animals with large wide heads, short legs and an aquatic or semi aquatic lifestyle (they probably were pretty clumsy on land). They moved around using mainly their muscular tail with rays similar to that of fish.

Fossil and restoration of Tiktaalik. Photo by Linden Tea.

Similarly to current amphibians, they presented a lateral line (sensory organ that allows fish to detect vibrations and movement underwater) and were able to breathe through their skin (they lost the cosmoid scales of their ancestors). Also, the eggs were laid in the water, from which the tadpoles emerged and later on, they suffered a metamorphosis process to become adults just like current amphibians. Subsequently ichthyostegalians gave rise to the rest of amphibian groups.

Skeletons of Ichthyostega and Acanthostega, two typical ichthyostegalians.

Clade Reptiliomorpha

Reptiliomorphs were the ancestors of amniotes and appeared about 340 million years ago. Most of them were usually large and heavy animals, which presented more advanced adaptations to live on land (laterally-placed eyes instead of dorsally-placed ones and a knobby more impervious skin). Even though, reptiliomorphs still laid their eggs in the water and had larval-stages with gills. It wouldn’t be until the late Carboniferous period when the first amniotes (animals that could lay their eggs on dry land) would emancipate completely from water.

Mounted skeleton of Diadectes a large herbivorous reptiliomorph from the American Museum of Natural History, photo by Ghedoghedo.

Order Temnospondyli

This group is one of the possible candidates to being the ancestors of modern amphibians. This is the most diverse group of primitive amphibians and it survived until the early Cretaceous period, about 120 million years ago. The temnospondyls varied greatly in shape, size and lifestyle.

Restoration of Eryops megacephalus a large temnospondylian predator, by Dmitry Bogdanov.

Most of them were meat-eaters, but some were terrestrial predators, some were semi aquatic and some had returned completely to water. Even though, all species had to return to water to breed for the fertilization was external; while the female was laying clutches of eggs in the water, the male released the sperm over them.

Mounted skeleton of Koskinonodon a 3 metres long temnospondyl, from the American Museum of Natural History, photo by Lawrence.

Within the temnospondyls we can find some of the biggest amphibians that ever lived, such as Prionosuchus, with an estimated length of 4,5 meters and about 300 kilograms of weight. Also, even though their skin was not covered with scales, it wasn’t completely smooth like in modern amphibians.

Restoration of Prionosuchus by Dmitry Bogdanov.

It is believed that this group could be the sister-taxon of modern amphibians, even though there’s one last group which could be a candidate to that post.

Order Lepospondyli

Lepospondyls were a small group of primitive animals which appeared at the early Carboniferous and disappeared at the late Permian period. Even though lepospondyls were not as numerous and smaller than the temnospondyls, they presented a wide range of body shapes and adaptations.

Restoration of Diplocaulus magnicornis, of about 1 metre long was the biggest of all lepospondyls, by Nobu Tamura.

The first lepospondyls looked superficially like small lizards, but subsequently lots of groups suffered processes of limb reduction or loss.

Restoration of Pelodosotis, an advanced lepospondyl, by Dmitry Bogdanov.

The relationship of the lepospondyls with the rest of tetrapods isn’t very clear. Different hypothesis go from some authors arguing that they are a group separated from the labyrinthodonts, some thinking that they are the ancestor of current amphibians and reptiles, and some even saying that they are the ancestors of only a portion of modern amphibians.

Restoration of Lysorophus, a Permian lepospondyl, by Smokeybjb.

As we can see, the classification of primitive amphibians can be an extremely complex thing. On this entry I tried to make a summary of the most important groups of ancient amphibians and, on the next one, we’ll center on the evolution of modern amphibians, the so-called “lissamphibians”, and we’ll look in more detail all the controversies surrounding these curious animals.

REFERENCES

The following sources have been consulted in the elaboration of this entry:

• Cover photo: Tyler Keillor and Beth Rooney.
• http://www.ncbi.nlm.nih.gov/pmc/articles/PMC1691537/pdf/14667343.pdf
• http://www.biology.ualberta.ca/courses.hp/biol606/papers/Ahlberg+1998.pdf
• http://usf.usfca.edu/fac_staff/dever/tetrapod_review.pdf
• http://icb.oxfordjournals.org/content/early/2007/08/13/icb.icm055.full.pdf+html
• http://www.usfca.edu/fac_staff/dever/tetropodevol.pdf
• http://icb.oxfordjournals.org.sci-hub.org/content/5/2/287

Spinosaurus: the first aquatic dinosaur?

Recently, the BBC documentary series “Planet Dinosaur” has premiered on TVE2. In this series the latest paleontological discoveries concerning the biology of dinosaurs are explained. On my last entry we talked about the theropod dinosaurs, one of which is the spinosaur, one of the largest predators that have ever existed. On this entry I’m going to explain some of the facts that paleontology has revealed about the lifestyle of this creature.

TAXONOMY

The spinosaur (scientific name Spinosaurus aegyptiacus) belonged to the Spinosauridae family, a group of specialized theropods which appeared during the late Jurassic and became extinct about 93 million years ago during the late Cretaceous. This group was characterized by being relatively large theropods, with conic teeth and long snouts similar to crocodiles, and elongated neural spines through its back forming a sail-like structure (that’s where the name “Spinosauridae“ comes from, meaning spine reptiles).

Comparition of the different sizes of various spinosaurids by Scott Hartman. From right to left: Irritator challengeri, Baryonyx walkeri, Suchomimus tenerensis and Spinosaurus aegyptiacus.

Some of the more famous members on this family are, the Baryonyx from Europe, which had long curved claws on its hands to capture the fish it fed on, similar to its close relative the Suchomimus from northern Africa. Furthermore, there was the smaller Irritator of about 3 metres tall found in Brasil and finally, the Spinosaurus from northern Africa, which measuring between 12 and 18 metres long and wheighing between 7 and 20 tons, was one of the biggest predators to ever walk on land.

HABITAT AND DISTRIBUTION

The genus Spinosaurus was distributed in the zone of what is now the north of Africa. This genus lived during the Cretaceous, appearing about 112 million years ago and disappearing about 97 million years ago.

Map of the World 94 million years ago by Joshua Doubek, during the middle Cretaceous period.

During that period, the northern part of Africa was a very humid zone with high temperatures and lots of wetlands. Spinosaurs probably lived in areas with large rivers and mangrove forests next to the sea, where tidal movements flooded its habitat during certain seasons of the year. This is in accordance with the vision that spinosaurids preferred wet semiaquatic habitats with plenty of great fish to prey upon.

Reconstruction from 2010 of Spinosaurus aegyptiacus by Dmitry Bogdanov.

Currently there are two possible spinosaur species. The most famous is Spinosaurus aegyptiacus from Egypt, the species of which we have more information. A possible second species is Spinosaurus maroccanus from Morocco, which some authors consider simply as a subpopulation of Spinosaurus aegyptiacus.

FUNCTION OF NEURAL SPINES

Spinosaurs were discovered in 1912 from a fossil which included its characteristic dorsal spines. These spines grow up to a length ten times that of the vertebra from which they emerged.

The scarcity of spinosaur fossils means that the function of the spines is still a mystery for science, although there are some hypothesis. One of these is that the spines formed a “sail” along the back of the animal which was highly irrigated and helped the animal’s thermoregulation, as such a big animal probably would have had problems losing heat. Therefore its sail would have helped the spinosaur to evade overheating, orienting it towards fresh winds to cool down.

Reconstruction of the skeleton of a subadult spinosaur (Japan Museum, photo by Kabacchi).

Another hypothesis tells us that the spines held a hump-like structure similar to that of camels, which the animal would have used as a fat reserve system to store fat to withstand periods with little available feeding resources.

Lots of paleontologists think that both hypothesis could be correct and that the spinosaur used the sail both to regulate its body temperature and also to store fat to resist periods of low prey abundance. It is also possible that the sail made the spinosaur appear bigger than it actually was and that they used it during mating rituals similar to those of the modern peacock.

FEEDING STRATEGIES

The Spinosaurus‘s skull shows adaptions to a piscivorous diet. The snout is longer and slender than on other theropods. Aside from this, observing the snout of Spinosaurus it has been seen that it presents a series of little holes similar to those found on crocodiles. It is thought that these structures indicate the presence of pressure receptors which helped them detect the movement of their preys underwater.

Upper jaw of Spinosaurus from the Museo di Storio Naturale di Milano, where the holes which possibly contained the pressure receptors can be seen.

While the teeth of most carnivorous theropods where curved and serrated on their posterior part to tear flesh, spinosaur teeth were conic in shape and had no serration, more similar to those of crocodiles. These teeth were more useful for catching and holding fast and slippery prey and to prevent them from escaping (for example, a fish). Also, various Spinosaurus fossils have been found to have between their theeth scales and bones of large prehistoric fish which probably populated many rivers during the Cretaceous period.

Reconstruction by Joschua Knüppe of two Mawsonia species, the rests of which have been found between the teeth of Spinosaurus.

Nevertheless, it is generally believed that the spinosaur was probably an opportunistic predator, feeding mainly on fish, also hunting small dinosaurs when it had the opportunity and stealing prey from smaller predators using its great size to intimidate them.

POSTURE AND LIFESTYLE

Spinosaurs have traditionally been represented as bipedal animals, as most similarly-sized theropods have. Eventhough most fossils are actually pretty incomplete, it is known that its forelegs were more developed than in most theropods, having long curved claws.

Traditionally it was thought that Spinosaurus hunted in a manner similar to a grey heron, roaming through zones of shallow water, sinking its long snout underwater to detect prey using the pressure receptors, and catching fish with its jaws. It then, probably used its front legs as hands to tear its prey to small pieces easy to swallow.

Reconstruction by Joschua Knüppe of Spinosaurus aegyptiacus in hunting posture.

At the end of 2014 a new Spinosaurus fossil was discovered which has changed the view we had on this animal. For the first time, scientists found a fossil which shows the structure of the hind legs of this dinosaur and they have observed a number of characteristics not found in any other theropod not even in other spinosaurids. This fossil shows that the hind legs of Spinosaurus were much more massive than those of other theropod dinosaurs, in which the bones are usually hollow to make them more agile (like present day birds). Also, in this fossil the hind legs are actually much shorter in relation to the size of the animal than in any other theropod, leading some scientists to think that Spinosaurus was actually a quadrupedal animal. This has made some paleontologist think that maybe the lifestyle of the spinosaurs was much more similar to that of a crocodile and that they spent much more time living in water than on land, making the Spinosaurus the first known aquatic dinosaur.

Reconstruction by Rodrigo Vega of Spinosaurus based on the skeleton found in 2014.

Anyway, many paleontologists argue that the biology of a species cannot be based on a single fossil and advise caution when generalizing to the whole species (the fossil could belong to an adult and a juvenile that died together or could even come from an individual which had suffered some kind of embryonic malformation that kept its legs from developing normally). Paleontology is a science in which with every new discovery we can unravel the tree of life and the evolution of the different groups of living beings. With a little of luck, future discoveries will enable us to clarify the anatomy of Spinosaurus aegyptiacus and define the lifestyle of such a unique and extraordinary reptile.

REFERENCES

The following sources have been consulted in the elaboration of this entry:

• Cover photo: Davide Bonadonna.
• http://palaeos.com/vertebrates/theropoda/spinosauridae.html
• http://www.geol.umd.edu/~tholtz/G104/lectures/104therop.html
• http://palaeo.gly.bris.ac.uk/palaeofiles/fossilgroups/dinosauria/Theropoda.html
• http://www.scielo.br/pdf/aabc/v83n1/v83n1a06.pdf

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The kingdom of the reptiles: what is a dinosaur?

Dinosaurs (superorder Dinosauria, “terrible reptiles”) are a group of reptiles which dominated all terrestrial ecosystems during the Mesozoic (Secondary Era or “the Age of Reptiles”). Even today, to most people there’s still some confusion over what a dinosaur is and what is not, and the term “dinosaur” is often used to refer to all the reptiles that evolved during the Secondary Era. In this entry I’ll try to give account of some of the different groups of reptiles that appeared during the Mesozoic and I’ll explain the classification of the different dinosaurian groups and some of their adaptations.

MESOZOIC REPTILES: DINOSAURS AND LOTS MORE

The rise of the dinosaurs was possible thanks to a mass extinction phenomenon which occurred 251 million years ago (Permian-Triassic extinction event). That phenomenon annihilated up to 96% of marine species and up to 70% of terrestrial species in that time, leaving lots of empty ecological niches to be inherited by new animal species.

Modified graphic from Rohde & Muller (2005) showing the great massive extinction. The darker zone corresponds to the Mesozoic period.

During the Triassic period (in the early Mesozoic) many different groups of reptiles evolved. One of these groups was the Dinosauria, which at that moment was far from being the dominant group of terrestrial animals. Some other reptilian groups of that time were the terrestrial rauisuchians (clade Rauisuchia) and fully aquatic groups like the sauropterygians (superorder Sauropterygia) and the ichthyopterygians (superorder Ichthyopterygia).

Reconstructions by Dmitry Bogdanov of Prestosuchus (a rauisuchian, top), Nichollsia (a suropterygian, left bottom) and Platypterigius (an ichthyopterygian, right bottom).

A second mass extinction in the late Triassic and the early Jurassic put an end to most of the dominant reptile groups, allowing the yet small dinosaurs to expand and evolve, along with some new groups like the crocodilomorphs (superorder Crocodylomorpha, ancestors of crocodilians), the flying pterosaurs (order Pterosauria).

Reconstructions by Dmitry Bogdanov of Dakosaurus (a crocodilomorph, top) and Scaphognathus (a pterosauria, bottom).

As we can see, dinosaurs are only one of many reptile groups that evolved during the Mesozoic. During the Jurassic period, dinosaurs diversified into many different groups, but they were mostly restricted to terrestrial ecosystems, which they would rule until their practical extinction 65 million years ago at the end of the Cretacic period.

DINOSAUR CLASSIFICATION

The first dinosaurs evolved around 231 million years ago during the mid-Triassic period. They were small in size and were characterized by their limb’s posture, which contrary to most reptiles, grew vertically elevating their body from the ground. That gave them more agility and a more active lifestyle.

Top: Skeleton of Eoraptor, one of the oldest known dinosaurs (Museum of Japan, photo by Kentaro Ohno). Bottom: Representation of the posture common among most reptiles (left) and the posture characteristic of dinosaurs (right).

Then dinosaurs diverged into two different orders: the Saurischia and the Ornithischia. These two groups were distinguished by the structure of their pelvis; saurischians conserved a pelvis more closely similar to that of the other reptiles, while the ornithischians evolved a pelvis superficially similar to that of modern birds.

Representation showing the structure of saurischian hips (left) and ornitischian hips (right). The animals represented are facing left.

ORNITISCHIA: BIRD HIPPED

Evolutionary tree of Ornithischia, modified by Zureks.

Ornithopoda (“bird feet”): Ornithopods were the most diverse group of Ornithischia, characterized by their three-toed feet similar to that of birds. They were herbivores that could combine bipedal and quadruped walking. Among them we can find the Iguanodon, one of the first dinosaurs to be discovered by science.

Iguanodon feet (right) and reconstruction by O. C. Marsh (1896).

Ornithopods acquired many different adaptations; some groups had duck-like bills to feed on aquatic vegetation, others developed specialized hands with a sharp thumb and an opposable little finger to grasp the plants they fed on. Many groups developed bony crests which are thought to be used both for species identification and for communication between members of the same species.

Reconstruction of Parasaurolophus (missing author), an ornithopod which presented a big hollow crest to amplify the sounds it made.

Marginocephalia (“fringed heads”): The so-called marginocephalians were a group of herbivorous dinosaurs related to the ornithopods characterized by a great cranial ossification. These can be divided into two separated groups:

Pachycephalosaurians (suborder Pachycephalosauria, “thick-headed reptiles”) were bipeds which had an extremely thick skull and a series of lateral osteoderms (keratin-covered ossifications) flanking it. It is believed that pachycephalosaurians resolved territorial fights and disputed reproductive rights via head-ramming, similar to goats.

Reconstruction of Pachycephalosaurus by Jordan Mallon.

The other members of the group are the ceratopsians (suborder Ceratopsia, “horned faces”), quadrupeds which presented; neck frills making their skulls look bigger and the “rostral bone”, which formed a beak-shaped structure on the mouth. Lots of species also developed facial horns which could protrude from the cheek-bone, the eyebrow or the neck frill.

Reconstruction of Rubeosaurus by Lukas Panzarin (left) and skull of Triceratops (right), photo by Zachi Evenor.

Thyreophora (“shield bearers”): This basal group of ornithischians was exclusively composed of quadruped herbivores characterized by the presence of heavy osteoderms that constituted their main defence. This group can be divided into:

Stegosaurians (suborder Stegosauria, “roofed reptiles”) were big herbivorous dinosaurs characterized by having two rows of dorsal osteoderms from the neck to the tail, which served as protection and helped them in their thermoregulation. Some species also developed caudal spines called “thagomizers” used as weapons to defend themselves from predators.

Mounted “thagomizer” at Denver Museum of Nature and Science (left) and reconstruction of Stegosaurus by Nobu Tamura (right).

Anchylosaurians (suborder Ankylosauria, “fused reptiles”) developed heavy bony armours that covered most of the body. Some of them, like the Ankylosaurus, developed big bony clubs at the end of the tail to fend off predators.

Reconstruction of Euoplocephalus by Nobu Tamura.

SAURISCHIA: REPTILE HIPPED

Evolutionary tree of Saurischia, modified by Zureks.

Sauropodomorpha (“reptile-shaped feet”): The sauropodomorphs are better known as the “long-necked dinosaurs”. That’s because they adapted to feed on the highest strata of vegetation.

Reconstruction of different sauropodomorphs (left to right): Camarasaurus, Brachiosaurus, Giraffatitan and Euhelopus.

Most species became large quadrupeds, with pillar-like legs similar to those of elephants and long necks to reach the leaves of the highest trees. Later species reached tremendous sizes, like the Amphicoelias which could grow up to 60 metres long.

Theropoda (“beast feet”): This last group is mostly known for two reasons. First of all is that this group includes some taxons of great predators like the Tyrannosauridae and Dromeosauridae families. The second reason is that theropods are the only dinosaurian group that includes living species, because modern birds are included in the suborder Theropoda.

Skeleton of Allosaurus from american museum collections (1915).

All theropods are bipedal and most of the Mesozoic species were carnivorous, with sharp replaceable teeth adapted to predation. Theropods present a saurischian pelvis but later on, birds evolved a hip structure more similar to that found in ornitischian dinosaurs.

Reconstruction by Davide Bonnadonna of the different clades that led to the aparition of birds (left to right): Neotheropoda, Tetanurae, Coelurosauria, Paraves and finally the Archaeopteryx, believed to be the first bird species that ever existed.

Some species had feathers to help thermoregulation. Birds from these groups evolved at the end of the Jurassic period.

REFERENCES

The following sources have been consulted in the elaboration of this entry:

• Cover photo: Gerhard Boeggemann.
• http://web.archive.org/web/20110612180650/http://www.hmnh.org/library/diplodocus/holland1910.html
• http://www.smithsonianmag.com/science-nature/tracking-the-origin-of-dinosaurs-70638045/?no-ist
• http://www.geol.umd.edu/~tholtz/G104/104Syl.html
• http://palaeo.gly.bris.ac.uk/palaeofiles/fossilgroups/dinosauria/index.html

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