Were dinosaurs slow and lumbering, or quick and agile? It depends largely on whether they were cold- or warm-blooded.

When dinosaurs were first discovered in the mid-19th century, paleontologists thought they were plodding beasts that relied on their environment to keep warm, like modern-day reptiles.

But research during the last few decades suggests that they were faster creatures, nimble like the velociraptors or T. rex depicted in the movie Jurassic Park, requiring warmer, regulated body temperatures.

Now, researchers, led by Robert Eagle of the California Institute of Technology, have developed a new way of determining the body temperatures of dinosaurs for the first time, providing new insights into whether dinosaurs were cold- or warm-blooded.

"Eagle and colleagues have applied the newest and most innovative techniques to answering the question of whether dinosaurs were warm- or cold-blooded," says Lisa Boush, program director in the National Science Foundation's (NSF) Division of Earth Sciences, which funded the research.

"The team has made important strides in discovering that the body temperature of dinosaurs was close to that of mammals, and that the dinosaurs' physiology allowed them to regulate that temperature. The result has implications for our understanding of dinosaurs' ecology--and demise."

By analyzing the teeth of sauropods--long-tailed, long-necked dinosaurs that were the biggest land animals ever to have lived--the scientists found that these dinosaurs were about as warm as most modern mammals.

"This is like being able to stick a thermometer in an animal that has been extinct for 150 million years," says Eagle, a geochemist at Caltech and lead author of a paper to be published online in the journal Science Express.

"The consensus was that no one would ever measure dinosaur body temperatures, that it's impossible to do," says John Eiler, a co-author and geochemist at Caltech. But using a technique pioneered in Eiler's lab, the team did just that.

The researchers analyzed 11 teeth, unearthed up in Tanzania, Wyoming and Oklahoma, that belonged to the dinosaurs Brachiosaurus and Camarasaurus.

They found that Brachiosaurus had a temperature of about 38.2 degrees Celsius (100.8 degrees Fahrenheit) and Camarasaurus had one of about 35.7 degrees Celsius (96.3 degrees Fahrenheit), warmer than modern and extinct crocodiles and alligators, but cooler than birds.

The measurements are accurate to within one or two degrees Celsius.

"Nobody has used this approach to look at dinosaur body temperatures before, so our study provides a completely different angle on the long-standing debate about dinosaur physiology," Eagle says.

The fact that the temperatures were similar to those of most modern mammals might seem to imply that dinosaurs had a warm-blooded metabolism.

But, the researchers say, the issue is more complex. Because sauropod dinosaurs were so huge, they could retain their body heat much more efficiently than smaller mammals like humans.

"The body temperatures we've estimated provide key information that any model of dinosaur physiology has to be able to explain," says Aradhna Tripati, a co-author who's a geochemist at University of California, Los Angeles and visiting geochemist at Caltech. "As a result, the data can help scientists test physiological models to explain how these organisms lived." The measured temperatures are lower than what's predicted by some models of dinosaur body temperatures, suggesting there is something missing in scientists' understanding of dinosaur physiology.

These models imply that dinosaurs were so-called gigantotherms, that they maintained warm temperatures by their sheer size.

To explain the lower temperatures, the researchers suggest that dinosaurs could have had physiological or behavioral adaptations that allowed them to avoid getting too hot.

The dinosaurs could have had lower metabolic rates to reduce the amount of internal heat. They could also have had something like an air-sac system to dissipate heat.

Alternatively, they could have dispelled heat through their long necks and tails.

Previously, researchers have only been able to use indirect ways to gauge dinosaur metabolism or body temperatures.

For example, they inferred dinosaur behavior and physiology by figuring out how fast dinosaurs ran based on the spacing of dinosaur tracks, studying the ratio of predators to prey in the fossil record, or measuring the growth rates of bone.

But these lines of evidence were often in conflict.

"For any position you take, you can easily find counter-examples," Eiler says. "How an organism budgets the energy supply it gets from food, and creates and stores the energy in its muscles--there are no fossil remains for that."

Eagle, Eiler and colleagues developed what's known as a clumped-isotope technique that shows that it's possible to determine accurate body temperatures of dinosaurs.

"We're getting at body temperature through a line of reasoning that I think is relatively bullet-proof, provided you can find well-preserved samples," Eiler says.

In this method, the researchers measured the concentrations of the rare isotopes carbon-13 and oxygen-18 in bioapatite, a mineral found in teeth and bone.

How often these isotopes bond with each other--or "clump"--depends on temperature.

The lower the temperature, the more carbon-13 and oxygen-18 bond in bioapatite. Measuring the clumping of these isotopes is a direct way to determine the temperature of the environment in which the mineral formed--in this case, inside the dinosaur.

"What we're doing is special in that it's thermodynamically-based," Eiler says. "Thermodynamics, like the laws of gravity, is independent of setting, time and context."

Because thermodynamics worked the same way 150 million years ago as it does today, measuring isotope clumping is a reliable technique, says Eiler.

Identifying the most well-preserved samples of dinosaur teeth was one of the major challenges of the analysis.

The scientists used several ways of finding the best samples. For example, they compared the isotopic compositions of resistant parts of teeth--the enamel--with easily altered materials like the fossil bones of related animals.

The next step, the researchers say, is to determine the temperatures of more dinosaur samples, and extend the study to other species of extinct vertebrates.

In particular, discovering the temperatures of unusually small and young dinosaurs would help test whether dinosaurs were indeed gigantotherms.

Knowing the body temperatures of more dinosaurs and other extinct animals would also allow scientists to learn more about how the physiology of modern mammals and birds evolved.

In addition to Eagle, Eiler and Tripati, co-authors of the paper are Thomas Tutken from the University of Bonn, Germany; Caltech undergraduate Taylor Martin; Henry Fricke from Colorado College; Melissa Connely from the Tate Geological Museum in Casper, Wyoming; and Richard Cifelli from the University of Oklahoma. Eagle also has a research affiliation with UCLA.

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