Arxiu d'etiquetes: carapace

Shell evolution with just four fossil turtles

Turtles are charming animals yet, while they look cute to most people, they’ve been racking the brains of palaeontologists for decades. The combination of apparently primitive features and an extremely specialized anatomy, has made the reconstruction of the origin and evolution of these reptiles a nearly impossible task. In this entry we’ll try to get a general idea about the evolution of one of the most striking characteristics of turtles (the shell) with only four examples of primitive “turtles”.

CURRENT AND EXTINCT RELATIVES

As we explained in an earlier entry, the origin of turtles is still debated among the scientific community. Turtles show some anatomic characteristics not found among any current vertebrate, which makes their phylogenetic origin confusing. One of the characteristics that has puzzled palaeontologist more is their skull.

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Skull of a loggerhead sea turtle (Caretta caretta) in which we can see the lack of temporal openings. Photo by David Stang.

While the rest of reptiles are diapsid (they present a pair of temporal openings at each side of the skull), turtles present a typically anapsid cranium (without any temporal openings). Yet, recent genomic studies have proved that it’s more likely that testudines (order Testudines, current turtles) descend from a diapsid ancestor and that through their evolution they reverted back to the primitive anapsid form. What is not so clear is if turtles are more closely related to lepidosaurs (lizards, snakes and tuataras) or to archosaurs (crocodiles and birds). The most accepted hypothesis is the second one.

Even if the origins of the testudines are still somewhat mysterious, most palaeontologists coincide in that they belong to the clade Pantestudines, which groups all those species more closely related to turtles than to any other animal. A group of reptiles that are also found inside the pantestudines are the sauropterygians like plesiosaurs and placodonts.

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Reconstruction by Dmitry Bogdanov of the sauropterygian Plesiosaurus, a distant relative of turtles.

EVOLUTION OF TESTUDINES

The rest of pantestudines help us to form an image of how turtles acquired such a specialized anatomy. But first, take a look at some of the characteristics of turtles:

  • A shell made up of two parts: the upper shell (carapace) which comes from the fusion of the vertebrae and the dorsal ribs and the lower shell (plastron) that originates from ventral ribs called “gastralia” (still present in some current reptiles).
  • While the rest of vertebrates present the scapula over their ribs, the turtle’s ribs (their carapace) cover the scapula.
  • The ability to hide their heads and limbs in their shells.
  • The absence of teeth; having instead horny ridges in their jaws.

As we’ll see, these characteristics were acquired very gradually.

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The carapace of a dead turtle, in which we can see how the ribs fuse with the vertebrae to form the shell. Photo by Fritz Flohr Reynolds.

Even if their relationship with turtles isn’t still very clear, Eunotosaurus africanus is the most ancient candidate to being a turtle’s relative. Eunotosaurus was a fossorial animal that lived 260 million years ago in South Africa. This animal had very wide dorsal ribs which contacted each other, which is thought to have served as an anchoring point for powerful leg muscles, used while digging. Also, similarly to current turtles, Eunotosaurus had lost the intercostal muscles and presented a reorganization of the respiratory musculature.

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Fossil of Eunotosaurus, in which the characteristically wide ribs can be seen. Photo by Flowcomm.

The oldest indisputable relative of turtles is Pappochelys rosiane from Germany (240 million years ago). The name “Pappochelys” literally means “grandfather turtle” as, before the discovery of Eunotosaurus it was the oldest turtle relative. Just like Eunotosaurus, it presented wide dorsal ribs in contact with each other. Also, its ventral ribs were already wider and thicker and its scapular girdle was placed below the dorsal ribs.

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Drawing by Rainer Schoch of the skeleton of Pappochelys in which we can see some of its characteristics. It is believed that Pappochelys was a semiaquatic animal that swam with the aid of its long tail.

The next step in the evolution of turtles is found 220 million years ago, during the late Triassic in China. Its name is Odontochely semitestacea, which means “toothed turtle with half a shell”. This name is due to the fact that, unlike true turtles, Odontochelys still had a mouth full of teeth and it only presented the lower half of the shell, the plastron. Even if it also had thick dorsal ribs, only paleontological proofs of the plastron have been found. Odontochelys was discovered in freshwater deposits, leads us to believe that at first it only developed the plastron to protect itself from predators attacking from below.

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Reconstruction by Nobu Tamura of Odontochelys semitestacea. It’s not considered to be a true turtle due to the fact that it only had half a shell.

The first testudine known to possess a complete shell is Proganochelys quenstedti from the Triassic period, 210 million years ago. It already presented many characteristics found in current turtles: the shell was completely formed, with carapace and plastron, its skull was anapsid looking and it had no teeth. However, Proganochelys wasn’t able to retract its head and legs inside its shell (even if this may be because of the horns it had). It also presented two extra shell pieces at both sides, which probably served to protect its legs.

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Reconstruction of Proganochelys from the Museum am Lowentor of Stuttgart. Photo by Ghedoghedo.

PRESENT DAY TURTLES

The order Testudines as we know it, appeared around 190 million years ago, during the Jurassic period. These current turtles are classified into two different suborders, which both separated quickly at the beginning of the evolution of testudines:

Suborder Pleurodira: This suborder is the smallest one as it only contains three current families, all native from the southern hemisphere. The main characteristic is the form in which they retract their neck laterally inside their shell, which leaves the neck exposed and makes the cervical vertebrae present a characteristic shape (Pleurodira roughly means “side neck”). Also, pleurodirans present 13 scutes in their plastrons.

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Photo by Ian Sutton of an eastern long-necked turtle (Chelodina longicollis), a typical pleurodiran.

Suborder Cryptodira: Cryptodirans comprise most turtles. While pleurodirans only include freshwater species (as the testudines common ancestor is thought to be), criptodirans include freshwater terrapins, terrestrial tortoises and sea turtles. Apart from only presenting between 11 and 12 scutes in their plastrons, their principal characteristic is the ability to retract their neck and to hide their heads completely in their shell (Cryptodira roughly means “hidden neck”). Cryptodirans are found in practically all the continents and oceans (except in the coldest habitats).

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Photo of an Alabama red-bellied turtle (Pseudemys alabamensis) by the U.S. Fish and Wildlife Service. In this photo we can see how cryptodirans hide their heads.

Even if there still are some questions to be answered about the evolution of turtles, we hope that with this little introduction to some of the most characteristic fossil “turtles”, you have had an overall view about how turtles got their shells. Whatever their origins are, we hope that the apparition of men isn’t what puts an end to the history of this group of slow but steady creatures.

REFERENCES

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

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How is the life of a marine turtle?

I have talked about marine turtles in some past posts. In concrete, about the loggerhead sea turtle (Caretta caretta). In the following weeks, I am going to talk more about this amazing marine animals. In particular, this week I will explain how is the life of a marine turtle, especially about the loggerhead sea turtle, and in the next one, I am talking about which are the threats that endanger these animals and about what we can do to save them. 

INTRODUCTION

Loggerhead sea turtle is one of the seven sea turtles on Earth. It has a worldwide distribution, being the most abundant species in the Mediterranean, and it can be identified by the presence of a carapace that measures between 80 and 100 cm long with 5 lateral scutes, so that the first of them is in contact with the nuchal scute. It is endangered according to IUCN (International Union for the Conservation of Nature). The loggerhead sea turtle feeds on jelly plankton like jellyfishes during the oceanic stage, but feeds on fishes and squids in the neritic stage. Additionally, they can consume salt water due to the presence of salt glands placed in the cranium. Like other sea turtles, it cannot hide its head and fins inside the carapace.

Claus d'identificació de la tortuga babaua (Caretta caretta) (Foto extreta de MarineBio).
Identification key for a loggerhead sea turtle (Caretta caretta) (Picture from MarineBio).

HOW IS THE LIFE OF A MARINE TURTLE?

In marine turtles, the reproductive cycles are circadian, it is that it happens regularly over the time. The periodicity depends on each species, but in the case of the loggerhead sea turtle usually is biannual, so it takes place every two years (but sometimes every three years). Anyway, this cycle is not strict because it is dependant on some factors like food availability or illnesses.

The gregarious behaviour of many species is explained for the ability to recognise the individuals of the same species. In order to recognise each other, most of the species use smell, but they can use sight and sound. During the mating, when the female accept the male, the male bites the female in the neck and in the anterior fins. The male put itself on the female and catches her with the nails of the anterior fins (in the case of the loggerhead, it has two nails per fin). Mating takes place in the sea and usually in the first hours of the day. Moreover, a female can be impregnated by several males.

Aparellament de la tortuga babaua (Caretta caretta) (Foto: OceanWide Images).
Mating of a loggerhead sea turtle (Caretta caretta) (Picture: OceanWide Images).

The moment when the marine turtles lay the eggs depend on the moon phases, tides, temperature and wind, but it usually happens during summer in sandy beaches. Females return to the beaches where they were born, coming from feeding grounds. They navigate using marine currents, temperature changes, magnetic signals and the sound and smell of the beach.

Depending on the features of the beach, this will be more or less suitable for each marine turtle species. The loggerhead prefers open and shallow beaches and bays, continental or insular, with a slope between 5-10º and with a calm swell. Moreover, these beaches have to be protected by bushes in the terrestrial part and by coral or rockery reefs in the marine part. They usually lay on the first terrace of the beach, in zones without plants and in the first attempt. All the sea turtles have in common the fact that the lay is done beyond the highest tide.

When the female finds the place, with the anterior fins do a cavity where to place its body (called bed) and next, with the posterior fins, dig out the nest and place the eggs. During the period from which the female leave the water and dig out the nest, the animal is very sensitive to bother and can stop doing the nest and come back to water. 

Sea turtles do several lays per year. In the case of loggerheads, they usually do between 2 and 4 lays per year, with 100 eggs that weights 40 grammes (this is 4 kilos per lay). Despite of this, we have to have in consideration that the number of eggs produced for a female is limited by the capacity of storage of the female, which is related with the size. Between each lay in the same reproductive cycle, the mating is not necessary because they can store sperm.

Tortuga en la fase de posta dels ous (Foto: Brandon Cole).
Turtle laying the eggs in a beach (Picture: Brandon Cole).

Eggs are incubated during 50-60 days under the sand of the beach (in the loggerhead). The hatching is synchronized and when the small turtles reach the surface in few minutes are oriented thanks to the beach slope, the sound of the waves and the light of the moon on the sea.

Cria d'una tortuga babaua (Caretta caretta) sortint de l'ou (Foto: Rewilding Europe).
Loggerhead sea turtle baby (Caretta caretta) (Picture: Rewilding Europe).

During the first days of life, turtles present a high buoyancy. In the first weeks, small turtles remain in marine currents and gyres, where food is abundant, so they have a pelagic life. If they are male, the most probable is that they will never touch the land. 

When they are born, the carapace is soft  and, for this reason, the number of individuals that will survive is just a 10% of which leave the egg due to the predators, like crabs, sharks and seagulls. During the first year only survives 10-30% of the animals. Year after year, the mortality rate decreases for the increase of size and the hardening of the carapace. A study has found that just 10 out of every 10.000 eggs will become adults and just one will die for age.

Adult de tortuga babaua (Caretta caretta) (Foto: Deviant Art).
Adult of a loggerhead sea turtle (Caretta caretta) (Picture: Deviant Art).

Sea turtles do long-distance migrations, specially in the young stage. When they abandon the beach where they were born, during the next 10 years, they will be travelling long distances. The migrations are between feeding and reproductive grounds.

Then, the cycle restarts with the newborns.

REFERENCES

  • Cardona L, Álvarez de Quevedo I, Borrell A, Aguilar A (2012). Massive Consumption of Gelatinous Plankton by Mediterranean Apex Predators. PLoS ONE 7(3): e31329. doi:10.1371/journal.pone.0031329
  • Consejería de Medio Ambiente de la Junta de Andalucía (2014). Varamientos de Especies Marinas Amenazadas. Guías prácticas voluntariado ambiental.
  • CRAM: Caretta caretta
  • Dodd, C. Kenneth, Jr. 1988. Synopsis of the biological data on the Loggerhead Sea Turtle Caretta caretta (Linnaeus 1758). U.S. Fish Wildl. Serv., Biol. Rep. 88(14). 110 pp.
  • IUCN: Caretta caretta 
  • Márquez, R (1996). Las tortugas marinas y nuestro tiempo. México: IEPSA
  • Smith, T & Smith R (2007). Ecología. Pearson Educación (6 ed.)

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