Analysis of the nasal region in fossils of three theropod dinosaurs ( Nanotyrannus, Ornithomimus, and Dromaeosaurus ) and one ornithischian dinosaur ( Hypacrosaurus ) showed that their metabolic rates were significantly lower than metabolic rates in modern birds and mammals. In extant endotherms and ectotherms, the cross-sectional area of the nasal passage scales approximately with increasing body mass M at M 0.72 . However, the cross-sectional area of nasal passages in endotherms is approximately four times that of ectotherms. The dinosaurs studied here have narrow nasal passages that are consistent with low lung ventilation rates and the absence of respiratory turbinates.

Nasal respiratory turbinates are complex, epithelially lined structures in nearly all birds and mammals that act as intermittent countercurrent heat exchangers during routine lung ventilation. This study examined avian respiratory turbinate function in five large bird species (115-1, 900 g) inhabiting mesic temperate climates. Evaporative water loss and oxygen consumption rates of birds breathing normally (nasopharyngeal breathing) and with nasal turbinates experimentally bypassed (oropharyngeal breathing) were measured. Water and heat loss rates were calculated from lung tidal volumes and nasal and oropharyngeal exhaled air temperatures (T ex) ' Resulting data indicate that respiratory turbinates are equally adaptive across a range of avian orders, regardless of environment, by conserving significant fractions of the daily water and heat budget. Nasal Tex of birds was compared to that of lizards, which lack respiratory turbinates. The comparatively high nasal Tex of the lizards in similar ambient conditions suggests that their relatively low metabolic rates and correspondingly reduced lung ventilation rates may have constrained selection on similar respiratory adaptations.

Reptiles and birds possess septate lungs rather than the alveolar-style lungs of mammals. The morphology of the unmodified, bellowslike septate lung restricts the maximum rates of respiratory gas exchange. Among taxa possessing septate lungs, only the modified avian flow-through lung is capable of the oxygen–carbon dioxide exchange rates that are typical of active endotherms. Paleontological and neontological evidence indicates that theropod dinosaurs possessed unmodified, bellowslike septate lungs that were ventilated with a crocodilelike hepatic-piston diaphragm. The earliest birds ( Archaeopteryx and enantiornithines) also possessed unmodified septate lungs but lacked a hepatic-piston diaphragm mechanism. These data are consistent with an ectothermic status for theropod dinosaurs and early birds.

In terms of their diversity and longevity, dinosaurs and birds were/are surely among the most successful of terrestrial vertebrates. Unfortunately, interpreting many aspects of the biology of dinosaurs and the earliest of the birds presents formidable challenges because they are known only from fossils. Nevertheless, a variety of attributes of these taxa can be inferred by identification of shared anatomical structures whose presence is causally linked to specialized functions in living reptiles, birds, and mammals. Studies such as these demonstrate that although dinosaurs and early birds were likely to have been homeothermic, the absence of nasal respiratory turbinates in these animals indicates that they were likely to have maintained reptile-like (ectothermic) metabolic rates during periods of rest or routine activity. Nevertheless, given the metabolic capacities of some extant reptiles during periods of elevated activity, early birds were probably capable of powered flight. Similarly, had, for example, theropod dinosaurs possessed aerobic metabolic capacities and habits equivalent to those of some large, modern tropical latitude lizards (e.g., Varanus), they may well have maintained significant home ranges and actively pursued and killed large prey. Additionally, this scenario of active, although ectothermic, theropod dinosaurs seems reinforced by the likely utilization of crocodilian-like, diaphragm breathing in this group. Finally, persistent in vivo burial of their nests and ap-*

Ultraviolet light analysis of a fossil of the theropod dinosaur Scipionyx samniticus revealed that the liver subdivided the visceral cavity into distinct anterior pleuropericardial and posterior abdominal regions. In addition, Scipionyx apparently had diaphragmatic musculature and a dorsally attached posterior colon. These features provide evidence that diaphragm-assisted lung ventilation was present in theropods and that these dinosaurs may have used a pattern of exercise physiology unlike that in any group of living tetrapods.

Longisquama insignis was an unusual archosaur from the Late Triassic of central Asia. Along its dorsal axis Longisquama bore a series of paired integumentary appendages that resembled avian feathers in many details, especially in the anatomy of the basal region. The latter is sufficiently similar to the calamus of modern feathers that each probably represents the culmination of virtually identical morphogenetic processes. The exact relationship of Longisquama to birds is uncertain. Nevertheless, we interpret Longisquama's elongate integumentary appendages as nonavian feathers and suggest that they are probably homologous with avian feathers. If so, they antedate the feathers of Archaeopteryx, the first known bird from the Late Jurassic.

Skeletal ontogeny in extant archosaurians (crocodilians and birds) indicates that the morphology of the perinatal pelvic girdle is an indicator of overall developmental maturity [that is, altriciality (nestbound) versus precociality (mobile and relatively independent)]. Comparison of the skeletal anatomy of perinatal extant archosaurians and perinatal dinosaurs suggests that known dinosaur hatchlings were precocial. These data are consistent with the overall similarity in nesting behavior of dinosaurs and modern crocodilians.

Dinosaurs were among the most distinctive and successful of all land vertebrates. Attempts at reconstructing their biology have become commonplace. However, given the absence of closely comparable living models, deciphering their physiology necessarily remains speculative and determination of their metabolic status has been particularly problematical. Nevertheless, many paleontologists have advocated the notion that they were probably ''warm-blooded'' (endothermic), thus providing a model supposedly essential to the interpretation of these animals as having led particularly active, interesting lives. Those suppositions notwithstanding, the apparent absence of respiratory turbinates in dinosaurs, as well as likely ectothermic patterns of thermoregulation in very early birds, argues strongly that these animals were unlikely to have achieved the metabolic status of modern terrestrial endotherms. These data are not necessarily inconsistent with current models of active lifestyles of dinosaurs.