How Basal is Endothermy to Tetrapods?

A phenology of the evolution of endothermy in birds and mammals

Author:

Lovegrove

Abstract:

Recent palaeontological data and novel physiological hypotheses now allow a timescaled reconstruction of the evolution of endothermy in birds and mammals. A three-phase iterative model describing how endothermy evolved from Permian ectothermic ancestors is presented. In Phase One I propose that the elevation of endothermy – increased metabolism and body temperature (Tb) – complemented large-body-size homeothermy during the Permian and Triassic in response to the fitness benefits of enhanced embryo development (parental care) and the activity demands of conquering dry land. I propose that Phase Two commenced in the Late Triassic and Jurassic and was marked by extreme body-size miniaturization, the evolution of enhanced body insulation (fur and feathers), increased brain size, thermoregulatory control, and increased ecomorphological diversity. I suggest that Phase Three occurred during the Cretaceous and Cenozoic and involved endothermic pulses associated with the evolution of muscle-powered flapping flight in birds, terrestrial cursoriality in mammals, and climate adaptation in response to Late Cenozoic cooling in both birds and mammals. Although the triphasic model argues for an iterative evolution of endothermy in pulses throughout the Mesozoic and Cenozoic, it is also argued that endothermy was potentially abandoned at any time that a bird or mammal did not rely upon its thermal benefits for parental care or breeding success. The abandonment would have taken the form of either hibernation or daily torpor as observed in extant endotherms. Thus torpor and hibernation are argued to be as ancient as the origins of endothermy itself, a plesiomorphic characteristic observed today in many small birds and mammals.

Evidence of a High Metabolic Rate in Archosaurs

Palaeohistological Evidence for Ancestral High Metabolic Rate in Archosaurs

Authors:

Legendre et al

Abstract:

Metabolic heat production in archosaurs has played an important role in their evolutionary radiation during the Mesozoic, and their ancestral metabolic condition has long been a matter of debate in systematics and palaeontology. The study of fossil bone histology provides crucial information on bone growth rate, which has been used to indirectly investigate the evolution of thermometabolism in archosaurs. However, no quantitative estimation of metabolic rate has ever been performed on fossils using bone histological features. Moreover, to date, no inference model has included phylogenetic information in the form of predictive variables. Here we performed statistical predictive modelling using the new method of phylogenetic eigenvector maps on a set of bone histological features for a sample of extant and extinct vertebrates, in order to estimate metabolic rates of fossil archosauromorphs. This modelling procedure serves as a case study for eigenvector-based predictive modelling in a phylogenetic context, as well as an investigation of the poorly known evolutionary patterns of metabolic rate in archosaurs. Our results show that Mesozoic theropod dinosaurs exhibit metabolic rates very close to those found in modern birds, that archosaurs share an higher ancestral metabolic rate than that of extant ectotherms, and that this derived high metabolic rate was acquired at a much more inclusive level of the phylogenetic tree, among non-archosaurian archosauromorphs. These results also highlight the difficulties of assigning a given heat production strategy (i.e. endothermy, ectothermy) to an estimated metabolic rate value, and confirm findings of previous studies that the definition of the endotherm/ectotherm dichotomy may be ambiguous.

Were Mosasaurs Warm Blooded?

Endothermic mosasaurs? Possible thermoregulation of Late Cretaceous mosasaurs (Reptilia, Squamata) indicated by stable oxygen isotopes in fossil bioapatite in comparison with coeval marine fish and pelagic seabirds

Authors:

Lynn Harrell et al

Abstract:

The thermoregulatory style of Late Cretaceous mosasaurs has become a highly controversial subject in vertebrate palaeontology. These extinct marine reptiles have previously been described as poikilothermic, endothermic or gigantothermic. Here we analyse three genera of mosasaurs from the Mooreville Chalk in Alabama (USA) of differing body mass, and compare their δ18OPO4 derived body temperatures (Tb) with those of coeval poikilothermic fish (Enchodus) and endothermic pelagic seabirds (Ichthyornis). Results show that all mosasaurs, Clidastes (Tb = 33.1°C), Platecarpus (Tb = 36.3°C), and Tylosaurus (Tb = 34.3°C), had elevated average body temperatures in relation to those of the fish (Tb = 28.3°C) and were closer to those of Ichthyornis (Tb = 38.6°C). The temperatures calculated for Enchodus compare well with previously reported temperature estimates for the Mooreville Chalk and the Tb of Ichthyornis compares well with temperatures that have been reported for modern seabirds, suggesting that this method provides accurate results. Finally, although there are small differences of body temperature among mosasaur genera, these are independent of size, and thus inferred body mass, suggesting that mosasaurs were not gigantotherms, but rather endotherms.

The sixth sense in mammalians forerunners as a clue to the evolution of endothermy

The sixth sense in mammalians forerunners: Variability of the parietal foramen and the evolution of the pineal eye in South African Permo-Triassic eutheriodont therapsids

Authors:

Benoit et al

Abstract:

In some extant ectotherms, the third eye (or pineal eye) is a photosensitive organ located in the parietal foramen on the midline of the skull roof. The pineal eye sends information regarding exposure to sunlight to the pineal complex, a region of the brain devoted to the regulation of body temperature, reproductive synchrony, and biological rhythms. The parietal foramen is absent in mammals but present in most of the closest extinct relatives of mammals, the Therapsida. A broad ranging survey of the occurrence and size of the parietal foramen in different South African therapsid taxa demonstrates that through time the parietal foramen tends, in a convergent manner, to become smaller and is absent more frequently in eutherocephalians (Akidnognathiidae, Whaitsiidae, and Baurioidea) and nonmammaliaform eucynodonts. Among the latter, the Probainognathia, the lineage leading to mammaliaforms, are the only one to achieve the complete loss of the parietal foramen. These results suggest a gradual and convergent loss of the photoreceptive function of the pineal organ and degeneration of the third eye. Given the role of the pineal organ to achieve fine-tuned thermoregulation in ectotherms (i.e., “cold-blooded” vertebrates), the gradual loss of the parietal foramen through time in the Karoo stratigraphic succession may be correlated with the transition from a mesothermic metabolism to a high metabolic rate (endothermy) in mammalian ancestry. The appearance in the eye of melanopsin-containing retinal ganglion cells replacing the photoreceptive role of the pineal eye could also have accompanied its loss.

How did Warmed Bloodedness Originate?

Reptile thermogenesis and the origins of endothermy

Author:

Tattersall

Abstract:

Extant endotherms have high rates of metabolism, elevated body temperatures, usually tight control over body temperature, and a reasonable scope for further increases in metabolism through locomotor activity. Vertebrate ectotherms, on the other hand, rely on behavioural thermoregulation and cardiovascular adjustments to facilitate warming, and generally lack specific biochemical and cellular mechanisms for sustained, elevated metabolism. Nevertheless, the ancestral condition to endothermy is thought to resemble that of many extant reptiles, which raises the question of the origins and selection pressures relevant to the transitional state. Numerous hypotheses have emerged to explain the multiple origins of endothermy in vertebrates, including thermoregulatory, locomotory, and reproductive activity as possible drivers for these sustained and elevated metabolic rates. In this article, I discuss recent evidence for facultative endothermy in an extant lepidosaur, the tegu lizard. Since lepidosaurs are a sister group to the archosaurs, understanding how a novel form of endothermy evolved will open up opportunities to test the compatibility or incompatibility of the various endothermy hypotheses, with potential to elucidate and resolve long contentious ideas in evolutionary physiology.

Late Permian Cynodont Cynosaurus Lacked a Pineal eye, may Have Been Endothermic

Physiological implications of the abnormal absence of the parietal foramen in a late Permian cynodont (Therapsida)

Authors:

Benoit et al

Abstract:

The third eye (pineal eye), an organ responsible for regulating exposure to sunlight in extant ectotherms, is located in an opening on the dorsal surface of the skull, the parietal foramen. The parietal foramen is absent in extant mammals but often observed in basal therapsids, the stem-group to true mammals. Here, we report the absence of the parietal foramen in a specimen of Cynosaurus suppostus, a Late Permian cynodont from South Africa (SA). Comparison with Procynosuchus delaharpeae, a contemporaneous non-mammalian cynodont from SA, demonstrates that the absence of this foramen is an abnormal condition for such a basal species. Because seasonality was marked during the Late Permian in SA, it is proposed that the third eye was functionally redundant in Cynosaurus, possibly due to the acquisition of better thermoregulation or the evolution of specialized cells in the lateral eyes to compensate for the role of the third eye.

Appears Dimetrodon Relative Ophiacodon was Warm Blooded

Most people know that ‘warm-bloodedness’ is a characteristic of mammals. This trait actually encompasses a suite of physiological processes that help to maintain a relatively high, constant body temperature using heat generated internally. A new study by Christen Don Shelton of the University of Cape Town, South Africa and his colleague, Martin Sander at the University of Bonn, Germany, presented at this year’s Society of Vertebrate Paleontology meeting, shows that this character may have shown up in the ancestors of modern mammals far earlier than was previously thought.

One associated effect of being ‘warm-blooded’ is a relatively fast growth-rate. Mammals (and birds, who are also ‘warm-blooded’) tend to grow much faster than ‘cold-blooded’ vertebrates, like fish and reptiles. This fast growth rate is in turn associated with a particular type of bone growth pattern, called fibrolamellar bone (FLB). Both mammals and birds have FLB, and Shelton and his colleague investigated its presence in an early fossil relative of mammals, Ophiacodon.

Ophiacodon is distantly related to the large sail-back reptile, Dimetrodon, that people may be familiar with, and lived in North America around 280-300 million years ago. Although it didn’t look particularly mammal-like (you could be forgiven for calling it a ‘lizard’), it has many characters that link it with mammals. The highly vascularized tissue had previously been observed in Ophiacodon, but its presence had been written off to ecological factors, like its aquatic lifestyle, rather than to its fundamental physiology. Shelton looked at a number of bones of Ophiacodon of individuals of different ages, and found that the more likely explanation is that Ophiacodon was at least partly ‘warm-blooded’

link.

Determining Dinosaur Endothermy Based on the Temperature the Egg Shells Formed

Were dinosaurs really fast, aggressive hunters like the ones depicted in the movie "Jurassic World"? Or did they have lower metabolic rates that made them move more like today's alligators and crocodiles? For 150 years, scientists have debated the nature of dinosaurs' body temperatures and how those temperatures influenced their activity levels.

New research by UCLA scientists indicates that some dinosaurs, at least, had the capacity to elevate their body temperature using heat sources in the environment, such as the sun. They also believe the animals were probably more active than modern-day alligators and crocodiles, which can be active and energetic, but only for brief spurts.

The researchers also found evidence that other dinosaurs they studied had lower body temperatures than modern birds, their only living relatives, and were probably less active.

The research is published today in the journal Nature Communications.

Led by Robert Eagle, a researcher in the department of earth, planetary and space sciences in the UCLA College, the scientists examined fossilized dinosaur eggshells from Argentina and Mongolia. Analyzing the shells' chemistry allowed them to determine the temperature at which the eggshells formed -- information that had not been previously known.

"This technique tells you about the internal body temperature of the female dinosaur when she was ovulating," said Aradhna Tripati, a co-author of the study and a UCLA assistant professor of geology, geochemistry and geobiology. "This presents the first the direct measurements of theropod body temperatures."

The Argentine eggshells, which are approximately 80 million years old, are from large, long-necked titanosaur sauropods, members of a family that include the largest animals to ever roam the Earth. The shells from Mongolia's Gobi desert, 71 million to 75 million years old, are from oviraptorid theropods, much smaller dinosaurs that were closely related to Tyrannosaurus rex and birds.

Sauropods' body temperatures were warm -- approximately 100 degrees Fahrenheit, according to the study. The smaller dinosaurs had substantially lower temperatures, probably below 90 degrees.

link.

paper link.

Late Triassic Rhynchosaur Hyperodapedon Bone Growth Suggests Archosauromorphs Basally Grew Fast

New insights from bone microanatomy of the Late Triassic Hyperodapedon (Archosauromorpha, Rhynchosauria): implications for archosauromorph growth strategy

Author:

Mukherjee

Abstract:

Bone microanatomy of multiple postcranial skeletal elements of several individuals of Hyperodapedon collected from India is reported. This reveals that fibrolamellar bone tissue is predominant in the mid- and inner cortices, whereas the peripheral region of the cortex is composed of either parallel-fibred and/or lamellar bone. The pattern of primary osteons mostly ranges between laminar and subplexiform. Such predominance of fibrolamellar bone tissue in the cortex suggests an overall fast growth, which slowed down considerably later in ontogeny. Four distinct ontogenetic stages are identified based on the bone microstructure. During the juvenile stage, growth was fast and continuous, but it became punctuated during the early and late sub-adult stages. In adult individuals, growth was slow and showed periodic interruption but did not stop completely, suggesting that Hyperodapedon had an indeterminate growth strategy. Interelemental histovariations affecting cortical thickness, organization of the vascular network, incidence of growth rings and extent of secondary reconstruction are noted. Throughout ontogeny, the femora show higher cortical thickness than humeri and tibiae, suggesting differential appositional growth rate between the skeletal elements. Differences in cortical thickness are noted in the ribs, which suggest differential functional constraints based on anatomical site-specific occurrences. Although fibrolamellar bone tissue became progressively more dominant towards the archosaurs, there are considerable variations in the growth patterns of the archosauromorphs. This is exemplified by the bone microstructure of Hyperodapedon, which deviates from the generalized slow-growth pattern proposed for all basal archosauromorphs, suggesting that rapid growth was already present in the archosauromorphs. The cortical thickness of various long bones of Hyperodapedon bears similarity with that of several extant terrestrial quadrupeds, suggesting that Hyperodapedon was essentially a terrestrial quadruped.

Olenekian/Anisian Triassic Ichthyosaur Utatsusaurus was Probably Warm Blooded

Osteohistology of the Early Triassic ichthyopterygian reptile Utatsusaurus hataii: Implications for early ichthyosaur biology

Authors:

Nakajima et al

Abstract:

Ichthyosaurs were highly adapted to a marine lifestyle, as shown by their fish-like body shape and their assumed active swimming abilities and high metabolic rates. However, the processes of adaptation to an aquatic life in the early stages of this lineage remain poorly understood. Here, we present the first osteohistological data concerning the most basal ichthyopterygian yet known, Utatsusaurus hataii, from the Lower Triassic of Japan. The cancellous bone structure suggests adaptation to active swimming in an open marine environment. Moreover, the possible occurrence of rapidly deposited bone tissue, in a fibrolamellar complex, suggests a higher metabolic rate than in modern poikilothermic reptiles, and therefore a trend toward homeothermy. This basal ichthyosaur, with its elongate body, was already more adapted to an aquatic lifestyle than expected from its morphology, and the process of adaptation to a marine lifestyle was already well advanced by the Early Triassic.

Were Dinosaurs Mesotherms?

Evidence for mesothermy in dinosaurs

Authors:

Grady et al

Abstract:

Were dinosaurs ectotherms or fast-metabolizing endotherms whose activities were unconstrained by temperature? To date, some of the strongest evidence for endothermy comes from the rapid growth rates derived from the analysis of fossil bones. However, these studies are constrained by a lack of comparative data and an appropriate energetic framework. Here we compile data on ontogenetic growth for extant and fossil vertebrates, including all major dinosaur clades. Using a metabolic scaling approach, we find that growth and metabolic rates follow theoretical predictions across clades, although some groups deviate. Moreover, when the effects of size and temperature are considered, dinosaur metabolic rates were intermediate to those of endotherms and ectotherms and closest to those of extant mesotherms. Our results suggest that the modern dichotomy of endothermic versus ectothermic is overly simplistic.

Evolution of Feather Color in Dinosaurs & Birds

Melanosome evolution indicates a key physiological shift within feathered dinosaurs

Authors:

Li et al

Abstract:

Inference of colour patterning in extinct dinosaurs has been based on the relationship between the morphology of melanin-containing organelles (melanosomes) and colour in extant bird feathers. When this relationship evolved relative to the origin of feathers and other novel integumentary structures, such as hair and filamentous body covering in extinct archosaurs, has not been evaluated. Here we sample melanosomes from the integument of 181 extant amniote taxa and 13 lizard, turtle, dinosaur and pterosaur fossils from the Upper-Jurassic and Lower-Cretaceous of China. We find that in the lineage leading to birds, the observed increase in the diversity of melanosome morphologies appears abruptly, near the origin of pinnate feathers in maniraptoran dinosaurs. Similarly, mammals show an increased diversity of melanosome form compared to all ectothermic amniotes. In these two clades, mammals and maniraptoran dinosaurs including birds, melanosome form and colour are linked and colour reconstruction may be possible. By contrast, melanosomes in lizard, turtle and crocodilian skin, as well as the archosaurian filamentous body coverings (dinosaur ‘protofeathers’ and pterosaur ‘pycnofibres’), show a limited diversity of form that is uncorrelated with colour in extant taxa. These patterns may be explained by convergent changes in the key melanocortin system of mammals and birds, which is known to affect pleiotropically both melanin-based colouration and energetic processes such as metabolic rate in vertebrates, and may therefore support a significant physiological shift in maniraptoran dinosaurs.

Euparkeria Grew Like a Warm Blooded Animal

Evidence for high bone growth rate in Euparkeria obtained using a new paleohistological inference model for the humerus

Authors:

Lucas J. Legendre, Loic Segalen & Jorge Cubo

Abstract:

The study of bone growth rate and metabolic rate evolution in archosaurs (crocodiles, dinosaurs including birds, and pterosaurs) and close outgroups has become a subject of major interest among paleontologists in recent years. In this paper, we estimate the bone growth rate of Euparkeria using a new statistical inference model for the humerus. We modified the taxonomic range of extant species used in previous studies, on which we performed quantitative measurements of histological features and bone growth rates. Bone growth rate values estimated for Euparkeria are crucial in understanding the ancestral condition for archosaurs because this taxon is considered the closest relative to the archosaur crown group. We obtained an instantaneous growth rate of 6.12 μm/day, suggesting that Euparkeria shared with other non-archosaurian archosauromorphs (Prolacerta, Proterosuchus, and Erythrosuchus) a condition of high growth rate compatible with endothermy. This derived state may have been inherited by some Triassic crurotarsans, as suggested by the high instantaneous bone growth rate (14.52 μm/day) estimated in this study for Postosuchus. Jurassic crurotarsans may have lost endothermy during the transition from terrestrial habitats and active predation to aquatic habitats and sit-and-wait predation behaviors, so that Cretaceous crocodiles may be secondarily ectothermic, as suggested by δ18O values. In conclusion, we provide new evidence for the hypothesis of an ancestral endothermic state for the last common ancestor of archosaurs, and show that non-archosaurian archosauromorphs and Triassic crurotarsans may have been characterized by a thermometabolism more similar to that of dinosaurs than to that of lepidosaurs and turtles.

More Evidence of Dinosaur Endothermy

University of Adelaide research has shown new evidence that dinosaurs were warm-blooded like birds and mammals, not cold-blooded like reptiles as commonly believed.

In a paper published in PLoS ONE, Professor Roger Seymour of the University's School of Earth and Environmental Sciences, argues that cold-blooded dinosaurs would not have had the required muscular power to prey on other animals and dominate over mammals as they did throughout the Mesozoic period.

"Much can be learned about dinosaurs from fossils but the question of whether dinosaurs were warm-blooded or cold-blooded is still hotly debated among scientists," says Professor Seymour.

"Some point out that a large saltwater crocodile can achieve a body temperature above 30°C by basking in the sun, and it can maintain the high temperature overnight simply by being large and slow to change temperature.

"They say that large, cold-blooded dinosaurs could have done the same and enjoyed a warm body temperature without the need to generate the heat in their own cells through burning food energy like warm-blooded animals."

In his paper, Professor Seymour asks how much muscular power could be produced by a crocodile-like dinosaur compared to a mammal-like dinosaur of the same size.

Saltwater crocodiles reach over a tonne in weight and, being about 50% muscle, have a reputation for being extremely powerful animals.

But drawing from blood and muscle lactate measurements collected by his collaborators at Monash University, University of California and Wildlife Management International in the Northern Territory, Professor Seymour shows that a 200 kg crocodile can produce only about 14% of the muscular power of a mammal at peak exercise, and this fraction seems to decrease at larger body sizes.

"The results further show that cold-blooded crocodiles lack not only the absolute power for exercise, but also the endurance, that are evident in warm-blooded mammals," says Professor Seymour.

"So, despite the impression that saltwater crocodiles are extremely powerful animals, a crocodile-like dinosaur could not compete well against a mammal-like dinosaur of the same size.

"Dinosaurs dominated over mammals in terrestrial ecosystems throughout the Mesozoic. To do that they must have had more muscular power and greater endurance than a crocodile-like physiology would have allowed."

A Hypothesis on the Aquisition of Endothermy in Therapsids

The Role of Foraging Mode in the Origin of Therapsids: Implications for the Origin of Mammalian Endothermy

Author:

1. James A. Hopson (a,*)

Affiliations:

a. Department of Organismal Biology, University of Chicago, 1027 East 57th Street, Chicago, IL 60637, USA

*. Contact at jhopson@uchicago.edu.


Abstract:

The question of the adaptive basis for the origin of mammalian endothermy remains unresolved despite a great deal of research effort. Controversy continues over which physiological adaptations were of greatest importance in starting ectothermic nonmammalian synapsids of the Late Paleozoic on the path that culminated in modern endothermic mammals. Models of the selective basis for the origin of endothermy fall into two main categories: “thermoregulation first” and “aerobic capacity first.” Studies of lizards show a dichotomy between a low-energy “sit-and-wait” (SW) foraging mode in Iguania and a more energy-intensive “widely foraging” (WF) mode in Autarchoglossa. It is proposed that in the transition from basal synapsids (“pelycosaurs”) to therapsids, a shift from the primitive SW mode to the WF mode put the ancestors of mammals on the path to increased aerobic capacity and the ability to sustain high levels of foraging activity. Selection for increased energy expenditure disproportionately increased the amount of food energy consumed, thus improving foraging efficiency. A shift from reliance on anaerobic muscle metabolism for short but rapid dashes to capture prey to a reliance on aerobic metabolism for active searching for prey necessitated improvements of the cardiovascular system and lungs for increased aerobic capacity and greater stamina. Over time, therapsids became locked into high food requirements, which selected for improvements in aerobic metabolism, locomotor and food-processing ability, and neurosensory/behavioral specializations. Evidence of a link between maximum activity metabolism and resting (basal) metabolism in anurans and rodents suggests that further increases in aerobic activity metabolism required an increased basal metabolic rate, which led to high body temperatures and, ultimately, homeothermy. Therapsids show adaptations for increased activity, greater food-getting and food-processing ability, and higher metabolic rates than basal synapsids (“pelycosaurs”). It is argued that the “foraging mode” model is preferable to the “parental care” model of Farmer and the “correlated progression” model of Kemp for understanding the origin of mammalian endothermy.

Dino Sinuses Hint at Endothermy?

The Paranasal Air Sinuses of Predatory and Armored Dinosaurs (Archosauria: Theropoda and Ankylosauria) and Their Contribution to Cephalic Structure

Lawrence M. Witmer *, Ryan C. Ridgely

Department of Biomedical Sciences, College of Osteopathic Medicine, Ohio University, Athens, Ohio
email: Lawrence M. Witmer (witmerL@ohio.edu)

*Correspondence to Lawrence M. Witmer, Department of Biomedical Sciences, College of Osteopathic Medicine, Ohio University, Athens, OH 45701
Fax: 740-593-2400

Funded by:
National Science Foundation; Grant Number: NSF BSR-9112070, NSF IBN-9601174, NSF IBN-0343744, NSF IOB-0517257
Ohio University College of Osteopathic Medicine

Abstract
The paranasal air sinuses and nasal cavities were studied along with other cephalic spaces (brain cavity, paratympanic sinuses) in certain dinosaurs via CT scanning and 3D visualization to document the anatomy and examine the contribution of the sinuses to the morphological organization of the head as a whole. Two representatives each of two dinosaur clades are compared: the theropod saurischians Majungasaurus and Tyrannosaurus and the ankylosaurian ornithischians Panoplosaurus and Euoplocephalus. Their extant archosaurian outgroups, birds and crocodilians (exemplified by ostrich and alligator), display a diversity of paranasal sinuses, yet they share only a single homologous antorbital sinus, which in birds has an important subsidiary diverticulum, the suborbital sinus. Both of the theropods had a large antorbital sinus that pneumatized many of the facial and palatal bones as well as a birdlike suborbital sinus. Given that the suborbital sinus interleaves with jaw muscles, the paranasal sinuses of at least some theropods (including birds) were actively ventilated rather than being dead-air spaces. Although many ankylosaurians have been thought to have had extensive paranasal sinuses, most of the snout is instead (and surprisingly) often occupied by a highly convoluted airway. Digital segmentation, coupled with 3D visualization and analysis, allows the positions of the sinuses to be viewed in place within both the skull and the head and then measured volumetrically. These quantitative data allow the first reliable estimates of dinosaur head mass and an assessment of the potential savings in mass afforded by the sinuses.

So far the above only occurs in endotherms...in part also for heat exchange. hmmmm...

Couldn't resist the pix. There are more with the paper!

Nat Geo has a page too. So does Paleoblog.