There was never anyone quite like him before and it is very unlikely that his kind will be seen again. Baron Franz von Nopcsa (Figure B14.8.1) was one ofthe first paleontologists who saw to it that dinosaurs were interpreted in their full biological context. For this, he is generally regarded as the founder of the field of paleobiology. From him, we've learned about the unusual dinosaur fauna from Transylvania, that part of western Romania where his noble family's estate was located. This skeletal material formed the mainstay ofNopcsa's research, including soft tissue reconstruction and its relevance to jaw mechanisms, paleoecological reconstructions of the region as a Late Cretaceous island, and the evolution of the dwarf dinosaurs that lived on the island. He also published extensively on the early evolution of birds from small predatory dinosaurs, the origin of new evolutionary conditions due to disease, the pituitary gland and large size in dinosaurs, and the relationship between bone histology, growth rates, and thermoregulation among dinosaurs. A polyglot, Nopcsa published not only in German, but also regularly in English, Hungarian, and French.
Remarkable though these achievements were, they were conducted against a background of scientific and political involvement in the founding of the state ofAlbania. Nopcsa was captivated by the geography and people of this stark, yet beautiful land of the western Balkans. He began working there in 1906 and by the end of his career had published some still-current monographs on the geography, geology, and ethnography of Albania and its people.
By 1912, Austria-Hungary was exceedingly worried about safe travel to the Mediterranean and saw Nopcsa's knowledge of the geography of the area to be tactically important to them. So Nopcsa became a spy during the first and second Balkan wars, the first of which was to establish the new country of Albania from the dying Ottoman Empire in Europe, and the second was to prevent neighboring states from absorbing its territory. At that time, the new country needed a king, so Nopcsa volunteered for the job, suggesting to the Austro-Hungarian army chief of staff that he would fund the war effort with money he would obtain from marrying the daughter
of some American millionaire and would pledge Albania as an ally to the Empire in exchange for recognition as King. As far as is known, his proposal received no response, although he continued his spy work in Romania during World War I.
Despite his international activities, Nopcsa was a private man. He lived most of his life in Vienna, except for two years as Director of the Hungarian Geological Survey in Budapest. Living with him was his secretary, friend, and lover, Bajazid Elmas Doda, an Albanian he met in 1906. Transylvania was ceded to Romania after World War I and the Nopcsa estate was lost. Thereafter, Nopcsa's mental health declined and early in the morning of 15 April, 1933, he dosed Bajazid's tea with sleeping powder and then shot him. Going into his work room, Nopcsa wrote a suicide note and then killed himself.
In all four aspects, the changes wrought by cladistic analyses in our understanding of archo-saurs in general, and dinosaurs in particular, were nothing short of revolutionary.
Thecodontia. "Thecodontia" (theco - socket; odon - tooth; teeth set in sockets) was a term invented in 1859 by Richard Owen to group primitive archosaurs (see Figure 13.4). Thinking cladistically, however, "Thecodontia" was diagnosed using the same diagnostic characters as Archosauria; therefore, it must include all archosaurs - not just primitive ones. So the group "Thecodontia" was superfluous when there was already a group called Archosauria. Cladistic analysis, therefore, propelled the abandonment of what had for 120 years been considered a very important group of animals.
Dinosaur origins. Recall that Seeley divided dinosaurs into two groups - ornithischians and saurischians (on the basis of their pelvic anatomy) - and that he viewed these as having separate origins within "Thecodontia." The elegant 1986 cladistic work of Jaques A. Gauthier (Figure 14.12), now at Yale University, however, provided ample corroboration of a mono-phyletic Dinosauria, identifying upward of 10 derived features uniting all dinosaurs with each other (see Chapter 4). Since then, numerous cladistic analyses of both new and old taxa have confirmed that dinosaurs share a single, most recent common ancestor, itself a dinosaur. Ornithischia and Saurischia - each mono-phyletic - are more closely related to each other than they are to anything else.
Ornithischian and saurischian relationships. There are some other differences between the precla-distic view of dinosaurs and a more modern one. For example, before cladograms, all large, carnivorous dinosaurs were grouped together within the evocatively named Carnosauria; Coelurosauria was kept for the smaller forms. Cladistic analysis, however, paints a very different picture: one in which several different lineages of large-sized theropods, with their short arms and huge heads, evolved independently. The large theropod Carnotaurus is as far from other large (tetanuran) theropods as it is possible for one theropod to be from another. And Coelurosauria, once the exclusive domain of small, light-bodied forms, is now home to the mighty Tyrannosaurus rex as well. The clado-grams tell us that when theropods grow very large, they independently tend to take a similar form: large head, small arms, typically with reduced numbers of fingers, and long tails. What was it about their behavior or basic design that produced that shape?
Cladograms also revealed the important links between ceratopsians and pachy cephalosaurs (once thought to be a kind of ornithopod), confirmed suspected links between
ankylosaurs and stegosaurs, and situated the enigmatic therizinosaurs (once thought to be troodontids, or ornithischians, or who knows what else!) within Theropoda.
Birds as dinosaurs. Ostrom, as we have seen, constructed a compelling anatomical case for birds as dinosaurs in his 1974 paper on Archaeopteryx. In 1986, Gauthier published his now-classic paper on saurischian monophyly, in which he addressed Ostrom's observations from a cladistic viewpoint. Gauthier's analysis forms the backbone of our treatment of bird origins in Chapter 10 and, as we've seen, overwhelmingly confirms the fundamental dinosaurian ancestry of birds. With so many important evolutionary insights afforded by the use of cladograms, there should be no wonder why we have emphasized a cladistic approach throughout this book.
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