Arxiu d'etiquetes: crocodile

Reptiles and mammals: same origin, different stories

Did mammals evolve from reptiles? The truth is they didn’t. Reptiles and mammals both have independent evolutionary histories that separated soon after the apparition of the so-called amniotic egg, which allowed the babies of these animals to be born outside of water. Previously, we talked about the origin of vertebrates and about how they managed to get out of the sea to start walking on land for the first time. In this entry we’ll explain how the ancestors of reptiles and mammals, the AMNIOTES, became independent of the aquatic medium and became the dominant land animals.

THE AMNIOTIC EGG

The characteristic that unites reptiles and mammals in the same group is the amniotic egg. While amphibian eggs are relatively small and only have one inner membrane, the eggs of amniotes are much bigger and present various membranes protecting the embryo and keeping it in an aqueous medium. The outer layer is the eggshell which, apart from offering physical protection to the embryo, prevents water loss and its porosity allows gas interchange. Beneath the eggshell we can find the next membranes:

512px-Crocodile_Egg_Diagram.svgDiagram of a crocodile egg: 1. eggshell 2. yolk sac 3. yolk (nutrients) 4. vessels 5. amnion 6. chorion 7. air 8. alantois 9. albumin (white of the egg) 10. amniotic sac 11. embryo 12. amniotic fluid. Image by Amelia P.
  • Chorion: The first inner membrane, which offers protection and, together with the amnion, forms the amniotic sac. Also, being in contact with the eggshell, it participates in gas interchange, bringing oxygen from the outside to the embryo and carbon dioxide from the embryo to the outside.
  • Amnion: Membrane that surrounds the embryo and constitutes a part of the amniotic sac. It offers an aqueous medium for the embryo and connects it with the yolk sac (a structure that brings food and that is also found in fish and amphibians).
  • Allantois: The third layer, it is used as a storage for nitrogen waste products, and together with the chorion, helps in gas interchange.
512px-Amphibian_Egg_Diagram.svgDiagram of an amphibian egg: 1. jelly capsule 2. vitelline membrane 3. perivitelline fluid 4. yolk 5. embryo. Image by Separe3g.

All these different kinds of membranes eliminate the need amphibians had of laying their eggs in water. Also, unlike amphibians, amniotes don’t go through a gilled larval stage, but are instead born as miniature adults, with lungs and legs (at least those that have them). All these made the first amniotes completely independent of the aquatic medium.

AMNIOTE ORIGINS

The first amniotes evolved around 312 million years ago from reptiliomorph tetrapods. At the end of the Carboniferous period lots of tropical forests where the great primitive amphibians lived disappeared, leaving a colder and drier climate. This ended with many of the big amphibians of that time, allowing the amniotes to occupy new habitats.

Solenodonsaurus1DBReconstruction of Solenodonsaurus janenschi, one of the candidates in being the first amniote, which lived around 320-305 million years ago in what is now the Czech Republic. Reconstruction by Dmitry Bogdanov.

CHARACTERISTICS

These early amniotes had a series of characteristics that set them apart from their semiaquatic ancestors:

  • Horny claws (amphibians don’t have claws) and keratinized skin that prevents water loss.
  • Bigger large intestine and higher density of renal tubules to increase water reabsorption.
  • Specialized lacrimal glands and a third membrane in the eye (nictitating membrane) which keep the eye wet.
  • Larger lungs.
  • Loss of the lateral line (sensory organ present in fish and amphibians).

The skeleton and musculature also evolved offering better mobility and agility on a terrestrial medium. The first amniotes presented ribs that encircled their body converging at the sternum, making their inner organs more secure, and a series of muscular receptors offered them better agility and coordination during locomotion.

AMNIOTE SKULLS

Traditionally, the different amniotes were classified based on the structure of their cranium. The characteristic used to classify them was the presence of temporal openings (fenestrae), by which we have three groups:

  • Anapsids (“no arches”): No temporal openings (turtles).
Skull_anapsida_1Diagram of an anapsid skull, by Preto(m).
  • Synapsids (“fused arches”): With only one temporal opening (mammals).
Skull_synapsida_1Diagram of a synapsid skull, by Preto(m).
  • Diapsids (“two arches”): With two temporal openings (reptiles, including birds).
Skull_diapsida_1Diagram of a diapsid skull, by Preto(m).

Previously it was believed that the first amniotes presented an anapsid skull (without openings, like turtles) and that subsequently they separated into synapsids and diapsids (the temporal openings formed “arches” that offered new anchor points for the jaw’s musculature). Yet, it has been discovered that this three-group classification is not valid.

Even though we still believe that the first amniotes were anapsid, it is currently known that these, soon after their apparition, separated into two different lineages: the synapsids (clade Synapsida) and the sauropsids (clade Sauropsida).

SYNAPSIDA

This lineage includes mammals and their amniote ancestors. Even though the first synapsids like Archaeothyris looked externally like lizards, they were more closely related to mammals, as they shared one temporal fenestrae where the jaw muscles passed through.

Archaeothyris.svgDrawing of the skull of Archaeothyris, which is thougth to be one of the first synapsids that lived around 306 million years ago in Nova Scotia. Drawing by Gretarsson.

The ancestors of mammals were previously known as “mammal-like reptiles”, as it was thought that mammals had evolved from primitive reptiles. Currently it’s accepted that synapsids form a different lineage independent of reptiles, and that they share a series of evolutionary trends that makes them closer to modern mammals: the apparition of different kinds of teeth, a mandible made of one single bone, the vertical posture of their limbs, etc…

Dimetrodon_grandisReconstruction of Dimetrodon grandis, one of the better known synapsids, from about 280 million years ago. Reconstruction by Dmitry Bogdanov.

Even though most modern mammals don’t lay eggs and give birth to live offspring, all groups maintain the amniote’s three characteristic membranes (amnion, chorion and allantois) during embryonic development.

SAUROPSIDA

Sauropsids include current reptiles and their amniote ancestors. Currently, in many scientific papers the word “sauropsid” is used instead of “reptile” when discussing phylogenies, as the sauropsids also includes birds. The first sauropsids were probably anapsids, and soon after their appearance they separated into two groups: the Parareptilia which conserved anapsid skull, and the Eureptilia which include the diapsids (current reptiles and birds).

Traditional_ReptiliaEvolutionary tree of current vertebrates, in which green color marks the groups previously included inside reptiles. As you can see, the traditional conception of "reptile" includes the ancestors of mammals and excludes birds. Image by Petter Bøckman.

Diapsids are currently the most diversified group of land vertebrates. They diversified greatly in the late Permian period (about 254 million years ago), just before the Mesozoic (the Age of Reptiles). These can be divided into two main groups: the Lepidsaurs and the Archosaurs, both with representatives in our days.

LEPIDOSAURIA: SMALL AND PLENTIFUL

Lepidosaurs (literally “reptiles with scales”) appeared in the early Triassic (around 247 million years ago) and, even if most of them didn’t grow to big sizes, they are currently the largest group of non-avian reptiles. These are characterized by presenting a transversal cloacal slit, by having overlapping scales and shedding their skin whole or in patches and by other skeletal characters.

Rat_Snake_moulted_skinShed skin of a rat snake. Photo by Mylittlefinger.

The current lepidosaurs belong to one of two different orders:

  • Order Rhynchocephalia: That includes the two species of tuatara. Currently endangered, they are considered living fossils because they present skulls and characteristics similar to the Mesozoic diapsids.
Sphenodon_punctatus_(5)Photo of a tuatara (Sphenodon punctatus), by Tim Vickers.
  • Order Squamata: Current squamates include iguanas, chameleons, geckoes, skinks, snakes and other legless lizards. With more than 9000 living species, squamates are a large group with a wide array of adaptations and survival strategies.
Sin títuloPhotos of some squamates, from left to right and from top to bottom: Green iguana (Iguana iguana, by Cary Bass), king cobra (Ophiophaga Hannah, by Michael Allen Smith), Mexican mole lizard (Bipes biporus, by Marlin Harms) and Indian chameleon (Chamaeleo zeylanicus, by Shantanu Kuveskar).

ARCHOSAURIA: ANCIENT KINGS

Archosaurs (literally “ruling reptiles”) were the dominant group of land animals during the Mesozoic. These conquered all possible habitats until the extinction of most groups at the end of the Cretaceous period. Some of the extinct groups were the pseudosuchians (relatives of modern crocodiles, order Crocodylia), the pterosaurs (large flying reptiles) and the dinosaurs (excepting birds, clade Aves).

Massospondylus_Skull_Steveoc_86Drawing of the skull of the dinosaur Massospondylus in which we can see the different characteristic openings of diapsid archosaurs. Image by Steveoc 86.

As you see, both groups of modern archosaurs couldn’t be more different. Yet, crocodiles and birds share a common ancestor, and they are both more closely related with each other than with the rest of reptiles.

Yellow-billed_stork_kazingaPhoto of two species of modern arcosaurs: a Nile crocodile (Crocodylus niloticus) and a yellow-billed stork (Mycteria ibis). Photo by Tom Tarrant.

AND WHAT ABOUT TURTLES?

Turtles (order Testudines) have always been a group difficult to classify. Turtles are the only living amniotes with an anapsid skull, without any post-ocular opening. That’s why previously they had been classified as descendants of primitive amniotes (clade Anapsida, currently disused) or as primitive anapsid sauropsids (inside the Parareptilia clade)

KONICA MINOLTA DIGITAL CAMERASkeleton of the extinct tortoise Meiolania platyceps which lived in New Caledonia until 3000 years ago. In this photo it can be seen the compact cranium without openings. Photo by Fanny Schertzer.

Recent molecular studies have revealed that turtles are actually diapsids that lost their temporal openings secondarily. What still divides the scientific community is if testudines are more closely related to Lepidosauromorphs (lepidosaurs and their ancestors) or to Archosauromorphs (archosaurs and their ancestors).

Leopard_tortoiseIndividual leopard tortoise (Stigmochelys pardalis) from Tanzania. Photo by Charles J. Sharp.

As you have seen, the evolution of amniotes is an extremely complex matter. We hope that with this entry some concepts have been clarified:

  1. Mammals (synapsids) come from an evolutionary lineage different from that of reptiles (sauropsids).
  2. Sauropsids include traditional reptiles (lepidosaurs, archosaurs and turtes) and birds (inside archosaurs).
  3. There’s still so much to investigate about the placement of turtles (testudines) in the evolutionary tree of sauropsids.
Figure_29_04_03Modified diagram about the evolutionary relationships of the different amniote groups.

REFERENCES

During the elaboration of this entry the following sources have been consulted:

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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.

Sin título
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).
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).
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.

eorpohd
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.

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

ORNITISCHIA: BIRD HIPPED

ORNITHISCHIA
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.

IGUANODOOOONT
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.

parasauro
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.

Pachycephalosauria_jmallon
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.

CERAAA
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.

STEGo
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.

Euoplocephalus_BW
Reconstruction of Euoplocephalus by Nobu Tamura.

SAURISCHIA: REPTILE HIPPED

SAURISCHIA
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.

Macronaria_scrubbed_enh
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.

AllosaurusAMNH5753
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.

Davide-Bonnadonna-theropod-size-evolution-600-px-tiny-July-August-Darren-Naish-Tetrapod-Zoology
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:

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