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.

Sinàpsids: Abans que els dinosaures dominessin la Terra

Abans de que els dinosaures dominessin la Terra, a finals del Paleozoic la terra estava ocupada pels sinàpsids. Els sinàpsids (la línia d’amniotes que inclou als mamífers) van ser un grup molt exitós que va ocupar la majoria de nínxols des del final del Carbonífer i fins al Pèrmic, però al final del Paleozoic la majoria de famílies es van extingir a causa de l’extinció masiva del Pèrmic-Triàssic (fa uns 252 milions d’anys) de la qual només els mamífers van sobreviure fins a l’actualitat. En aquesta entrada veurem alguns dels grups de sinàpsids més peculiars, els quals van portar a l’evolució dels mamífers actuals.

CARACTERÍSTIQUES I TENDÈNCIES EVOLUTIVES

El clade Synapsida inclou als mamífers i a tots els altres amniotes més emparentats amb aquests que amb els rèptils. Els sinàpsids van ser els primers amniotes en diversificar-se i van aparèixer fa uns 320 milions d’anys, a meitats del Carbonífer. Aquests primers sinàpsids es caracteritzaven per la presència d’una única fenestra temporal darrera de cada òrbita, a través de la qual passen els músculs mandibulars. Synapsida vol dir literalment “arcs fusionats” fent referencia als arcs zigomàtics (perquè abans els científics creien que els sinàpsids havien evolucionat de rèptils diàpsids i per tant es creia que els seus arcs s’havien “fusionat”).

Dibuix d’un crani de Archaeothyris, en el que podem veure algunes de les característiques dels sinàpsids, com la fenestra temporal i dents caniniformes. Dibuix de Gretarsson.

Altres característiques que van aparèixer al llarg de la seva evolució són:

• Diferenciació de dents amb formes diferents (heterodonts).
• Mandíbula inferior formada per menys ossos.
• Adquisició d’una postura més erecte i un metabolisme endoterm.

Els primers grups de sinàpsids més primitiu i “reptilians” s’anomenen informalment pelicosaures, mentres que les formes més avançades s’anomenen teràpsids (clade Therapsida que de fet, van evolucionar dels pelicosaures). Com veurem, l’evolució dels sinàpsids és del tipus de “un grup que inclou al següent grup el qual inclou al següent grup”.

Arbre evolutiu dels amniotes modificat de Kenneth D. Angielczyk (2009).

ORÍGEN DELS SINÀPSIDS, ELS PELICOSAURES

Reconstrucció de Cotylorhynchus, un caseasaure que creixia fins als 3 metres de llarg. Dibuix de Dmitry Bogdanov.

Els primers sinàpsids presentaven potes que els sortien dels costats, cossos baixos que arrossegaven per terra i probablement eren ectoterms. Si ens fixem en la morfología cranial, els primers grups de sinàpsids van ser els caseasaures (clade Caseasauria) caracteritzats pels seus caps petits, un musell superdesenvolupat i els seus cossos enormes (probablement eren criatures lentes i ecotèrmiques). Tanmateix, si ens fixem en l’esquelet postcranial, els primers sinàpsids foren dos grups anomenats varanòpids i ofiacodòntids (famílies Varanopidae i Ophiacodontidae) els quals s’assemblaven als varans (evolució convergent) i, mentre que els primers eren animals força petits i àgils, els segons van desenvolupar formes més grans amb caps enormes.

Dibuixos del varanòpid Varanodon (superior) i l’ofiacodòntid Ophiacodon (inferior). Dibuixos de Dmitry Bogdanov.

Just abans de l’aparició dels teràpsids (més avançats), els dos últims grups de “pelicosaures” van evolucionar i van ocupar la majoria d’ecosistemes terrestres. Ambdós grups compartien una alta vela que els recorria l’esquena (semblant a la del Spinosaurus) formada per les espines neurals. Durant la vida, aquesta vela probablement estaria coberta de pell i tindria força vasos sanguinis. Tot i que es creu que aquests animals encara eren ectoterms, aquesta vela probablement s’utilitzava per guanyar o perdre calor més eficaçment.

Reconstrucció de diferents espècies d’edafosàurids del gènere Ianthasaurus, mostrant la seva vela característica. Dibuixos de Dmitry Bogdanov.

El primer d’aquests grups és la família Edaphosauridae. A diferència de la majoria de sinàpsids basals, els edafosàurids eren herbívors i juntament amb els caseasaures, foren dels primers grans amniotes en adoptar un estil de vida vegetarià. Les veles dels edafosàurids estaven cobertes de tubercles espinosos, la funció dels quals encara no està clara del tot.

Esquelet de Edaphosaurus del Field Museum de Chicago, on es poden veure els tubercles a les espines neurals. Imatge de Andrew Y. Huang (2011).

L’altre grup, la família Sphenacodontidae, era el grup germà dels teràpsids, dintre del clade Sphenacodontia. Mentre que la resta de clades de pelicosaures tenien les dents pobrament fixades a les mandíbules, els esfenacodonts tenien les dents profundament insertades. La majoria d’esfenacodòntids eren carnívors, amb fortes mandíbules i dents caniniformes ben desenvolupades. Algunes espècies van esdevenir els majors depredadors terrestres just abans de l’aparició dels teràpsids.

Reconstrucció de l’esfenacodòntid Dimetrodon, de Dmitry Bogdanov.

ELS PRIMERS TERÀPSIDS

Esquelet de Biarmosuchus, un teràpsid basal en el qual podem veure la seva postura més erecte. Imatge de Ghedoghedo.

Els teràpsids (clade Therapsida, “arcs de bèsties”) van aparèixer fa uns 275 milions d’anys i van reemplaçar als pelicosaures com a animals terrestres dominants a mitjans del Pèrmic. Els primers teràpsids ja tenien una postura més erecte, a diferència de les extremitats laterals dels pelicosaures. A més, les seves fenestres temporals eren més grans, fent les seves mandíbules més potents.

Reconstrucció de Estemmenosuchus, un dinocèfal del qual s’han trobat impresions fòssils de la pell, pel que se sap que estava cobert de pell llisa i glandular sense escames. Dibuix de Mojcaj.

Els teràpsids es van diversificar àmpliament i van desenvolupar algunes adaptacions extraordinàries. Els dinocèfals (clade Dinocephalia, “caps terribles”) van desenvolupar protuberàncies cefàliques òssies que es creu estaven relacionades amb algún tipus de comportament de competició entre mascles. Un altre grup, els anomodonts (clade Anomodontia, “dents anormals”), es caracteritzava per no tindre dents, excepte un parell de canins superiors (que probablement estaven coberts per un bec). Els anomodonts fóren el grup germà dels teriodonts.

Reconstrucció de Placerias, un anomodont que podia arribar a pesar fins a una tona. Dibuix de Dmitry Bogdanov.

THERIODONTIA I ELS PRIMERS DENTS DE SABRE

Els teriodonts (clade Theriodontia “dents de bèstia”) van esdevenir el grup de sinàpsids més exitós. Els tres grups principals probablement s’assemblaven força als mamífers, amb postures totalment erects, un paladar ossi secondari que els permetia respirar mentre empassaven o subjectaven una presa i dents heterodonts (dents incisiviformes, caniniformes i molariformes). El grup de teriodonts més primitius eren els gorgonopsis (clade Gorgonopsia, família Gorgonopsidae). Tots els membres d’aquest grup eren carnívors i depredadors actius, com revel·len les seves dents de sabre. Tot i que la majoria eren de mides modestes, els més grans arribaven als 3 metres de llarg i tenien canins de fins a 15 centímetres.

Reconstrucció de Inostrancevia, el gènere de gorgonòpsid més gran, depredant a Scutosaurus, un pararèptil. Dibuix de Dmitry Bogdanov.

Un segon grup, els terocèfals (clade Therocephalia “cap de bèstia”) eren força més avançats que els gorgonòpsids tot i que no van adquirir les mides dels seus cosins. Les seves potes s’assemblaven als dels primers mamífers, presentaven petits orificis als ossos que probablement aguantaven vibrises en llavis carnosos i tot apunta a que ja eren endoterms.

Reconstrucció d’un parell de Pristerognathus, un gènere de terocèfals en els quals podem veure algunes característiques més mamíferes. Dibuix de Dmitry Bogdanov.

Tant els gorgonòpsids com els terocèfals desapareixeren a finals del Pèrmic. L’únic grup de teràpsids que va sobreviure al llarg del Mesozoic i que coexistí amb els dinosaures van ser els cinodonts.

PETITS CINODONTS

Els cinodonts (clade Cynodontia “dents de gos”) van aparèixer a finals del Pèrmic i es van diversificar àmpliament juntament amb els arcosaures. Tot i que no està demostrat, la majoria de paleoartistes representen als cinodonts coberts de pèl, ja que les proves suggereixen un metabolisme endoterm. Algunes característiques dels cinodonts eren:

• Mandíbula inferior formada únicament per l’òs dentari, mentre que els altres òssos mandibulars van esdevenir els ossets de l’oïda (l’articular, el quadrat i l’angular van evolucionar en el martell, l’enclusa i l’estrep).
• Dents complexes: incissius per subjectar, canins per perforar, i premolars i molars per mastegar.
• Només dos jocs de dents (difiodonts), enlloc de dents que es renoven constantment (polifiodonts, com la majoria de rèptils).
• Grans cavitats cerebrals. S’han trobat alguns caus fòssils de diferents cinodonts, revel·lant comportaments socials complexes.

Reconstrucció de Thrinaxodon, un cinodont excavador amb bigotis i pèl. Imatge de Nobu Tamura.

Tot i que competien amb els arcosaures, algunes formes primitives es van fer força grans. Per exemple, alguns carnívors com Cynognathus tenien caps grans i medien 1 metre de llarg, mentre Trucidocynodon tenia la mida d’un lleopard. Tanmateix, la tendència evolutiva faria que els cinodonts es féssin cada cop més petits, com la familia Brasiliodontidae la qual, com la majoria de cinodonts, va viure a l’ombra dels dinosaures i altres rèptils més grans. Els brasiliodontids es creu que foren el grup germà dels Mammaliaformes (els mamífers i els seus parents més recents).

Reconstrucció de Brasilitherium, un dels cinodonts no mamífers més avançats, que només media 12 cm de llarg. Dibuix de Smokeybjb.

Finalment, els mamífers van aparèixer a finals del període Triàsic, fa uns 225 milions d’anys. Els primers mamaliaformes probablement eren animals insectívors, nocturns i semblants a les musaranyes. Es creu que aquest estil de vida nocturn és el que va promoure l’aparició del pelatge, ja que en els teràpsids l’endotèrmia va aparèixer abans que el pèl. Aquests mamaliaformes probablement tinguéssin glàndules mamàries per alimentar a les seves críes quan aquestes no tenien dents, però probablement no tinguéssin mugrons com els monotremes actuals.

Reconstrucció de Megazostrodon, un petit mamaliaforme que representa força bé la transició de cinodonts a mamífers moderns. Imatge de Udo Schröter.

Després de l’extinció de la majoria d’arcosaures a finals del període Cretàcic, els sinàpsids supervivents van ocupar els nínxols ecològics buits. Els mamífers han dominat el món des de llavors, conquerint la terra, el mar i fins i tot l’aire, però això no hagués estat possible sense les diferents adaptacions adquirides pels primers sinàpsids al llarg de la seva evolució. Gràcies a ells, els humans i tots els altres mamífers som actualmente els animals dominants del planeta.

REFERÈNCIES

Durant l’elaboració d’aquesta entrada s’han consultat les següents fonts:

• 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/
• Imatge de portada de Mauricio Antón.

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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