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November 7, 2014

University of the Witwatersrand

Through the careful study of modern and early fossil tortoise,researchers now have a better understanding of how tortoises breatheand the evolutionary processes that helped shape their unique breathingapparatus and tortoise shell.

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Origin of the unique ventilatory apparatus of turtles: How the tortoise'sribs got embedded in its shell

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Credit: Emma R. Schachner [Click to enlarge image]

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A Computed Tomography rendering of a snapping turtle (Chelydra serpentina)showing the skeleton (white), lungs (blue), and abdominal muscles (red and pink)used to ventilate the lungs. Because turtles have locked their ribs up into theiconic turtle shell, they can no longer use their ribs to breathe as in most otheranimals and instead have developed a unique abdominal muscle based system.

hrough the careful study of modern and early fossil tortoise,researchers now have a better understanding of how tortoises breatheand the evolutionary processes that helped shape their unique

breathing apparatus and tortoise shell. The findings published in a paper,titled: Origin of the unique ventilatory apparatus of turtles, in the scientificjournal, Nature Communications, on Friday, 7 November 2014, helpdetermine when and how the unique breathing apparatus of tortoises evolved.

Lead author Dr Tyler Lyson of Wits University'sEvolutionary Studies Institute, the SmithsonianInstitution and the Denver Museum of Nature andScience said: "Tortoises have a bizarre body plan andone of the more puzzling aspects to this body plan isthe fact that tortoises have locked their ribs up into theiconic tortoise shell. No other animal does this and thelikely reason is that ribs play such an important role inbreathing in most animals including mammals, birds,

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crocodilians, and lizards."

Instead tortoises have developed a unique abdominalmuscular sling that wraps around their lungs andorgans to help them breathe. When and how this mechanism evolved has beenunknown.

"It seemed pretty clear that the tortoise shell and breathing mechanism evolved intandem, but which happened first? It's a bit of the chicken or the egg causalitydilemma," Lyson said. By studying the anatomy and thin sections (also known ashistology), Lyson and his colleagues have shown that the modern tortoise breathingapparatus was already in place in the earliest fossil tortoise, an animal known asEunotosaurus africanus.

This animal lived in South Africa 260 million years ago and shares many uniquefeatures with modern day tortoises, but lacked a shell. A recognisable tortoise shelldoes not appear for another 50 million years.

Lyson said Eunotosaurus bridges the morphological gap between the early reptilebody plan and the highly modified body plan of living tortoises, making it theArchaeopteryx of turtles.

"Named in 1892, Eunotosaurus is one of the earliest tortoise ancestors and is knownfrom early rocks near Beaufort West," said Professor Bruce Rubidge, Director of theEvolutionary Studies Institute at Wits University and co-author of the paper.

"There are some 50 specimen of Eunotosaurus. The rocks of the Karoo are remarkablein the diversity of fossils of early tortoises they have produced. The fact that we findEunotosaurus at the base of the Karoo succession strongly suggest that there aremore ancestral forms of tortoises still to be discovered in the Karoo," Rubidge added.

The study suggests that early in the evolution of the tortoise body plan a gradualincrease in body wall rigidity produced a division of function between the ribs andabdominal respiratory muscles. As the ribs broadened and stiffened the torso, theybecame less effective for breathing which caused the abdominal muscles to becomespecialised for breathing, which in turn freed up the ribs to eventually -- approximately50 million years later -- to become fully integrated into the characteristic tortoise shell.

Lyson and his colleagues now plan to investigate reasons why the ribs of earlytortoises starting to broaden in the first place. "Broadened ribs are the first step in thegeneral increase in body wall rigidity of early basal tortoises, which ultimately leads toboth the evolution of the tortoise shell and this unique way of breathing. We plan tostudy this key aspect to get a better understanding why the ribs started to broaden."

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The above story is based on materials provided by University of the Witwatersrand.Note: Materials may be edited for content and length.

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University of the Witwatersrand. "Origin of the unique ventilatory apparatus ofturtles: How the tortoise's ribs got embedded in its shell." ScienceDaily.ScienceDaily, 7 November 2014..

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Journal Reference:

1. Tyler R. Lyson, Emma R. Schachner, Jennifer Botha-Brink, Torsten M. Scheyer,Markus Lambertz, G. S. Bever, Bruce S. Rubidge, Kevin de Queiroz. Origin of theunique ventilatory apparatus of turtles. Nature Communications, 2014; 5: 5211DOI: 10.1038/ncomms6211

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