The Work of FRANZ Baron NOPCSA (1877-1933): Dinosaurs, Evolution and Theoretical Tectonics

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L Jb. Geol. B.-A. ISSN 0016-7800 Band 127 Heft 2 S.187-203 Wien, August 1984

By DAVID B. WEISHAMPEL & WOLF-ERNST REIF*) With 2 figures and 1 table

Dinosaurier Kiefermechanik Systematik Histologie Paläobiologie Archaeopteryx Evolution Orthogenese Neolamarckismus Biogeographie Vulkanismus Orogenese Kontinentalverschiebung

Table of Contents

Summary, Zusammenfassung 187 1. Introduction ...... 188 2. Dinosaur Systematics and Paleobiology : : : : : : . '. : : '. : : : : : : : : : : : : : : : : : 189 3. Biogeography of the Cretaceous Siebenbürgen Island 190 4. Dinosaur Jaw Mechanics 190 5. Sexual Dimorphism , 191 6. Paleohistology , . . .. 191 7. Origin of Flight , , : : : : : : : : : : : : : : : : : : : : 192 8. Kerunia 192 9. Evolution 193 10. Theoretical Tectonics 195 11. Conclusions , 198 Literature , 199

Summary lieh untersucht. NOPCSAist im Wesentlichen bis heute als Di- The scientific studies of FRANZBaron NOPCSAhave received nosaurierspezialist bekannt. Es zeigt sich jedoch, daß NOPCSA only minor modern analysis, despite their relevance to early sich mit seinen Publikationen und Vorträgen entschieden ge- works in dinosaur paleobiology, neo-Lamarckian evolution, gen einseitiges Spezialistentum wandte. NOPCSAwar viel mehr theoretical tectonics and paleobiogeography. NOPCSA'Scon- ein sehr vielseitiger Theoretiker, der seine eigentliche Leistung temporary impact in these areas varies from progressive and in der Synthese getrennter Arbeitsgebiete, im Übertragen von insightful to eccentric and wrong. We analyse NOPCSA'Sstu- Konzepten auf neue Anwendungsgebiete und im Auffinden und dies in terms of their contell1Porary impact on the natural Lösen von Anomalien und Paradoxien sah. Wir untersuchten /' sciences and their relevance to modern research. Several in- NOPCSASIdeen zu einzelnen Problemen, ihre Rezeption durch teresting facets of his papers emerge: The mere existence of Zeitgenossen und ihr Nachwirken bis in die heutige Zeit. NOPCSA'Svolcanological hypotheses, including rifting and sub- NOPCSASkühne Konzeptionen haben in den meisten Fällen zu duction, his extensions and corrections to neo-Lamarckian fruchtbaren Diskussionen angeregt, auch wenn sich einige sei- evolution, functional morphology of the origin of avian flight, ner Lösungen sehr bald als unhaltbar erwiesen haben. and his advocacy of broad eclectic research programmes. Als Reptilpaläontologe beschrieb und revidierte NOPCSA nicht nur zahlreiche Dinosaurier-Taxa, er führte auch zahlrei- Zusammenfassung che neue Gesichtspunkte ein (Sexualdimorphismus, Biogeo- graphie, Schädelkinetik) und führte andere erfolgreich fort Aus dem umfangreichen Werk von FRANZ Baron NOPCSA (Biomechanik, Knochenhistologie). NOPCSAsetzte sich vehe- werden die Arbeiten und Bücher über fossile Reptilien, Inver- ment für das Modell der Vogel-Entstehung aus biped laufen- tebraten (Kerunia), Evolutionstheorie und theoretische Tektonik den (nicht arborikolen) Reptilien ein. analysiert. NOPCSASLeben und Werk wurde in kurzen Nachru- Bei der Interpretation von Kerunia (einer Hydrozoe aus dem fen (LAMBRECHT,1933; EDINGER, 1934, 1955; F. E. SUESS, Eozän) entwickelte NOPCSAein elegantes (wenn auch fal- 1933; WOODWARD,1934), in einer Biographie (KUBACSKA, sches) Modell einer Symbiose zwischen einem schalen losen 1945) und in Handbuchartikeln (LAMBRECHTet aI., 1938; ZAP. Cephalopoden und einer Hydrozoe. FE, 1978) beschrieben, aber noch nie wissenschaftsgeschicht- In einer Zeit, wo der Neolamarckismus von den meisten Bio- logen und Paläontologen unkritisch akzeptiert wurde, erkannte *) Authors' addresses: Prof. Dr. DAVIDB. WEISHAMPEL,Depart- NOPCSAdie Schwierigkeiten dieser Evolutionstheorie. Er ent- ment of Biological Sciences, Florida International University, wickelte eine Korrektur des Neolamarckismus, ohne in die Tamiami Campus, Miami, Florida 33199, USA; Dr. habil. dogmatische Einseitigkeit der Orthogenese-Theorie zu verfal- WOLF-ERNSTREIF, Institut für Geologie und Paläontologie, Uni- len, mit der einzelne andere Autoren den Neolamarckismus zu versität Tübingen, D-7400 Tübingen, BRD. überwinden versuchten. Er fand den Mechanismus, der die LA.

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MARCKschenThesen (Gebrauch und Nichtgebrauch der Orga- ne; Vererbung erworbener Eigenschaften) nicht in Frage steil- te, aber gleichzeitig das Aussterben von Taxa verständlich machte, in der "Arrostie". Arrostie ist die lang anhaltende Stö- rung des Hormonsystems, die zunächst zu (adaptiven) Verän- derungen in Körpergröße und Körperbau, aber letziich zum Aussterben führt. Nach NOPCSASTod überwogen zunächst die Einflüsse der Orthogenese-Theorien bis sich die Synthetische Theorie durchsetzte. NOPCSASKorrektur des Neolamarckismus blieb daher weitgehend unbekannt. Trotz seiner umfangreichen Erfahrungen in der Geologie Al- baniens blieb NOPCSAein Außenseiter in der Diskussion theo- i .. ~' . retischer tektonischer Probleme. Er demonstrierte die Frucht- ."-" 'to barkeit des Deckenmodelles und wandte die WEGENERSehe Ia Kontinentalverschiebungstheorie auf Probleme der Biogeogra- '~'7, -r phie an. In einer weitgehend unbekannt gebliebenen Arbeit I (1927a) erklärte er Orogenesen als gigantische Überschiebun- gen von Sialplatten und zeigte, daß Magmatismus vom atlanti- schen, vom pazifischen und vom gemischten Typ an verschiedene Regionen und Entwicklungsumstände eines Orogens und an Spalten, die beim Auseinanderdriften von Kontinentalblök- ken entstehen, gebunden ist. NOPCSA hat damit wichtige Aspekte der Plattentektonik vorweg genommen. NOPCSASletzte Arbeit ist ein Schema vom Ablauf der Erdge- schichte, das auf der Vorstellung orogenetischer Zyklen und auf der Kontinentalverschiebungstheorie basierte. ~- ,.,

1. Introduction "Personalia: Died: Dr. FRANZBaron NOPCSAin Vien- na on the 25th of April at the age of 56. For a long time he directed the Hungarian Geological Survey and made merit for himself as an unusually manysided and spiri- ted scholar, especially in the investigation of Albanian geology and the paleontology of reptiles." This anony- mous obituary, published in 1933 in the Centralblatt für Mineralogie, Geologie und Paläontologie, summarizes all that is commonly known about FRANZBaron NOPCSA VON FElSÖ-SZllVAs, a familiar figure in vertebrate paleontology (Fig. 1). However, even allowing for a few obituaries and memorials, NOPCSAis very poorly known today. Indeed his work has never been studied from a Fig. 1: FRANZ Baron NOPCSA(Drawing by F. MARTON,1926 [from KUBACSKA,1945]). theoretical point of view and it is our purpose to outline the highlights of his work as a progenitor to today's concept of paleobiology and continental drift. NOPCSAcontinued to acknowledge DOllO'S contribution Born 3 May, 1877, near Hatszeg, Hungary (now part to paleontology in numerous subsequent papers. of western Roumania), NOPCSAattended and received NOPCSApublished a total of 158 papers (101 of them his Abitur from the Theresianum in Vienna and went on in paleontology) before his death in 1933. These works to the University of Vienna in 1898. NOPCSA'Sfirst con- span such topics as paleobiology and taxonomy of lo- tact with vertebrate paleontology was rather coinciden- wer tetrapods (particularly dinosaurs, but also including tal; in 1895 fossil reptile bones were discovered by his turtles, squamates and pterosaurs), bird flight, evolutio- sister on the family estate near Szentpeterfalva and nary theory, paleophysiology, paleohistology, archaeo- these NOPCSAshowed to Prof. Dr. E. SUESSat the Uni- logy, ethnology, geology and Albanian geography. versity of Vienna. SUESSapparently planned formal ex- NOPCSA'Sprimary research areas were dinosaur paleo- cavations but they were never undertaken. On his se- biology and systematics (42 papers), although he also cond visit with SUESS, NOPCSAwas told to conduct a discussed other archosaurs (5 papers on crocodilians, study of the Szentpeterfalva material by himself. This 4 on pterosaurs), lepidosaurs (11 papers), therapsids (4 work culminated in NOPCSA'Sfirst paleontological con- papers), turtles (7 papers), ichthyosaurs (1 paper) and tribution, on a new hadrosaurid dinosaur from the Sie- amphibians (1 paper). benbürgen region of Hungary, at the Vienna Academy NOPCSAseems to have had a good understandig of of Science on 21 June, 1899. Also in 1899, NOPCSA his international audience. He published primarily in made his first journey to Albania, a country which fasci- German (114 papers) including the bulk of his tectonic nated him for the rest of his life. work and in part reflecting his close personal ties with While still at the Theresianum, NOPCSAmet OTHENIO Vienna and its academy early in his career. However, ABEL. ABEL, then a graduate assistant to E. SUESS,be- important papers which strike an international pose came well known along with NOPCSAas one of Eur- (e. g. 1905e, 1907, 1923e, 1926b, 1928c, 1929a, 1934) ope's earliest paleobiologists (EHRENBERG,1975; REIF, were commonly written in English for both English- 1980). NOPCSAgraduated from the University of Vienna language and German-language journals (31 papers). in 1904 and in the same year met LouIS DOllO in Brus- He published 9 papers (primarily on geology but also sels. Influenced by DOllO even before this meeting, on paleontology) in Hungarian, his native tongue, and

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additionally 5 papers in French on fossil material found Table 1: VON HUENE'S and NOPCSA'S Dinosaur Taxonomies. in Normandy and Central France. VON HUENE (1914) NOPCsA (1923) In 1920, NOPCSAwas offered membership in the Geo- logical Survey of Roumania which he declined. How- Ramus Sauromorpha [Subclass Archosauria] ever, he became Director of the Royal Hungarian Geo- (= diapsids) logical Survey in 1925. This he resigned in 1929. He Supraorder Dinosauria continued his paleontological and geological work until Order Saurischia Order Saurischia 1933, when he committed suicide on 25 April. Suborder Coelosauria Suborder Pachypodosauria During his lifetime, NOPCSAkept a diary which was (= prosauropods) read by K. LAMBRECHTand A. T. KUBACSKA,two of his Suborder PachYPodosauria biographers. Presumably from his diary and other me- (= carnosaurs, prosauro- morabilia, NOPCSAwrote an autobiography which was pods, sauropods) never published (KUBACSKA,1945, p. 275). The only Suborder Coelosauria available material on NOPCSAare the few obituaries and Suborder Megalosauria memorials (LAMBRECHT,1933; LAMBRECHTet aI., 1938; Suborder Sauropoda EDINGER,1934, 1955; F. E. SUESS, 1933; WOODWARD, Order Ornithischia Order Orthropoda 1934, ZAPFE, 1978) and KUBACSKA'sbiography (1945). Suborder Ornithopoda Suborder Poposauria It is unknown to us whether the NOPCSAestate still (= Poposaurus) exists in the National Museum of Hungary in Budapest. Suborder Thyreophora Suborder Ornithopoda Additionally, the VONHUENEarchives in the Institut und (= stegosaurs, ankylosaurs) Museum für Geologie und Paläontologie, Universität Suborder Thyreophora Tübingen, contains 45 postcards and letters from FRANZ (=ceratopsians, stegosaurs, BARONNOPCSAto FRIEDRICHVON HUENE. ankylosaurs) Suborder Ponderopoda (= ceratopsians) 2. Dinosaur Systematics and Paleobiology schians and ornithischians (Orthopoda of NOPCSA),sug- Probably NOPCSA's broadest and most accessible gested that at the supraordinal level these two groups works for the present generation of reptilian paleontolo- were more closely related to each other than to any ar- gists are his studies of the taxonomy and phylogenetic chosaur group. NOPCSA'sand VONHUENE'staxonomies relationships among fossil reptiles. His best known stu- of dinosaur relationships are given in Table 1. Since the dies, Die Familien der Reptilien (1923d), The Genera of 19205, much work has been done on "within-group" ta- Reptiles (1928c) and Osteologia reptilium recentium et xonomy and phylogenetics of dinosaurs (much by VON fossilium (1931) helped establish taxonomic diversity HUENE,but also by COLBERT,GILMORE,LULL, PARKSand among fossil reptiles during the early part of this cen- STERNBERG,to name a few). Yet the debate over phylo- tury. genetic relations of and between dinosaurs and their NOPCSA'stomes on reptile taxonomy were based in subdivisions, begun ultimately between OWENand SEE- large part on more than 50 papers on the anatomy and LEY, goes on unabated today (cf. BAKKER& GALTON, taxonomy of widely diverse groups of fossil reptiles. 1974; BAKKER,1980; BONAPARTE,1976; CHARIG,1976, These range from original descriptions of new taxa 1982). The issues of ancestor-descendant versus si- (1900, 1903a, 1923f, 1929c, 1930), revised accounts of ster-group relationships and vertical versus horizontal old taxa (1902, 1903a, 1905a,b,c, 1911, 1912, 1923g), classification schemes are still cloudy and it will take in- and studies of phylogenetic relationships among repti- cisive and broad research on all archosaur groups be- les (1903b, 1910, 1925b, 1932c). The constant stream fora a clear solution to dinosaur taxonomy and phyloge- of NOPCSA'sresearch on fossil reptiles parallels that of netic is in sight. the other prominent reptilian paleontologist in Europe At a time when descriptive studies of dinosaur re- and NOPCSA'sgood friend and long-standig colleague, mains had become commonplace, NOPCSAwrote a se- FRIEDRICHFREIHERRVONHUENEof Tübingen, Germany. ries of innovative papers which combined osteological NOPCSAwas a frequent and favorite guest of the VON description with discussion of 50ft anatomy in fossil ar- HUENEfamily (U. VON HUENE,pers. comm., 1983) and chosaurs (e. g. 1900, 1902, 1903a). Most of his work for 25 years they exchanged correspondence on re- pertained to cranial musculature and neuroanatomy in spective research projects (KUBACSKA,1945), reflecting dinosaurs which, as is well known, lack modern relati- a continued exchange of ideas in their studies of fossil ves from which to extrapolate 50ft-tissue anatomy, phy- reptiles. siology, or behavior. Influenced most strongly by DOL- Since both NOPCSA'sand VON HUENE's main area of LO's (1883, 1884) studies of the cranial and skeletal re- research was on dinosaurs, it is interesting to note that constructions of Ig u a nod 0 n , NOPCSAdiscussed the their work diverged dramatically on the question of di- anatomy implicit not only in his dinosaur material from nosaur relationships and the concept of the Dinosauria Siebenbürgen (1900, 1902, 1903a, 1929b), but also in- as a natural group. VONHUENE,beginning in 1914, pro- cluding ankylosaur material from North America vided new and important support for SEELEY'S(1887) (1928a). Although he never published many details of separation of OWEN'S(1842) Dinosauria into the Sauri- the reptilian postcrania, these ideas were certainly not schia (including carnivorous and the gigantic sauropod foreign to him (letter of 3 October 1929 to VON HUENE dinosaurs) and the Ornithischia (remaining herbivorous discusses sauropod muscle reconstructions as a means groups). In doing 50, VONHUENEalways considered the to analyze body posture, limb movement and gait; Ku- Saurischia and Ornithischia to be distinct groups wi- BACSKA,1945). NOPCSA'sstudies, along with those of thout close common ancestry. NOPCSA(1917b, 1922, his contemporaries LULL (1908), OSBORN (1912), 1923d). accepting the dichotomy implicit between sauri- BROWN (1914), RUSSELL(1935) and LULL & WRIGHT

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(1942), represent an emerging interest in dinosaur bio- STANNIUS,1856). NOPCSAdid not elaborate the details logy that transcends the then common osteological des- of jaw mechanics in hadrosaurids (1900), nor in other criptions of taxa. These were the works which tackled ornithopod dinosaurs (1902, 1903a), but rather discus- the problem of understanding animals long since extinct sed his ideas as asides in descriptive sections of his and so different from their extant relatives as to exclude early and otherwise standard monographs. Nor did he most inferences about how these animals functioned follow up on his jaw mechanics work after 1903 or pro- and how they fitted into their ecological setting. It is on vide a clear understanding of the implications of his these initial studies that much of today's dinosaur pa- model of intracranial mobility in this group of dinosaurs. leobiology is built. NOPCSA'Ssuggestion that at least some kinds of dino- 3. Biogeography saurs had a movable quadrate might have been buried in the annals of descriptive paleontology had it not be- of the Cretaceous Siebenbürgen Island en for the detailed work on reptilian cranial morphology As alluded to in the introduction, NOPCSA'Soriginal by J. VERSLUYS.In his monograph (1910, 1912) which work on the Siebenbürgen fauna of Roumania (Ma- came out of a 2-year visit to the U. S. A., VERSLUYS estrichtian; JELETZKY,1963) was the mainstay of his di- analyzed the broad question on intracranial mobility in- nosaur work, producing important studies of the anato- cluding streptostyly in reptiles and birds. In these pa- my and taxonomy of 8 or so species of dinosaurs and pers VERSLUYSrejected NOPCSA'Smodel of quadrate other reptiles. Because of its location and the peculiar movement in hadrosaurs. However, in 1923, in a detai- aspects of its fauna, the Siebenbürgen material also led study of the famous hadrosaurid "mummy" in the lent itself to discussion of the relationship between is- Naturmuseum Senckenberg, Frankfurt am Main, VERS- land faunas, body size, species richness and evolutio- LUYS reversed his earlier opinion. He outlined a new nary atavism. Late in his career, NOPCSA(1923c, 1934) jaw mechanisms which incorporated NOPCSA'Soriginal began synthesizing taxonomic, ecologic and biogeogra- views of fore-aft quadrate mobility and included an ad- phic data to present a model of Siebenbürgen as an ditional medial rotation of the mandibles about their isolated Late Cretaceous island. NOPCSAthen argued long axes. In Europe, the interesting theme of intracra- that the fauna consisted of isolated descendants of nial mobility was carried on by VON KRIPP(1933c). Ho- much earlier and more cosmopolitan terrestrial reptiles. wever, he departed to a considerable degree from both Body size is lower among the Siebenbürgen animals re- NOPCSA'Sand VERSLUYS'models. VONKRIPP'Swork on lative to their ancestors als well as their closest con- jaw mechanisms in hadrosaurids stems from his then- temporary relatives on the mainland and, NOPCSApoin- current kinematic analyses of intracranial mobility of ted out, they are less diverse and retain the primitive bird skulls (1933a, b) and seems to represent a side-is- nature of their ancestors. NOPCSA'Swork and its impli- sue for him. Yet his work was to break largely with the cation to other Late Cretaceous terrestrial communities tradition of using joint morphology as the basis for erec- in Europe (viz. LAPPARENT,1947; PARIS&TAQUET,1973; ting a mechanism; rather VONKRIPPused kinematic dia- LAPPARENT& AGUIRE,1956; SEELEY,1881, 1883) have grams of 3-dimensional mechanisms modeled on joint been largely ignored since the early part of this century morpholgy and joint position to graphically simulate the (although see MOLNAR, 1980). These faunas and the working of the jaws. Through this new approach VON Siebenbürgen fauna are in great need of reexamination. KRIPPjustified a new jaw mechanism for hadrosaurids. NOPCSA'Sbroad paleoecologic and evolutionary outlook He rejected fore-aft quadrate mobility and medial rota- on his unusual Siebenbürgen fauna is both heterodox tion of the mandibles, instead outlining a jaw mecha- and interesting for its time, yet by today's view of island nism based on lateromedial movement of the quadrate biogeography often incomplete or wrong-headed. and lateral rotation of the mandibles. Surprisingly, VON KRIPP'Smechanism is based on the same Senckenberg 4. Dinosaur Jaw Mechanics hadrosaurid "mummy" which had been the foundation of VERSLUYS'Smechanism. VON KRIPP'Swork however, That analysis of vertebrate jaw mechanisms is impor- represents a more rigorous attack on the problem of tant to trophic paleoecology and evolutionary character modeling jaw mechanism in extinct animals by graphi- transformation is now ably expressed by the many pa- cally analyzing coordinated quadrare and mandibular pers on the subject (ALLIN, 1976; BARGHUSEN,1968; movement in conjuction with observations of tooth BRAMBLE,1978; DEMAR& BARGHUSEN,1972; GREAVES, wear. 1978). Although most deal with the evolution of the NOPCSA'Searly work on hadrosaurid jaw mechanics mammalian jaw, considerable work has been done on had the effect of establishing a polarity between jaw systems in reptiles, the modern approach having its European and North American paleobiologists working roots in the studies of STANNIUS(1856), VERSLUYS,LAK- on the same problem. The North American response JER and LUTHER.However, the early work of NOPCSA, (LAMBE,1920; LULL& WRIGHT,1942; OSTROM,1961) re- based on dinosaur material from Transsilvania, Hunga- jected intracranial mobility in favor of rigid skull mecha- ry, influenced to a great degree the ideas of these and nisms. LAMBErejected movement of the quadrate on the other authors, especially on intracranial mobilitv in re- basis of joint morphology and structural constraints cent and fossil reptiles. within the skull in his description of the hadrosaurid Ed- NOPCSA'S first paleontological paper (1900) consi- montosaurus regalis. Rather, he suggested that hadrosau- dered the possibility of intracranial mobility in the ha- rid jaw mechanisms consisted of simple jaw closing and drosaurid dinosaur Telmatosaurus (= Limnosaurus) transsylva- nearvertical shearing of the lower teeth past those in nicus, now stored in the British Museum (Natural Histo- the upper jaws. LAMBE'Smechanism was followed with- ry). Based in part on comments by MARSH(1893) and out comment in the first survey work on hadrosaurids by analogy with modern lacertilians, NOPCSAsuggested by LULL & WRIGHT(1942). that the quadrate bone was able to swing in a fore and OSTROM(1961) also rejected intracranial mobility in aft direction (streptostyly; a term coined originally by hadrosaurids based on structural constraints and in-

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stead indicated that jaw movement occurred slowly by LA, 1979; HORNER, 1982). NOPCSA'S sexual dimorphism means of fore-aft translation at the jaw joint. Like VON hypothesis however had several fatal flaws. Both KRIPP, OSTROM used several aspects of tooth wear to STERNBERG (1935) and RUSSELL (1946) pointed out that support his hypothesis. NOPCSA'S males and females rarely occur sympatrically NOPCSA'S model of hadrosaurid jaw mechanics as over wide stratigraphic intervals and hence are of little well as those of VERSLUYS, VON KRIPP and others, have taxonomic or paleobiologic value. That is not to say that recently been reanalyzed by WEISHAMPEL (1981, in the analysis of sexual dimorphism among fossil reptiles press a, in press b) using 3-dimensional computer ana- has had a waning history since NOPCSA'S time. Indeed, lyses. These previously mentioned mechanisms fail to the practice of identifying sexes and growth series model tooth wear in accordance with hadrosaurid real among these animals has been fruitfully applied to se- dentitions. Rather, hadrosaurid mechanisms consist of veral reptile groups in order to rectify taxonomic and mobile upper jaws, (maxillae and several other facial paleoecologic problems. Immediately following Nopc- bones) which can rotate laterally along a slightly in- SA'S work, several studies used presumed secondary clined hinge joint. Only in this way was it possible for sexual characteristics to qualitatively sex fossil reptiles hadrosaurids to chew side-to-side while the jaws re- and establish growth series: BROILI & SCHRÖDER (1937) mained isognathous (i. e. producing bilateral occlusion), on Dicynodon; BROWN & SCHLAIKJER (1940) on Protocera- something NOPCSA failed to observe. tops, and VAUGHN (1966) on Seymouria. These isolated studies, the now classical synthesis on sexual selection 5. Sexual Dimorphism and the fossil record by DAVITASHVILI (1961), and Hop- SON'S (1975) work on sexual selection and hadrosaurid For paleontologists who have worked on dinosaurs, dinosaurs round out the qualitative treatment of sexual NOPCSA is perhaps best known for his 1929 (a) work on dimorphism in fossil reptiles since NOPCSA'S time. Ac- supposed sexual dimorphism in these animals, un- cess to highspeed computers have made quantitative doubtedly because the paper is in English and because studies of growth and sexual dimorphism in fossil repti- the geological raw data are so at odds with the anato- les almost commonplace. The most important of these mic basis for his theory. It is rarely discussed, however, (in one sense, descendant pieces of NOPCSA with a nu- that NOPCSA'S interest in the impact of sexual dimor- merical bent) include DODSON (1975, 1976) on lambeo- phism on reptilian taxonomy extends back to 1905( e) saurine hadrosaurids and Protoceratops, respectively, and when he adduced evidence for the presence of sauro- GRINE & HAHN (1978), GRINE et al. (1978) and BRADU & pod 0 s pen is from an enigmatic bone which was pre- GRINE (1979) on the therapsid Diademodon. It is likely, viously interpreted as a clavicle (HATCHER, 1901). The that sexual dimorphism, once introduced by NOPCSA to identification of this bone in terms of its primary sexual crudely explain the shapes of bones or skeletal configu- characteristics was a novel anatomic approach but met rations in dinosaurs, will appear in more studies of evo- with considerable resistance. HOLLANDS (1906), TOR- lutionary ecology and taxonomy in fossil reptiles, NIER (1909), and ABEL (1910) all rejected NOPCSA'S among other organisms. Despite being absolutely contention, the last two authors calling the bone an epi- wrong in his work on sexual dimorphism in these ani- sternal or first rib, respectively. NOPCSA again returned mals, NOPCSA'S legacy is certain to be fulfilled. to the subject of the sauropod os penis in 1918, but merely reiterated his earlier arguments. No one has 6. Paleohistology since advocated or refuted NOPCSA'S hypothesis; in- , deed it is never discussed in present-day literature. Starting in 1924, NOPCSA became interested in bone Hence, it seems that NOPCSA'S idea that an os penis histology as a means to identify reptilian taxa on the was present in sauropods is a dead issue. Yet, what is basis of small scraps of bone. His published work, of overriding importance is that he tried to understand which began in 1925, was hardly more advanced than how sexes among fossil reptiles might be identified in a such special-interest studies as MANTELL (1850a, b), biologically reasonable manner. OWEN (1859) and BROILI (1922), in which whole bones NOPCSA came back to the sauropod os penis in later or bone fragments were sectioned to document either work on sexual dimorphism and dinosaur taxonomy. internal morphology or identify the animal. Similarly, the Rather than arguing from the basis of supposed primary comprehensive work of QUECKETT (1855) and SEITZ sexual characteristics, as in his 1905 paper, NOPCSA (1907) was intended for anatomic and taxonomic com- (1915, 1928a, 1929a) turned to identifying what he be- parison, but over a wide variety of animals. The primary lieved were secondary sexual characteristics. He ar- source for NOPCSA'S slides was his own reptile material gued that the fossil record of dinosaurs is dominated from the Siebenbürgen area, but in 1924 VON HUENE with examples of sympatric species pairs. From these sent additional material from the Tübingen collections pairs, he itemized the then obvious dimorphism. Skull to NOPCSA to be thin-sectioned (letter of 1 August, shape, vertebral form and relative size of limb bones 1924; KUBACSKA, 1945). NOPCSA also received bone were ascribed to secondary sexual characteristics. The specimens from the British Museum (Natural History), shape of the distal ischium was related to the attach- the Royal Ontario Museum and the American Museum ment of penile musculature. By combining species of of Natural History. By 1925, NOPCSA had amassed a different genera into male and female pairs, he thought collection of more than 500 histologic slides pertainig to to establish a revised dinosaur taxonomy. In his short 86 fossil reptile genera (letter to VON HUENE, 25 March discussion (1929a), NOPCSA identified several small ha- 1925; KUBACSKA, 1945). As mentioned above, NOPCSA'S drosaurid genera as subadults of other taxa (cf. NOPCSA first paleohistology paper (NOPCSA, 1925; not listed in & HEIDSlECK, 1933; DODSON, 1975) and argued for such RICQLtS, 1980) intended to provide histologic discrimi- sophisticated social behaviour as display, social hierar- nation of fossil reptiles from Nigeria, based on frag- chies and parental care among several groups of orni- ments of femora, humeri and unidentifiable bone. Ho- thopods (for a modern treatment, see HORNER & MAKE- wever, in later studies he recognized that histologic

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structure may change from site to site within the skele- there exists a fundamental dichotomy in flight mecha- ton due to a variety of factors, among them stress and nism that can arise from either a quadrupedal or a bipe- growth rates. NOPCSAultimately chose to section ribs of dal ancestor. Quadrupedal ancestors evolve a flight ap- fossil reptiles, arguing that their mechanical loading paratus consisting of both fore- and hindlimbs plus ex- would be similar among all groups regardless of size or tensive patagium. In this way, the patagium effectively growth rates (letter to VON HUENE, 6 June 1929; Ku- supports the body at its center of gravity and flight con- BACSKA,1945). In one of his last two histologic papers sists first in gliding and secondarilay in powered flight. done in collaboration with E. HEIDSlECK,NOPCSAused Bipedal locomotion allows tow-fold limb specialization: rib histology to solve the problem of differentiating juve- 1. the development of hindlimb construction for curso- niles from small adults, recognizing conspecific juveni- rial locomotion and les and adults which had hitherto been assigned to dif- 2. independent forelimb specialization, including the ferent species (NOPCSA& HEIDSlECK,1933). development of a small patagium and thus foresha- Apparently NOPCSAhoped to conduct a thorough hi- dowing the avian wing. stologic study of all fossil reptiles (see numerous comments in KUBACSKA,1945), but its future was cut short Since only the forelimb in birds, including Archaeopte- by NOPCSA'Sdeath in 1933. The whereabouts of his hi- ryx, is involved in the flight apparatus, NOPCSAargued stologic thin sections are unknown to us. Such work that birds must have evolved from a bipedal cursorial was not to resume again until the 1950s and 60s with ancestor. As a likely consequence, powered flight arose the survey work of ENLOW & BROWN (1956, 1957, first, while gliding developed as a secondary achieve- 1958). Current work, often as a tool to understanding ment. In support of his theory, NOPCSAreferred to Archa- paleophysiology from bone microstructure, can be eopteryx and modern birds, citing highly modified cursio- found in studies of RICQLES(e. g. 1980). rial features of the hindlimb (e. g. reduction of pedal di- gits to a tridactyl condition, digitigrade stance, elonga- ted tibia and fibula relative to the femur) in comparison 7. Origin of Flight to the less modified forelimb construction and the long The discovery of Archaeopteryx brought to the paleonto- neck. Before the forelimb evolved into a wing, the hind- logical forefront the concept of evolutionary intermedia- limb developed as a cursorial functional complex, and tes, in this case at the class level between Reptilia and hence, birds must have been bipedal cursors before Aves. This unique status has generated two partly-de- they could fly. pendent evolutionary questions. First, what are the phy- In answer to the commonly held view that the avian logenetic relationships of Archaeopteryx? Second, what furcula represent fused paired clavicles in reptiles, he can Archaeopteryx reveal about the evolutionary history of advanced the idea (1923e, 1929b) that this homology avian flight? OSTROM, (1976), TARSITANO& HECHT does not exist. Rather the furcula sprang de novo as an (1980), HECHT & TARSITANO(1983) and THULBORN& ossificatioan of a tendon extending from the scapuloco- HAMLEY(1982) review in some detail the first question, racoid to the sternum and acting as an important sup- for which it has been suggested that bird origins can be port between the sternum and shoulder girdle. That found among such different groups as pseudosuchian such a tendon would have been loaded in tension and thecodontians, crocodilians, ornithopod dinosaurs and the furcula in compression seems not to have occurred theropod dinosaurs. NOPCSA'S work on bird origins to him and currently the furcula is believed to represent (1907, 1923e, 1927c, 1929b) does not deal explicitly co-ossified clavicles of dermalorigin (VANTYNE & BER- with phylogenetics; he denoted avian ancestry wihtin a GER, 1959). rather vague dinosaur-like group of animals. It is not NOPCSAtouched brieflyon the selective advantage of clear whether he considererd this group as belonging feathers in 1929. He agreed with. earlier authors that among pseudosuchians (thecodontians) or among dino- feathers had not initially evolved as a funtional re- saurs. His 1929 comments about a relationship with sponse to flight. Rather, he explored the possibility that "Proterosaurus" [= ?Proterosuchus] are vague and, we consi- their initial selective advantage was insulatory. Because der, misleading. What is important in NOPCSA'Sdiscus- he believed that the dinosaur (or dinosaur-like) bird di- sion of avian origins is his consideration of the evolu- chotomy may have taken place in the Permian, glacial tion of bird flight. conditions then-existing would have placed a premium In 1907, the prevailing theory of the origin of bird on insulatory structures - such as feathers - in con- flight maintained that it arose among tree-dwelling repti- junction with small size and elevated metabolic rates. les (MARSH, 1880; HURST, 1895; PYCRAFT,1894a, b, NOPCSA'Srun n i n g pro - a vis hypothesis was critici- 1896). The process of branch-hopping led to gliding zed in Europe and North America (HAY, 1910; ABEL, structures and finally to wings involved in powered 1912; BEEBE,1915; STEINER,1917) but later was igno- flight. In all cases, thecodontians were envisioned as red after the publication of HEILMANN'Streatise on the the group that spawned arboreally-derived flight. origin of birds (1927). This work championed both the In direct opposition to the arboreal theory of avian arboreal theory and thecodontian ancestry. flight and evolution from thecodontians, NOPCSAelabo- It is not well understood how much influence Nopc- rated the concept of a cursorial origin, whose progeni- SA'S works have had on discussions of the origin of tors as mentioned above were small, lightweight, dino- avian flight, so long has HEILMANN'Sstudy been in the saur-like animals. Although not the first to argue curso- vanguard. Later writings only elaborate the ancestry of rial origin or dinosaur/dinosaur-like ancestry for birds Archaeopteryx (DE BEER, 1954, 1964) or adaptive aspects (viz. WILLISTON,1879) it is clear that NOPCSAwas the of the arboreal theory (BOCK, 1965). However, if he is most ardent and articulate supporter of, as he termed it, correct, OSTROM'S(1973, 1974, 1976) theory of a cur- the "running pro-avis" concept. In contrast to WILLI- sorial, predaceous origin for bird flight, vindicates STON,NOPCSAsubstantiated his theory through several NOPCSA'Sopinion of the relationship between bipedality new lines of evidence. He argued that in vertebrates and flight. They differ with regard to the adaptive basis

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for the development of feathered wings. For NOPCSA, nally settled by EBERHARD FRAAS (1911) who published the flapping of proto-wings increased running speed; for a specimen from a Recent Hydractinia calcarea which had OSTROM, they were used as insect nets. However, phy- overgrown a serpulid worm and which had the typical logenetic analyses of Archaeopteryx by TARSITANO & Kerunia-shape. HECHT (1980; see also HECHT & TARSITASNO, 1983) cast doubt on OSTROM'S conclusions about the relationship 9. Evolution of Archaeopteryx and they further suggest that the origin of avian flight requires an intermediate arboreal stage. It is unknown whether NOPCSA had a more than aver- A final twist to the avian flight story comes from a age familiarity with evolutionary theory, i. e. with the very recent biomechanical study of the physics of classical publications on Darwinism and on Lamarckism ground-originating flight (CAPLE, et aI., 1983). They ad- and with current discussions. However, there is hardly vocate, a la NOPCSA, a cursorial proavis but on sophisti- any doubt that NOPCSA knew the papers by JAEKEL cated mechanistic grounds. This work suggests, if no- (1894, 1902, 1913), HENNIG (1928, 1932) and a few thing else, that it is insufficient to use simple scenarios other authors. In the 1920s when NOPCSA published his and functional morphology to explain the origin of flight important papers, neo-Lamarckism had lost its monoli- from, either the ground-up or tree-down. thic importance and was gradually being replaced by the theory of orthogenesis. Standpoints which combi- 8. Kerunia ned both views (i. e. neo-Lamarckism and orthogenesis) were regarded as quite acceptable by some au- One of the most charming papers by NOPCSA is that thors. In neo-Lamarckism, evolutionary change is on Kerunia (NOPCSA, 1905d). Kerunia cornuta (Eocene, brought about by the need of the organism; in other Fayum District, Egypt) is now known as a symbiosis be- words evolution is completely opportunistic and adapt- tween a pagurid hermit crab, which inhabits a snail- ive. In orthogenesis, evolutionary change is brought shell, and a hydractinian coelenterate. The hydractinian about by an internal program in the organism; in other settles on the snail-shell and with its growing skeleton words it is not opportunistic, but preprogrammed and increases the "living chamber" of the pagurid (DACQUE, not necessarily adaptive (REIF, in press). Compromises 1921, p. 471). Kerunia was discovered by the Swiss pa- between both points of view seem to be feasible. leontologist K. MAVER-EvMAR (1900, 1901). MAVER-Ev- The reason for the various attempts between 1910 MAR regarded it as a cephalopod. OPPENHEIM (1902) and 1930 to liberate paleontology from a dogmatic neo- discovered its true hydractinian nature on the basis of Lamarckism is apparent. Evolution could not be inter- surface structures but was not interested in the fact that preted as completely opportunistic, otherwise one (as he assumed) Kerunia originally overgrew a mollusc would not observe extinction, long-term linear trends, . shell which was later dissolved. Rather he was inter- and harmful characters (the famous sabre-tooth tiger ested in taxonomic problems (should the species re- comes to mind). ceive the generic name Kerunia or Hydractinia?) and in its Two solutions to this paradox are important in Ger- regional and stratigraphic distribution. man literature: ABEL'S theory of 1928 and 1929 which NOPCSA found virtues in both these views of MAVER- laid the ground for orthogenesis and NOPCSA'S theory EVMAR and of OPPENHEIM. He had collected Kerunia in which corrected neo-Lamarckism without accepting or- Egypt (during a trip with his mother and his sister in thogenesis and which is hence a unique theory. 1904). He showed the specimens to R. BULLEN NEW- To solve the paradox (of opportunism and adaptation TON, a specialist on Eocene cephalopods at the British on the one hand and extinction, long-term trends and Museum, who pointed out the great resemblance be- seemingly harmful characters on the other), ABEL sug- tween Kerunia and the Eocene cephalopod Belosepia. With gested an all-encompassing theme, the principle of a cross-section of a complete specimen (Figs. 1 and 2 inertia, which underlies all evolution. This principle in NOPCSA, 1905d), NOPCSA demonstrated the similarity combines in the growth pattern between Belosepia and Kerunia. Be- cause of the high regularity of the growth-pattern of Ker- 1. EIMER'S law of orthogenesis, unia, NOPCSA clearly excluded the model of a hydracti- 2. COPE'S law of the unspecialized ancestor, nian overgrowing a gastropod. Thin sections and sur- 3. DOLLO'S law of irreversibility and face structures clearly pointed out to NOPCSA that Ker- 4. ROSA'S law of the increasing reduction of variability unia is a hydractinian. NOPCSA found only one way out (REIF, in press). of this paradox and (despite the fact that he is comple- From the mere enumeration of the laws which were tely wrong) it is a fascinating model which he devel- combined, it is quite obvious that ABEL'S principle could oped: Kerunia is both a cephalopod and a hydractinian! It explain everything, adaptation as well as extinction, is a s y m bio s i s of a Belosepia-like cephalopod and a opportunism in evolution as well as longterm trends. hydractinian in which the hydractinian takes over com- The principle simply states that the evolutionary pro- pletely the task of skeleton-building for the cephalopod. cess has such a high inertia (in the strictest physical After the death of the cephalopod the hydractinian con- sense) that it can easily run beyond the adaptive realm tinues mineralizing and forms the typical irregular into the area of hypertrophied organs and extinction. shape in which no specimen is exactly alike another. NOPCSA'S solution was quite different. In one of his NOPCSA clearly pointed out that this model could be first papers (1923a, see also 1926a) of the series, tested with the recent hydractinian-pagurid hermit crab NOPCSA delt with DOLLO'S law of irreversibility (the term symbiosis. had been coined by ABEL). He showed under which Several authors after NOPCSA confirmed that Kerunia conditions evolution is irreversible (e. g. when a charac- was a hydractinian colony and H. DOUVILLE (1907) even ter is so strongly fixed that it cannot be altered) and ad- found the snail shell around which the hydractinian had ditionally gave several cases of reversible evolution in grown as predicted by OPPENHEIM. The problem was fi- fossil reptiles: if an organism faces an evolutionary

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dead-lock, it can only bß saved from extinction by a re- 2. Mechanically-induced convergence (e. g. torpedo versal to an embryonic stage. The life history of an indi- shape in aquatic vertebrates) is caused by natural vidual, seen as a distorted recapitulation of the history selection on newly acquired characters which are of its whole phylum, explains why a limited reversal of transmitted to the offspring. evolution can occur. 3. Arrostic changes: pachyostosis is caused by a defi- As early as 1917, NOPCSAapproached his major ciency of air-supply in aquatic animals. The changes theme, the relation between body size and the size of during the evolution of dinosaurs (size, tooth shape, the pituitary fossa in dinosaurs, several years before pattern of bone calcification) can be explained by a the growth hormone was identified in the pituitary disturbance of the hormone system. The distur- gland. With incrasing size of the pituitary fossa during bance, and this is NOPCSA'Snew idea, is caused by evolution, the bones of the extremities become more a change in food and in vitamin supply, leading to masive. In an extensive review of literature on endocri- hereditary changes in physiology. nologic diseases, NOPCSAcame to the conclusion that 4. Cases of inheritance or traumatic irritations. Tumors, there is a relationship between these phenomena and skin diseases (like keratosis) and traumatic lesions acromegaly. This is a hyperfunction of the pituitary can, if massively accumulated, be regarded as prim- gland, which is connected with an increase in size of ordial stages of horns, antlers and snout thickenings the pituitary fossa and which leads to gigantism and to in mystriosuchid phytosaurs. By a change in function extreme ossification in humans. Acromegaly was known the primordial stages can be developed as fully to be sometimens hereditary and that it leads to a de- functional structures. As growth and healing pheno- crease in fertility and vitality. NOPCSAwas very satisfied mena are accompanied by an activity of hormones, by this because this suggested an answer to why dino- there is again no doubt that the above-mentioned saurs died out. He did not elaborate on the question of disturbances become heritable by simply postulating how dinosaurs might have acquired acromegaly nor that hormones "effect [affect] the constitution of the how it became hereditary (NOPCSAapparently fully be- germplasm" (p. 646). lieved in LAMARCK'Sprinciple of the inheritance of ac- 5. Inheritance of acquired characters can never be quired characters). tested in experiments with living organisms. All ex- In 1923 (NOPCSA,1923b), he discussed osteological periments even over many generations seem like phenomena in marine tetrapods which he explained by "child's play" (p. 648) from a paleontological point of reference to the human diseases pachY9stosiS and view. "Even so much as one hundred generations osteosclerosis. "Pachyostosis" and "osteosclerosis" (in means nothing in the eyes of a paleontologist" (p. NOPCSA'Sinterpretation) occur in marine tetrapods 648). There is no doubt that NOPCSAis right here! which have only recently invaded aquatic habitats. It is There is no inductive experimental way through how- a physiological disturbance which is caused by the new ever many generations to disprove the inheritance of milieu and which is often adaptive by influencing the acquired characters. It is only by our understanding buoyancy of the animals. In fully adapted marine tetra- of the functional interrelationships between DNA, the pods, "pachyostosis" and "osteosclerosis" are secon- different RNAs and various enzymes that we can de- darily lost. The process of acquiring "pachyostosis" and ductively falsify the model of inheritance of acquired "osteosclerosis" in aquatic tetrapods is usually harm- characters. less, because the results are spread over many genera- NOPCSAsupported his case by citing numerous publi- tions. NOPCSAsuggested the term "Arrostie" for these cations by zoologists on successful experiments and on and the phenomena of gigantism in dinosaurs. The term the theoretical plausibility of the inheritance of acquired indicates that important evolutionary changes were characters. He assumed that the degree of fixation of a brought about by disease-like processes over many ge- character determines whether it can be altered by ex- nerations. ternal stimuli. These alterations are "phenotypical" be- A summary of his views on evolution is given in cause they occur via the phenotype. Changes are in- NOPCSA(1926b): "In a certain sense the factors of he- adaptive and lead to rapidly diverging lines (e. g. Lati- redity and evolution can only be studied on groups of pinnatidae and Longipinnatidae in ichthyosaurs). "After animals with a history going back to far remote geologi- this primary change, both groups evolve but slowly and cal times. They must also have recent representatives, exclusively on purely adaptive and mechanically-neces- so that their biology can be known at least to some ex- sitated lines" (p. 652). This is neo-Lamarckism at its tent. They must show a good amount of variation and best! How can one expect that random mutations can live in different media." There is no doubt that the lead to anything but inadaptive evolution? Evolution is higher vertebrates fulfill these requirements and that adaptive and hence we have to point out the appropria- this is the justification for NOPCSA'Spaper. It is very te mechanisms (among them, the evolutionary potential characteristic for the time when NOPCSAwrote these li- of what we would today call "heterochrony"). nes that paleontologists had no inkling of how important In the last chapter the "axiom of irreversibility" is re- laboratory genetics and population genetics were (or placed by the "much less rigid rule: the marks of would later become) for the theory of evolution. They evolution are generally ineradicable." Here did not regard it as arrogant, let along anactualistic, to NOPCSAcontinues to fight the dogmatism of ABEL'S claim that the fossil record provided the relevant causal "Law of DOllO" and again he is right; only with an un- factors for the process of evolution. derstanding (which we are approaching only today) of the function of the genome and epigenetic apparatus NOPCSAdelt with a variety of different problems: can one meaningfully discuss the question if irreversibi- 1. Parallel evolution, which is caused by similar envi- lity. ronmental stimuli on similarly structured tissues in A short version of the 1926(b) paper was published in closely related groups. These stimuli lead to newly German in Natur und Museum (1927b). Interestingly acquired heritable characters. enough this excerpt is accompanied by a neo-Lamar-

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ckian chapter from "Back to Methusala" by BERNHARD familiar not only with vertebrate anatomy, ecology, phy- SHAW(an arrangement that was made by the editor of logeny and biomechanics, but also with the large litera- the journal). ture of human pathology and endocrinology. ABEL, who developed his principle of inertia later, did not fully ac- We know from his biographers (EDINGER,1933; LAM- cept NOPCSA'Stheory. ABEL in his book of 1929 only BRECHT,1933; KUBACSKA,1945) that NOPCSAwas se- mentioned two less important descriptive papers by riously ill in 1928 when, on his invitation, the annual NOPCSA,but he did not quote from the more important meeting of the Paläontologische Gesellschaft took theoretical papers (NOPCSA,1917a, 1923a, b, 1926a, b. place in Budapest (1928b). NOPCSAread his welcoming 1927a, 1928b). Instead ABEL developed a view of evo- address from a wheelchair. It is important to know that lution which was much more schematic (not to say dog- NOPCSAbelieved that he would never be able to work matic) and which led directly to orthogenesis (REIF, in again (he was, however, wrong on this assumption) in press). It is fascinating to see how NOPCSAused ex- order to appreciate that this address has to be read as amples of teratology, pathology and endocrinology to his intellectual legacy. In the first paragraph NOPCSA discover how evolutionary novelties are brought about, emphasized both aspects of paleontology, "geo-paleon- always assuming that acquired characters are, under tology" and "paleo-zoology" (he did not use ABEL'S certain circumstances, heritable. If disturbances to nor- term paleobiology) only to demonstrate that both direc- mal physiology are slow, they can, at least normally, be tions are important. Natural Science is one unit and it is harmless and will be effective causes of evolutionary only our weakness that we separate individual discipli- change. In the long run, however, as in' the case of di- nes. "Der Nachteil der Zersplitterung der einheitlichen nosaurs, they can be one factor for the extinction of a Naturwissenschaft liegt darin, daß brach liegende taxon. It is quite obvious that NOPCSAprovided satisfac- Grenzbegiete entstehen." The whole paper that follows tory answers to all problems faced by dogmatic neo-La- is an entire argument against specialists who fight for marckism and that he avoided such dogmatism himself. the separation of disciplines and who never begin to The only problem is that he could not see in the matu- touch the "Grenzgebiete", in other words those who ring Mendelian genetics and in the tacitly developing avoid interdisciplinary work. population genetics the major lever arm which would NOPCSAthen gave a short survey of the history of pa- remove neo-Lamarckism from its eminent position with- leontology and its different approaches. He extensively in 10 years after his own death. pointed out the virtues of ABEL'S book on paleobiology (1911) and its neo-Lamarckian approach, only to con- clude that not all observable evolutionary changes are 10. Theoretical Tectonics explained by ABEL'S neo-Lamarckian model (see also NOPCSA'Scontributions to geology will be mentioned REIF, in press). Rather NOPCSAbelieved that ABEL neg- only partly here. It is well known that he wrote more lected that an organism and "hence every skeleton" is than 25 articles including a 700 page monograph on the not only a mechanically integrated but also physiologi- regional geology and stratigraphy of Albania. This will cally integrated unit (p. 8). He then summarized his re- not be discussed here. It suffices to mention that Nopc- sults on the inheritance of disturbances of the hormone SA, a former student of E. SUESS,was well trained in system and of the effects of oxygen supply. His main field geology and well informed about the tectonic and interest was no longer in the pituitary gland but in the geotectonic hypotheses of his time. To be expected, he thyroid gland, a discussion of which he was never able was progressive rather than conservative in application to complete due to his bad health. He concluded that of the nappe model to the reconstruction of the Dina- the mechanical adaptation of an organism during evolu- rids. In a letter to A. WEGENERof Aug. 23, 1928, he cal- tion is only possible to the degree that is allowed by its led himself a supporter of WEGENER'Stheory from the physiology; in other words, evolution is by no means as beginning (KUBACSKA,1945, p. 245). opportunistic as was claimed by orthodox neo-Lamar- The first paper which is of interest for us is the one ckians! Physiology plays such a dominating role that page note in which NOPCSApointed out that he had de- paralell evolution is quite common. A single species (he monstrated overthrusts in northern Albania in several gives as an example Mastodon) can originate from its an- papers since 1906 (NOPCSA,1921). NOPCSAcomplained cestral species in different areas (of Europe) at the sa- that L. KOBER,a well known alpine geologist, had clai- me time because the same environmental stimulus hits med in reviews of two NOPCSApapers and of a book by the same physiological system in many animals at the KOSSMAT,that he, KOBER, had discovered the nappe- same time. Arrostic processes can be strong, which structure of the Dinarids in 1914 and that NOPCSAhad makes taxonomy difficult (p. 15; it is not clear whether always negated ist. NOPCSA'Sdefense is clear but ag- NOPCSAthinks of saltational processes here). However, gressive and it shows that he feared for his acceptance if the arrostic processes are weak, mechanical adapta- by the establishment of structural geologists. It seems tion leads to "schöne und systematisch leicht über- as though he had every reason for this fear, because schaubare Entwicklungsreihen" (p. 15). NOPCSAcould the revolutionary ideas which he later published were not possibly have predicted the near end of the neo-La- systematically neglected. Most if not all of his papers marckian era when he exclaimed: "Meine Herren! Mit on paleontology and geology were "reviewed" in Ge 0- schwacher Hand habe ich heute versucht, einen schwe- logisches Zentral blatt. This, however, means not- ren Vorhang zu ziehen, um Ihnen ein neues Morgenrot hing because the "reviews" were simply short versions zu zeigen. Ziehen Sie, namentlich die Jüngeren von Ih- of the abstracts, or, at best, descriptions of the content nen, an diesem Vorhange nur kräftig weiter; Sie werden of the papers. We have been unable to find any of bemerken, das Morgenrot wird immer röter und Sie NOPCSA'Spapers on theoretical geology quoted in sum- werden einen Sonnenaufgang erleben." marizing books, textbooks, and those papers by well- This is NOPCSA'Sversion of neo-Lamarckism which known structural geologists which we have checked. fascinates by its interdisciplinary outlook. NOPCSAwas There is no doubt, however, that with respect to the re-

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I m

A B C

. . . Fig. 1. ~1V'"V'V'Vl 8auere E...ruptiva. ~r-:7:"l+++ hBilse. he E.rUIlt'lVI\. FI.gurenerkl~.rung: A Obere SIalregIOn, B Untere SiaJregi~n,.e Simaregion. I. Alte abgetragene Gebirge mit späterem Scbollenbau ,und ge- mIschten spateren EruptIonen. II. Junge FaltengebIrge mIt mnert!D Vulkankranz (links), überschobener Stirne (Mitte) und \'ortiefe (rechts). III. Alte Tafelländer mit Deckenergü8Ben. Fig. 2: ~OPC~A'Sm?del of magmati~ processes. A = ~pper sialic region; B = lower sialic region; C = simatic region; I = eroded m.ountal~regions Wlt~ postorogenetlc structural overpnnt and mixed postorogenetic volcanic eruptions; II = young fold-mountains with an Inner volcanic garland (left), an overthrusted front (center) and an outer trench (right); III = old continental plates with lava plateaus; vv = acidic eruptives; + + = alkaline eruptives (from NOPCSA,1927 a). gional geology of Albania, NOPCSAheld a virtual mono- magmatism. In many details his model agrees with the poly for several years which was accepted by most of modern view of magma origin. For all intents and pur- his colleagues. poses, NOPCSA'Sapproach and his diagram are highly In a letter of Jan. 2, 1925 to F. VON HUENE,NOPCSA original. He must have developed it himself, since it ap- exclaimed: "Hurrah! Hurrah! Hurrah! that you have Kar- pears that he cannot have "borrowed" it from other roo (South Africa) reptiles from Brazil. They must be sources. Despite his originality, NOPCSA' model does the same or nearly the same forms as in Africa. Unfor- not extend beyond the firm basis of WEGENER'SKonti- tunately I sent my manuscript on WEGENER'Scontinent- nentalverschiebungstheorie, namely the idea that sialic al drift to England away already". We were unable to plates "swim" on the simatic crust. This is certainly a discover which paper NOPCSAmeant and whether it was very important difference to the modern plate tectonics eventually published. concept. NOPCSA'Sfirst completely theoretical paper on tecto- Geotectonic models were widely discussed in the nics (1927a) takes continental drift for granted and do- 1920s. A critical summary is give by NÖLKE(1924) who es not even mention WEGENER.By quoting only a few concluded that among all the various models only the literature sources, NOPCSAdemonstrated that the main Theory of the Earth's Contraction (which had been structures of orogenies are gigantic overthrusts of sialic strongly supported by E. SUESS) could be seriously blocks. This model helped him, again by quoting only a held. The geophysicist KIRSCH(1928; 1938) summari- few references, to explain the mechanisms of three sed all data on radioactivity as an energy source for types of magmatism, each of which is restricted to cer- magmatism, convection in the Earth's mantle, and con- tain areas of a mountain region.The first type of mag- tinental drift. However, even in his second book KIRSCH matism which occurs on the inner, concave rim of did not quote NOPCSAnor did he develop any model of mountain regions is acidic and consists of trachyte, an- subduction. The British geologist ARTHUR HOLMESis desite, dacite and rhyolite. This type is found in the cir- usually praised for having first developed (in 1929) cum-Pacific and the Eurasian meridional mountain re- a model of convection in the Earth's mantle which gions: this is the Pacific-type of magmatism in SUESS' powered the drifting of crustal plates (HALLAM,1973, p. sense and NOPCSAexplained it by a melting of the sub- 26; SCHWARZBACH,1980, p. 98). However, it is only in- ducted sialic plate. The second type of magmatism is dicated by HALLAM(1973) but clearly shown by KIRSCH basaltic, occurring in the Pacific, in the Atlantic (Atlan- (1928, p. 76ff; KIRSCHseems to be completely forgotten tic-type of magmatism sensu SUESS)and on consolida- today) that the foundation to this theory was laid much ted sialic blocks (Syria, India, Australia, Antarctica, Ca- earlier. The roots can be found in HOLMES'own papers nada, Brazil). The magmas are derived from melting si- dating back at least to 1925 and JOLy's papers which matic crust. The third type is a mixed type consisting of date back much further (GREENE, 1982). The idea is peridotites, serpentinites, diabases and gabbros which summarised by F. E. SUESSin 1912. However this may occur in the center of eroded mountain regions. Ser- be, it seems to be true that HOLMES(1929) developed pentinites and peridotites characteristically can be an interesting and important model of convection. Oro- found in big overthrust zones. Peridotites, which chemi- geny in HOLMES'model is not a process of collision and cally are simatic (high Mg content), were squeezed into subduction, but simply a process of jolting of areas the overthrust zone from below during orogenesis. With where the convection current has a downward direction. details from his hypothesis, NOPCSAexplained why aci- Overthrusting and subduction of crustal plates are not dic magmatism is often ended by alkaline eruptions and included in HOLMES'model. alkaline magmatism often by acidic eruptions; a pheno- Only recently AMPFERER'SUnterströmungstheo- menon which he called "inversion". r ie has been "rediscovered" as one of the predeces- NOPCSAsupported his model of the different origins sors of WEGENER'S continental drift hypothesis (K. of magmas by a careful discussion of available geoche- SCHMIDT,1976; for dates on AMPFERER,see SCHWARZ- mical data. His diagram (1927a; see our fig. 2) is cer- BACH,1980). As early as 1906, AMPFERERargued stron- tainly the most spectacular part of his paper because it gly against the Theory of the Earth's Contraction and clearly shows subduction zones and their associated instead suggested that the mechanics of the Earths's

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crust can be explained by subcrustal magma flow. In gen strongly influenced German geology well into the 1911, AMPFERER& HAMMERsuggested a sucking-in of 1960s and 1970s (REIF, in press). crustal parts "erdeinwärts" (= into the mantle?) which The only tectonic synthesis which NOPCSA (1927) leads to a dense folding, overthrusting, and nappe for- quoted, but which we could not find referred to by other mation of crustal parts. authors, is that of SANDBERG(1924). SANDBERGseems AMPFERER(1925) very critically discussed WEGENER'S to have been an outsider in tectonics. He quoted only a theory. He did not accept that the sialic plates drift on few papers by himself which were published in German, the simatic layer, which would involve the acceptance French and English. He wrote his two-volume book in of shear zones on the ocean floor. Rather the plane of English which was translated into German by someone mass transport must lie much deeper than assumed by else. SANDBERGdeveloped a fixistic theory, rejecting all WEGENER.AMPFERERalso pointed out that the plates plate movement and also the Contraction Theory. All cannot be moved by outer forces but rather by inner tectonic processes ultimately are caused by vertical ones. If America drifts westward, as AMPFERERaccep- isostatic movements. The causes of the disturbance of ted, oceanic crust must sink into the depths, only to re- isostasy are differential heat flow, erosion, sedimenta- appear as oceanic crust in the Atlantic. This process of tion, melting, and compaction. NOPCSAdid not state mass transport is at the same time the motor for plate whether he agreed with SANDBERGor not; obviously he movement. (AMPFERERdid not point out that the ener- did not. NOPCSAonly quoted some volcanological and gy source is radioactivity!). This means that convection sedimentological data from SANDBERG. processes transport the American continent, at the One should comment on K. SCHMIDT'S(1976) "redis- same time s u c kin gin oceanic crust in the Pacific coveries" of AMPFERERand SCHWINNERthat these au- (which explains the deep sea trenches!) and reintegra- thors are not found in the English geological literature ting it by local eruptions in the Atlantic. AMPFERERdid nor in the English literature on the history of continental not provide any model of overthurst of sialic on simatic drift and plate tectonics. However, in German literature crust or of sialic on sialic crust. He strongly argued these authors and their theoretical contributions have against the assumption that the Andes can be explain- never been forgotten. A survey of the literature shows ed by a jolting of simatic crust or a sweeping together that both authors can be found in pre-war textbooks of the clastic detritus from the gradually eroded conti- (CLaas, 1936; BRINKMANN,1940; probably also in ear- nental block. He left it completely open as to how the lier editions of this book) and in text-books of West Andes formed and did not touch the problem of andesi- Germany (BRINKMANN,1976; ZElL, 1975; SCHMIDT-THO- tic volcanism. ME, 1968; 1972; WUNDERLICH,1966; 1968) and East In German-speaking countries a connection between Germany (KETTNER,1958; KRAUS, 1959; HOHL, 1971). radioactivity and geological processes was first summa- For all intents and purposes NOPCSA(1927a) was rized by F. E. SUESS(1912). In a paper which was also correct when he claimed that his model of orogeny and only recently "rediscovered" by K. SCHMIDT(1976) and magmatism is new. There are several reasons why this which was not quoted by NOPCSA,the geologist R. model was not accepted by other authors. The paper SCHWINNER(1920), Graz, Austria, developed a theory of appeared in a Hungarian version and in a German ver- thermal convection which strongly supports AMPFERER'S sion in the same journal (Földtani Közlöny) which was models and which explains magmatism and tectonic not widely read outside of Hungary. Only a short ver- processes. SCHWINNERaccepted radioactivity as the sion appeared in French in the memoirs of the Interna- energy source for all tectonic phenomena and introdu- tional Geological Congress in Madrid (1928d) (see Ku- ced the term "tectonosphere". The tectonosphere inclu- BACSKA,1945). NOPCSAwas never regarded as an au- des all outer layers of the Earth down to the level whe- thority on theoretical tectonics. Additionally it seems, at re there is a hydrostatic gravitational equilibrium. Ac- least for German literature, that in the 1930s conti- cording to SCHWINNERthe tectonosphere is 120 km nuous production of new geotectonic hypotheses which thick. SCHWINNERapplied models from meteorology (cy- was so characteristic for the 1920s had come to a halt. clones and anticyclones) to discuss the processes in KIRSCH,AMPFERER,SCHWINNER,STILLEand many others the tectonosphere and to explain orogenies and volca- continued to propose their models but no new author nism. He is most likely the first author to give schemati- appeared on the stage. cal diagrams of thermal convection of the earth and his WEGENER in the last (fourth) edition of his book model of orogeny is similar to that of HOLMES(1929). (1929) did not mention NOPCSA,for two possible rea- Volcanism is exclusively explained by a "Zerrungstekto- sons. First, he got the paper too late, since it probably nik" (extensional tectonics) of the crust. In other words, accompanied NOPCSA'Sletter of Aug. 23, 1928 (reprin- both specific features of NOPCSA'S(1927) model, over- ted in KUBACSKA,1945). Second, WEGENERwas not in- thrusting of sialic plates during orogenies and regional terested in the orogenetic and magmatic consequence differentiation of the different types of magmatism, are of his theory. He quoted SCHWINNER(1920; and also lacking. KIRSCH,1928) only with the idea that radioactivity could It is not surprising that the revolutionary publications be the source of energy for plate movement, but of WEGENER(1912, 1915) and of SCHWINNER(1920) thought that it was too premature to decide. were not mentioned by HANSSTILLEin his "G run d fra- We skip NOPCSA'S"Glossen zu E. HAARMANN'SOszil- gen der vergleichenden Tektonik" (1924). AM- lations-Theorie" (1932a). It is a very interesting polemi- PFERERis quoted only from his papers which describe cal paper but does not display too much of NOPCSA'S structural data, but not from his theoretical papers. own model. STILLEhimself favored the Contraction Theory and was From KUBACSKA(1945) we know how much work strongly fixistic; in other words, he rejected all plate NOPCSA spent on his "Zur Geschichte der Adria" movements, subductions, Unterströmungen, etc. STIL- (1932b) and how important this project was to him. In a LE'S personal authority and especially his G run d fra- very extensive review of paleontological, regional geo-

logical, biogeographical, gravimetric and seismic data ic, it is not surprising that the equatorial arc coincides NOPCSAshowed that the Adriatic See is not a geosyn- directly with the Jurassic equator and that the northern cline but a submerged craton (called "Adriatis"), name- end of the longitudinal belt is the Jurassic North Pole. If ly a promontory of the African craton. This model is we go back 220 million years, we find in the Variscian now generally accepted (CHANNEll et aI., 1979). CHAN- (late Paleozoic) mountains traces of two belts roughly NEll et al. (1979) show that the "Adriatis" (or "Adria") 40°-50° north and south which run parallel to the equa- problem dated back to SUESS(1883) and they mention tor. Very small traces are left of a longitudinal Variscian NOPCSA'Spaper but misrepresent his work by not poin- belt. Of the Caledonian orogeny another 220 million ting out that he was an adherent of continetal drift. years back, we find only parts of a mountain belt which originally probably formed a longitudinal belt. The distri- In his last paper which appeared posthumously in bution of glaciations throughout the Phanerozoic en- England (1934), NOPCSAdeveloped a unified picture of hances the regularity of this pattern. Four factors are global tectonics and paleogeography which he used as responsible for this development of the crust: the basis for discussion of the paleobiography of fossil reptiles and Stegocephalia. After he had submitted the 1. An eastward force which formed the longitudinal English paper, he wrote a German version of his global belts with their festoons. Its source is probably the tectonic concepts and sent it to Centralblatt six differential rotational velocity of the different layers weeks before his death. of the Earth. 2. The high rotational velocity of the Earth which by the NOPCSAgave simple but convincing data on paleobio- action of centrifugal force produced the two Varis- geography, paleoclimatology and the distributions of cian belts at middle latitudes. Later a lower rotatio- orogenies and Carboniferous glaciation to demonstrate nal velocity of the Earth together with cosmic factors that one has to accept continental drift. He avoided in- (gravity of Moon and Sun) formed the Alpine equa- troducing WEGENER'Stheory in a dogmatic way. It is an tiorial belt by compressing action of the centrifugal important addition to his magmatological model of 1927 forces. that he stated (1934, P. 108): "It seems likely that a 3. A migration of the poles which deforms the rotation- drifting asunder of land-masses would always be ac- elipsoid and which has strongest effects on those companied by eruptions of an Atlantic (basaltic) type. meridians on which they migrate. Such a combination ... is going on at the present in Gre- 4. Unknown, but intensive forces (among others also at African Rift Valley, on the borders of the Red Sea, "Polflucht?"), which lead to the breaking up of old and is beginning in the Mediterranean. It is curious that sialic blocks. at the beginning of such a process (for example in the African Rift Valley and the Jordan Valley) negative gra- It is not necessary to discuss the details of this mo- vity anomalies persist, but that they change later on to del. Attempts to pack all details of the Earth's history positive anomalies (as in the Red Sea and in the Medi- into one simple system were made before NOPCSAand terranean)." NOPCSA referred to HOLMES(1931) and after him (GREENE,1982). Many of his results and argu- pointed out that movement of continental land-masses ments of details of plate movement sound very modern. is feasible because of the possibility of the presence of In contrast to the majority of professional geologists, deep-lying currents. We will not discuss the many data NOPCSAwas an uncompromising "continental drifter" and arguments which NOPCSAused for reconstructing and he always fascinates by the creativity of his ideas plate movements. It suffices to mention that he critici- and by the fact that he could bring together data which sed WEGENER(1929) and other authors for assuming hitherto had been seen in isolation. He must have that continetal blocks drift parallel to each other. Rat- known AMPFERER'SUnterströmungstheorie but he never her, he pointed out, one has to assume rotations and commented on it. He was well aware of HOLMES'and not carry the fitting of the continents according to their KIRSCH'Sideas of radioactivity as a motor for crustal outlines too far (as WEGENERhad done) because during movement but unfortunately did not see the possibilities drifting and rotation, the outlines of the continental of this model and took recourse to ad hoc explanations blocks are modified. The model which NOPCSAdevel- like "Polflucht" a,nd centrifugal force. oped is highly speculative (which NOPCSAwould prob- In a very important contribution, SENGÖR(1982) dis- ably easily have admitted). He searched for a picture of cusses the 1920 -1950 schools of thought in global tec- the structure of the Earth and its development which is tonics. He shows that the distinction between "fixists" as highly patterned and regular as possible. He quoted and "mobilists" does not pinpoint the real issues be- attempts in the same direction by earlier authors but he tween the different schools of thought. Issues which went much further than they. In order to understand the characterize the (fixist) "KOBER-STillE school" are an- current structure of the crust, the Atlantic first must be tiuniformitarianism, catastrophism and a belief in an or- closed to reveal two orogenetic belts both of which derly, regular Nature (hence the search for regular tec- have the size of great circles. These are the equatorial tonic cycles, for regular intervals between glaciations, belt, which can be traced not only from Spain to East etc.). The mobilistic "WEGENER-ARGANDschool" on the Asia, but also across the Pacific to Peru and from there other hand had a strong uniformitarian tendency and back to Spain, and the longitudial belt, again a great believed in an inherently irregular Nature, in which circle, which surrounds the Pacific. Local disturbances probability rather than determinism was believed to be in the direction and structure fo both belts are ex- a realistic approach. SENGÖRshows that, despite the plained by local geological conditions. The significance fact that he was a fixist and accepted contraction theory of the belts is shown by magmatism but also by the di- (see also GREENE,1982), SUESS' uncompromising uni- stribution of earthquakes. A special feature of the longi- formitarianism made him a predecessor of the WEGE- tudinal belt is festoon-shaped island arcs on both sides NER-ARGANDschool. He not only paved the way for the of the Pacific which all have an eastward convexity. As nappists in the last quarter of the 19th century, but also both orogenetic belts formed since the middle Mesozo- for the mobilists in the early 20th century.

Despite the fact that no documents are known which many noteworthy theoretical contributions in geology illuminate SUESS'Sinfluence on NOPCSA,SENGÖR'Seva- and evolutionary biology. luation of SUESScontributes much to an understanding NOPCSA'Swork was most often accepted only if it was of NOPCSA'Sideas on theoretical geology. NOPCSAhad descriptive (viz. his dinosaur and geological work); only no problems accepting the nappist model and conti- some more esoteric studies were not purposefully ne- nental drift, making very fruitful use of both theories. glected in German-speaking nations and elsewhere. In- However, he preferred the determinism which characte- deed, he was invited to England on several occasions rized the KOBER-STILLEschool (which began in the early to study and deliver talks on dinosaurs and his travels. 1920s) to the indeterminism of the WEGENER-ARGAND For many of his readers of the last 30 years, primarily school. It is perhaps not by coincidence that this very among English-speaking workers and also due in large last paper tried to develop a highly ordered picture of part to a one-sided account of NOPCSA'Slife by EDINGER the history of the Earth which looks familiar if one (1955), repeated by COLBERT(1968), NOPCSAis known knows STILLE'Swork, but which would never have been primarily for his heterodox behavior. (Without a doubt, acceptable for the WEGENER-ARGANDschool. NOPCSAcontributed in many ways to his own unusual mystique). We argue here that he clearly was no exotic nor a complete outsider to vertebrate paleontology of the early 20th century. Because he was very well read 11. Conclusions and eclectic in his work, he contributed vast amounts of primary literature and synthetic views on theoretical Judging from what we know about his life and what tectonic geology, neo-Lamarckian evolution and paleo- can be gleaned from his works, NOPCSAas a scientist is biology of reptiles. Also throughout most of his career, somewhat of a paradox. He was the last of a line of he focused his attention on the conflict of maintaining Hungarian nobility and hence never knew the need for broad scientific perspectives in the face of the ever-in- employment before World War I. He was sent to the creasing subdivision of scientific disciplines. He spoke Theresianum, like other children of the nobility, to gain out loudly for tearing down the boundaries and filling a respectable education. However, at this time in his unoccupied interstices between disciplines: his specia- life, NOPCSAbroke with tradition. He did not continue on list versus generalist presentation in Budapest (1928b). in university with a degree in law, economics or agricul- Only through maintaining a manysided theoretical view ture; rather he furthered himself in natural sciences. into which many seemingly unrelated phenomena come Whether this pathway began through his coincidental to play could important issues be settled (viz. his work encounter with dinosaur bones from the family's estate on tectonics and biogeography). To continually subdi- (and then to SUESS)we do not know. Yet it is clear that vide science is to continually lose the connections NOPCSAadvanced himself in vertebrate paleontology at among associated bits of knowledge. Indeed, NOPCSA'S his own hands, as SUESSbefore him had done for him- overall research programme was not so much the tack- self in geology (GREENE, 1982). At the same time, ling of large empirical problems or the development of NOPCSAlearned his geology directly from SUESS. an encompassing theory. Nor was he simply or strictly Because of his financial independence, NOPCSAwas a solver of evolutionary, paleontological or geological not forced to seek a scientific education in order to be riddles. Rather, it seems that the overall purpose to his employed. An autonomous career in vertebrate paleon- work was to stress the bringing together of facts and tology was one of his choosing, and in the days before events which escape the perspective of the specialist. World War I, he felt free to move to Vienna, with its NOPCSAis perhaps the best remembered for campaign- important resources, rather than live on his Hungarian ing for a new eclectic and synthetic intellectual beha- estate. His heterodox life style and behavior as a scien- vior. tist can also be attributed to NOPCSA'Sfortunes. Be- cause he did not depend on the hierarchically struc- Acknowledgements tured and constraining university system, his attitudes We thank F. WESTPHAL,for critically reviewing the manu- and enterprises could remain relatively uncompromising script, and J. H. OSTROMand J. S. MciNTOSH,for information and his behavior bordering on the arrogant. Yet to cha- on Archaeopteryx and sauropods, respectively. U. VON HUENE racterize NOPCSAas arrogant is to overlook the obvious kindly provided information on F. BARONNOPCSAand F. VON problem of combining in one person a high level of in- HUENE.We are grateful to H. WORNERfor typing the manus- telligence and creativity not often tempered with the cript. ability of self-criticism. NOPCSA'S studies in tectonic geology, evolutionary biology, paleobiogeography and sexual dimorphism prove his ability to intelligently dis- References cover problems and solve them in remarkable ways. ABEL, 0.: Die Rekonstruktion des Diplodocus. - Abh. Zool.-Bot. This skill seems to have been lacking among many of Ges. Wien, 5, 1-59, Wien 1910. NOPCSA'Scontemporaries. Yet, in retrospect, NOPCSA ABEL, 0.: Grundzüge der Paläobiologie der Wirbeltiere. - sometimes failed to ponder the overall feasibility of his 708 pp., Stuttgart (Schweizerbart) 1912. hypotheses and arguments. Nor did he seem to care ABEL, 0.: Das biologische Trägheitsgesetz. - Biologia genera- much for selling his ideas; he often did not think very lis, 4, 1-102, Wien 1928. highly of the general readers of his papers. The inability ABEL, 0.: Paläobiologie und Stammesgeschichte. - 423 pp., Jena (Gustav Fischer) to criticize his own work acted both against and for 1929. ALLIN, E. F.: Evolution of the mammalian middle ear. - J. NOPCSA.Against because of outlandish and easily fal- Morph., 147, 403-437, Philadelphia 1976. sified ideas which he presented on paper; and for, be- AMPFERER,0.: Über das Bewegungsbild von Faltengebirgen. cause he excelled at assembling disparate ideas into - Jahrb. Geol. A.-A. 56, 539-622, Wien 1906. new frameworks. As such, NOPCSAwas one of the first AMPFERER,0.: Über Kontinentalverschiebungen. - Die Natur- great theorists in vertebrate paleontology and made wiss. 13, 669-675, Berlin 1925.

AMPFERER,O. & HAMMER,W.: Geologischer Querschnitt durch DAVITASHVILI,L.: [The Theory of Sexual Selection]. - 538 pp., die Ostalpen vom Allgäu zum Gardasee. - Jahrb. Geol. R.- Moskov (Akad. Nauk. SSSR) 1961 (in Russian). A. 61, 531-710, Wien 1911. DE BEER, G.: Archaeopteryx lithographica. - 68 pp., London (Brit. Anonymus: Personalia. - Centralbl. Min. Geol. Paläont., 1933, Mus. [Nat. Hist]) 1954. 368, Stuttgart 1933. DE BEER, G.: Archaeopteryx. -In: A. L. THOMSON(Ed.): A New Dictionary of Birds, 58-62, London (Nelson Co.) 1964. BAKKER, R. T.: Dinosaur heresy - dinosaur renaissance - DeMAR, R. & BARGHUSEN,H. R.: Mechanics and evolution of why we need endothermic archosaurs for a comprehensive the synapsid jaw. - Evolution, 26, 622-637, Lancaster theory of bioenergetic evolution. - In: THOMAS,R. D. K. & 1972. OLSON, E. C. (Eds.): A Cold Look at the Warm-Blooded Di- DODSON,P.: Taxonomic implications of relative growth in lam- nosaurs, 351-462, Boulder, Colorado (Westview Press) beosaurine hadrosaurs. - Syst. Zool.,24, 37-54, Lawrence 1980. 1975. BAKKER, R. T. & GALTON, P. M.: Dinosaur monophyly and a DODSON,P.: Quantitative aspects of relative growth and sexual new class of vertebrates. - Nature, 248, 168-172, London dimorphism in Protoceratops. - J. Paleont., 50, 929-940, Tul- 1974. sa 1976. BARGHUSEN,H. R.: The lower jaw of cynodonts (Reptilia, The- DOLLO,L.: Quatrieme note sur les dinosauriens de Bernissart. rapsida) and the evolutionary origin of mammal-like adductor - Bull. Mus. Roy. Hist. Nat. Belg., 2, 223-248, Bruxelles jaw musculature. - Postilla, 116, 1-49, New Haven 1968. 1883. BEEBE,C. W.: A Tetrapteryx stage in the ancestry of birds. - DOLLO,L.: Cinquieme note sur les dinosauriens de Bernissart. Zoologica, 2, 39-52, New York 1915. - Bull. Mus. Roy. Hist. Nat. Belg., 3, 129-146, Bruxelles BOCK,W. J.: The role of adaptive mechanisms in the origin of 1884. higher levels of organization. - Syst. Zool., 14, 272-287, DOUVILLE,H.: Sur Ie genre Kerunia. - Bull. Soc. Geol. France Lawrence 1965. (IV), 6, 129-141, Paris 1907. BONAPARTE,J.: Pisanosaurus mertii CASAMIQUELAand the origin of the Ornithischia. - J. Paleont., 50, 808-820, Tulsa 1976. EDINGER,T.: NOPCSA.- Anat. Anz., 79, 59-62, Jena 1934. BRADU,D. & GRINE,F. E.: Multivariate analysis of diademodon- EDINGER,T.: Personalities in Paleontology. Nopcsa. - Soc. tine crania from South Africa and Zambia. - S. Afr. J. Sei., Vert. Paleont. News Bull., 43, 35-39, New York 1955. 75, 441-448, Johannesburg 1979. EHRENBERG,K.: OTHENIOABEL'S Lebensweg. - 162 pp., Wien BRAMBLE,D. M.: Origin of the mammalian feeding complex: (Österr. Hochschülerschaft, Univ. Wien) 1975. models and mechanism. - Paleobiology, 4, 271-301, Law- ENLOW,D. H. & BROWN,S.O.: A comparative histological stu- rence 1978. dy of fossil and Recent bone tissues. Part I. - Texas J. Sei., BRINKMANN,R.: EMANUELKAYSER'SAbriß der Geologie. - Band 8, 405-443, Austin 1956. 1: Allgemeine Geologie, 282 S., 6. Auflage, Stuttgart (Ferdi- ENLOW,D. H. & BROWN,S.O.: A comparative histological stu- nand Enke) 1940. dy of fossil and Recent bone tissues. Part II. - Texas J. BRINKMANN,R.: Abriß der Geologie. - Band 1: Allgemeine Sei., 9, 186-214, Austin 1957. Geologie, 268 S., 10. Auflage, Stuttgart (Ferdinand Enke) ENLOW,D. H. & BROWN,S.O.: A comparative histological stu- 1967. dy of fossil and Recent bone tissues. Part III. - Texas J. BROILl, F.: Über den feineren Bau der "verknöcherten Sehnen" Sei., 10, 187-230, Austin 1958. (= verknöcherten Muskeln) von Trachodon. - Anat. Anz., 55, FRAAS, E.: Eine recente Kerunia-Bildung. - Mitt. K. K. Zool.- 465-475, Jena 1922. Botan. Gesellsch. Wien, 1911, 2-8, Wien 1911. BROILl, F. & SCHRÖDER,J.: Über einige neue Anomodontier aus der Tapinicephalus-Zone. - Sitzungsber. Math.-Naturwiss. GREAVES,W. S.: The jaw lever-system in ungulates: a new mo- Abt. Bayer. Akad. Wiss., 2, 118-136, München 1937. del. - J. Zoo I. Lond., 184, 271-285, London 1978. BROWN,B.: Anchiceratops, a new genus of horned dinosaurs from GREENE,M. T.: Geology in the Nineteenth Century. - 324 pp., the Edmonton Cretaceous of Alberta with discussion of the Ithaca and London (Cornell University Press), 1982. origin of the ceratopsian crest and the braincasts of Anchice- GRINE, F. E., HAHN, B. D. & Gow, C. E.: Aspects of relative ratops and Trachodon. - Bull. Amer. Mus. Nat. Hist., 33, growth and variability in Diademodon (Reptilia; Therapsida). - 539-548, New York 1914. S. Afr. J. Sei., 74, 50-58, Johannesburg 1978. BROWN,B. & SCHLAIKJER,E. M.: The structure and relationship GRINE,F. E. & HAHN,B. D.: Allometric growth in the Diademo- of Protoceratops. - Ann. N. Y. Acad. Sei., 40, 133-266, New dontinae (Reptilia; Therapsida): a preliminary report. - Pa- York 1940. laeont. Afr., 21, 161-166, Johannesburg 1978. HALLAM,A.: A Revolution in the Earth Sciences. From Contin- CAPLE, G., BALDA, R. P. & WILLIS, W. R.: The physics of lea- ental Drift to Plate Tectonics. - 127 pp., Oxford (Clarendon ping animals and the evolution of preflight. - Amer. Nat., Press) 1973. 121, 455-467, New York 1983. HATCHER,J. B.: Diplodocus (MARSH): its osteology, taxonomy CHANNELL,J. E. T., D'ARGENIO,B. & HORVAT,F.: Adria, the Af- and probable habits, with a restoration of the skeleton. - rican promontory, in Mesozoic Mediterranean paleogeogra- Mem. Carnegie Mus., 1, 1-63, 1901. phy. - Earth Science Reviews, 15, 213-292, Amsterdam HAY, O. P.: On the manner of locomotion of the dinosaurs, 1979. especially Diplodocus, with remarks on the origin of the birds. CHARIG,A. J.: "Dinosaur monophyly and a new class of verte- - Proc. Washington Acad. Sei., 12, 1-25, 1910. brates": a critical review. - In: A. d'A.BELLAIRS& C. B. Cox HECHT,M. K. & TARSITANO,S.: The paleobiology and phyloge- (Eds.): Morphology and Biology of Reptiles. - Linn. Soc. netic position of Archaeopteryx. - Geobios Mem. Sp., 6, Lond. Symp., 3, 6-104, London 1976. 141-149, Lyon 1983. CHARIG,A. J.: Problems in dinosaur phylogeny: a reasoned ap- HENNIG,E.: Zum Wesen und Problem der Anpassung. - Die proach to their attempted resolution. - Geobios Mem. Sp., Naturwissenschaften, 16 (29), 571-574, Berlin 1928. 6, 113-126, Lyon 1982. HENNIG,E.: Wesen und Wege der Paläontologie. - 512 pp., CLOOS,H.: Einführung in die Geologie. Ein Lehrbuch der inne- Berlin (Gebr. Borntraeger) 1932. ren Dynamik. - 503 pp., Berlin (Gebr. Borntraeger) 1936. HEILMANN,G.: The Origin of Birds. - 208 pp., New York (D. COLBERT,E. H.: Men and Dinosaurs. - 283 pp., New York Appleton Co.) 1927. (E. P. Dutton & Co.) 1968. HOHL, R.: Geotektonische Hypothesen. - In: Die Entwick- lungsgeschichte der Erde, Band I, 279-320, Hanau/Main DACQUE,E.: Vergleichende biologische Formenkunde der fos- (Werner Dausier) 1971 (originally published in the DDR). silen niederen Tiere. - 777 pp., Berlin (Gebr. Borntraeger) HOLLAND,W. J.: The osteology of Diplodocus MARSH. - Mem. 1921. Carnegie Mus., 2, 225-278, 1906.

HOLMES, A.: Radioactivity and earth movements. - Trans. LULL, R. S. & WRIGHT, N. E.: Hadrosaurian dinosaurs of North geol. Soc. Glasgow, 18, 559-606, 1929. America. - Geol. Soc. Amer. Sp. Paper 40, 1-135, Boulder HOLMES, A.: Radioaktivität und Geologie. - Verh. Nat. Ges. 1942. Basel, 41, 136 ff., Basel 1931. LUTHER, A.: Über die von N. trigeminus versorgte Muskulatur HOPSON, J. A.: The evolution of cranial display structures in der Amphibien. - Acta Soc. Sci. Fennicae, 44, 1-151, Hel- hadrosaurian dinosaurs. - Paleobiology, 1, 21 -43, Lawren- sinki 1914. ce 1975. MANTELL, G. A.: On the Pelorosaurus, an undescribed gigantic HORNER, J. R.: Evidence of colonial nesting and "site fidelity" terrestrial reptile, whose remains are associated with those among ornithischian dinosaurs. - Nature, 297, 675-676, of the Iguanodon and other saurians in the strate of Tilgate London 1982. Forest, in Sussex. - Phil. Trans. Roy. Soc. Lond., 140, HORNER, J. R. & MAKELA, R.: Nest of juveniles provides evi- 379-390, 1850(a). dence of family structure among dinosaurs. - Nature, 282, MANTELL, G. A.: On a dorsal dermal spine of the Hylaeosaurus re- 296-298, London 1979. HUENE, F. VON: Über die Zweistämmigkeit der Dinosaurier, mit cently discovered in the strate of Tilgate Forest. - Phil. Trans. Roy. Soc. Lond., 140, 391-392, 1850(b). Beiträgen zur Kenntnis einiger Schädel. - N. Jb. Min. Geol. MARSH, O. C.: Odontornithes: a monograph on the extinct too- Paläont., 37, 577-589, Stuttgart 1914. thed birds of North America. - Prof. Paper Engineer. Dept. HURST, C. H.: The structure and habits of Archaeopterix. - Nat. U.S. Army, 18, 1-201, Washington 1880. Sci., 6, 112-122, 180-186, 244-248, New York 1895. MARSH, O. C.: The skull and brain of Claosaurus. - Amer. J. Sci. sero 3, 55, 83-86, New Haven 1893. JAEKEL, 0.: Darwinismus und Descendenzlehre. - 18 pp., Ber- MAYER-EYMAR, K.: Un singulier cephalopod de l'Eocene d'E- lin (Himmel und Erde Verlag) 1894. gypte (Kerunia cornuta). - Eclog. Geol. Helvet., 6, 120-121 JAEKEL, 0.: Über verschiedene Wege phylogenetischer Ent- (Abstr.), Basel 1900. wicklungen. - Jena (G. Fischer) 1902. MAYER-EYMAR, K.: Interessante neue Gastropoden aus dem JAEKEL, 0.: Wege und Ziele der Paläontologie. - Paläont. Z., Untertertiär Ägyptens. - Vierteljahrschrift naturf. Ges. Zü- 1, 1-72, Stuttgart 1913. rich, 46, 31-32, 1901. JELETZKY, J. A.: The allegedly Danian dinosaur-bearing rocks MOLNAR, R. E.: A dinosaur from New Zealand. - In: CRESS- of the globe and the problem of the Mesozoic-Cenozoic WELL M. M. & VELLA, P. (eds.): 5th International Gondwana boundary. - J. Paleont., 37, 1005-1018, Tulsa 1963. Symp., 91-96, Rotterdam (A. A. Balkema) 1980.

NÖLKE, F.: Geotektonische Hypothesen. Eine kritische Zusam- KETTNER, R.: Allgemeine Geologie I. Der Bau der Erdkruste. - menstellung. - 128 pp., Berlin (Gebrüder Borntraeger) 412 pp. Berlin (East) (VEB Deutscher Verlag der Wissen- 1924. schaften) 1958. NOPCSA, F.: Dinosaurierreste aus Siebenbürgen. I. Schädel KIRSCH, G.: Geologie und Radioaktivität. Die radioaktiven Vor- von Limnosaurus transylvanicus novo gen. et spec. - Denkschr. gänge als geologische Uhren und geophysikalische Energie- Akad. Wiss. Wien, 68, 555-591, Wien 1900. quellen. - 214 pp., Wien und Berlin (Julius Springer) 1928. NOPCSA, F.: Dinosaurierreste aus Siebenbürgen. II. Schädelre- KIRSCH, G.: Geomechanik. Entwurf einer Physik der Erdge- ste von Mochlodon. - Denkschr. Akad. Wiss. Wien, 72, schichte. - 150 pp., Leipzig (Johann Ambrosius Barth) 149-175, Wien 1902. 1938. NOPCSA, F.: Weitere Schädelreste von Mochlodon. - Denkschr. KRAUS, E.: Die Entwicklungsgeschichte der Kontinente und Akad. Wiss. Wien, 74, 229-264, Wien 1903(a). Ozeane. - 258 pp., Berlin (Akademie Verlag) 1959. NOPCSA, F.: Origin of the Mosasaurus. - Geol. Mag., 1903, KRIPP, D. VON: Der Oberschnabel-Mechanismus der Vögel. - 119-121, London 1903(b). Morph. Jahrb., 71, 469-544, Jena 1933(a). NOPCSA, F.: Notes on the British dinosaurs. I. Hypsilophodon. - KRIPP, D. VON: Die Spezialisationsreihe der Störche, Reiher Geol. Mag., 1905, 203-208, London 1905(a). und Kormorane vom konstruktiven und biotechnischen NOPcsA, F.: Notes on the British dinosaurs. II. Polacanthus. - Standpunkt. - Morph. Jahrb., 72, 60-92, Jena 1933(b). Geol. Mag., 1905, 241-250, London 1905(b). KRIPP, D. VON: Die Kaubewegung und Lebensweise von Edmon- NOPCSA, F.: Notes on the British dinosaurs. II. Streptospondylus. tosaurus spec. auf Grund der mechanisch konstruktiven Ana- - Geol. Mag., 1905, 289-293, London 1905(c). lyse. - Palaeobiologica, 5, 409-422, Wien 1933(c). NOPCSA, F.: Kerunia, a symbiosis of a hydractinian with a ce- KUBACSKA, A. T.: FRANZ BARON NOPCSA. - 295 pp., Budapest phalopod. - Annals Mag. Nat. Hist. (7), 16, 95-101, Plate (Ungarisches Naturwiss. Mus.) 1945. III, London 1905(d). NOPCSA, F.: Remark on the supposed clavicle of the sauropo- LAKJER, T.: Studien über die Trigeminus-versorgte Kaumusku- dous dinosaur Diplodocus. - Proc. Zoo I. Soc. Lond., 1905, latur der Sauropsiden. - 154 pp., Copenhagen (C. A. Reit- 241 -250, 1905(e). zel) 1926. NOPCSA, F.: Neues aus Nordalbanien. - Centralbl. Geol. Pa- LAMBE, L. M.: The hadrosaur Edmontosaurus from the Upper Cre- läont., 1906, 69-76, Stuttgart 1905(f). taceous of Alberta. - Mem. Canadian Geol. Surv., 120, NOPCSA, F.: Ideas on the origin of flight. - Proc. Zool. Soc. 1- 79, Ottawa 1920. Lond., 1907, 223-236, 1907. LAMBRECHT, K.: FRANZ BARON NOPCSA t, der Begründer der NOPcsA, F.: On the systematic position of the Upper Cretace- Paläphysiologie, 3. Mai 1877 bis 25 April 1933. - Paläont. ous dinosaur Titanosaurus. - Geol. Mag., 1910, 261, London Zeitsehr., 15, 201-211, Stuttgart 1933. 1910. LAMBRECHT, K., QUENSTEDT, W. & QUENSTEDT, A.: Palaeontolo- NOPCSA, F.: Notes on the British dinosaurs. IV. Stegosaurus pris- gi. Catalogus bio-bibliographicus. - Fossilium Catalogus I, cus. - Geol. Mag., 1911, 109-115, 145-153, London 1911. Pars 72, 495 pp., s'Gravenhage (W. Junk) 1938. NOPCSA, F.: Notes on the British dinosaurs. V. Craterosaurus. - LAPPARENT, A. F. DE: Les dinosauriens du Cretace superieur Geol. Mag., 1912, 481-484, London 1912. du Midi de la France. - Mem. Soc. Geol. Fr. n. s., 26, NOPCSA, F.: Über Geschlechtsunterschiede bei Dinosauriern. 1-54, Paris 1947. - Centralbl. Min. Geol. Paläont., 1915, 385-388, Stuttgart LAPPARENT, A. F. DE & AGUIRE, E.: Presence de dinosauriens 1915. dans Ie Cretace superieur du Bassin de Tremp (Province de NOPCSA, F.: Die Riesenformen unter den Dinosauriern. - Cen- Levida, Espagne). - C. R. Soc. Geol. France, 1956, tralbl. Min. Geol. Paläont., 1917, 332-348, Stuttgart 261-262, Paris 1956. 1917(a). LULL, R. S.: The cranial musculature and the origin of the frill NOPCSA, F.: Über Dinosaurier I. Notizen über die Systematik in ceratopsian dinosaurs. - Amer. J. Sei. sero 4, 25, der Dinosaurier. - Centralbl. Min. Geol. Pläont., 1917, 387-399, New Haven 1908. 204-213, Stuttgart 1917(b).

NOPCSA,F.: Neues über Geschlechtsunterschiede bei Ortho- NOPCSA,F.: Notizen zur Großtektonik der Erde. - Centralblatt poden. - Centralbl. Min. Geol. Paläont., 1918, 186-198, Min. Geol. Paläont. 1933, 369-379, Stuttgart 1933. Stuttgart 1918(a). NOPCSA,F.: The influence of geological and climatological fac- NOPCSA,F.: Leipsanosaurus n. gen. Ein neuer Thyreophore aus tors on the distribution of non-marine fossil reptiles and Ste- der Gosau. - Földt. Közl., 48, 324-328, Budapest 1918(b). gocephalia. - Quart. J. Geol. Soc. Lond., 60, 76-140, Lon- NOPCSA,F.: Zur Arbeitsmethode Professor L. KOBERS.- Ver- don 1934. hand I. Geol. Staatsanstalt, 1921, 198-199, Wien 1921. NOPCSA,F. & HEIDSlECK,E.: On the histology of the ribs in im- NOPCSA,F.: Bemerkungen zur Systematik der Reptilien. - Pa- mature and half-grown trachodont dinosaurs. - Proc. Zool. läont. Zeitsehr., 5, 107-118, Stuttgart 1922. Soc. Lond., 1, 221-226, London 1933. NOPCSA, F.: Reversible und irreversible evolution. - Proc. Zool. Soc. Lond., 1923, 1045-1059, London 1923(a). OPPENHEIM,P.: Über Kerunia comuta MAYER-EYMAR,aus dem Eo- NOPCSA,F.: Vorläufige Notiz über Pachyostose und Osteoskle- cän Ägyptens. - Centralblatt Min. Geol. Paläont. 1902, rose einiger mariner Wirbeltiere. - Anat. Anz., 56, 44-49, Stuttgart 1902. 353-359, Jena 1923(b). OSBORN,H. F.: Crania of Tyrannosaurus and Allosaurus. - Mem. NOPCSA,F.: On the geological importance of the primitive rep- Amer. Mus. Nat. Hist. (n. s.), 1, 1-30, New York 1912. tilian fauna in the uppermost Cretaceous of Hungary. - OSTROM,J. H.: Cranial morphology of the hadrosaurian dino- Quart. J. Geol. Soc. Lond., 79, 100-116, London 1923(c). saurs of North America. - Bull. Amer. Mus. Nat. Hist., 122, NOPCSA,F.: Die Familien der Reptilien. - Fortsehr. Geol. Pa- 33-186, New York 1961. läont. Heft 2, 1-210, Jena 1923(d). OSTROM,J. H.: The ancestry of birds. - Nature, 242, 136, NOPCSA,F.: On the origin of flight in birds. - Proc. Zool. Soc. London 1973. Lond., 1923, 463-477, London 1923(e). OSTROM,J. H.: Archaeopteryx and the origin of flight. - Quart. NOPCSA, F.: Über eine neue Kreideschlange. - Paläont. Rev. BioI., 49, 27-47, Baltimore 1974. Zeitsehr., 5, 264-265, Stuttgart 1923(f). OSTROM,J. H.: Archaeopteryx and the origin of birds. - BioI. J. NOPCSA,F.: Notes on the British dinosaurs. VI. - Geol. Mag., Linn. Soc. Lond., 8, 91-182, London 1976. 1923, 193-199, London 1923(g). OWEN, R.: Report on British fossil reptiles. Part II. - British NOPCSA,F.: On the systematic position of Saurostemum and Tan- Assoc. Adv. Sei. Rept. 11th Meeting, Plymouth, 1841, gasaurus. - S. Afr. J. Sei., 21, 206-207, Johannesburg 1924. 60-204, 1842. NOPCSA,F.: On some reptilian bones from the Eocene of So- OWEN,R.: Monograph of the fossil Reptilia of the Wealden and koto. - Geol. Surv. Nigeria Occ. Paper 2, 1-15, 1925(a). Purbeck formations. Suppl. II. Crocodilia (Streptospondylus, etc. Wealden). - Monogr. Palaeontogr. Soc. Lond. II, NOPCSA,F.: Die Wirbelsäule von Rhabdodon und Orthomerus. - 20-44, 1859. Palaeont. Hungarica, 1, 273-304, Budapest 1925(b). NOPCSA,F.: Diskussionsbemerkungen zu G. J. VON FEJERVA- PARIS,J. P. & TAOUET,P.: Decouverte d'un fragment de den- RYSVortrag über das DOLLO'sche Gesetz. - Paläont. Z., 7, taire d'hadrosaurien (Reptilien dinosaurien) dans Ie Cretace 184, Stuttgart 1926(a). superieur des Petites Pyrenees (Haute-Garonne). - Bull. NOPCSA,F.: Heredity and evolution. - Proc. Zool. Soc. Part 2, Mus. Natl. Hist. Nat., 3, 17-27, Paris 1973. 1926, 633-665, London 1926(b). PYCRAFT,W. P.: Wing of Archaeopteryx. - J. Oxford Univ. Jr. Sei. NOPCSA,F.: Beiträge zur Verteilung der Eruptivgesteine. - Club, 1, 172-176, 1894(a). Földt. Közl., 56, 149-160, Budapest 1927(a). PYCRAFT,W. P.: The wing of Archaeopteryx I. - Nat. Sei., 4, NOPCSA, F.: Vererbung erworbener Eigenschaften. - Natur 350-360, 437-448, 1894(b). und Museum, 57, 124-128, Frankfurt 1927(b). PYCRAFT,W. P.: The wing of Archaeopteryx II. - Nat. Sei., 5, NOPCSA,F.: Bemerkungen über PETRONIEVICSseine Arbeiten 261-266, 1896. über Archaeopteryx. - Ann. Geol. Penins. Balk., 1927, 105-111, Beograd 1927(c). QUECKETT,J. T.: Descriptive and illustrated catalogue of the NOPCSA,F.: Scolosaurus cutleri, a new dinosaur. - Geol. Hung., histological series contained in the Museum of the Royal 1, 54-74, Budapest 1928(a). College of Surgeons II. Structure of the skeleton of verte- NOPCSA,F.: Festrede gehalten anläßlich des Besuchs der Pa- brate animals, 248 pp., London 1855. läontologischen Gesellschaft. - 15 pp., Magyar Kiralyi Föld- REIF, W.-E.: Paleobiology today and fifty years ago: a review tani Intezet Hazinyomdaja, Budapest 1928(b). of two journals. - N. Jb. Geol. Paläont. Mh. 1980, 361-372, NOPCSA,F.: The Genera of Reptiles. - Palaeobiologica, 1, Stuttgart 1980. 163-188, Wien 1928(c). REIF,W.-E.: The search for a macroevolutionary theory in Ger- NOPCSA,F.: Sur la distribution de principaux types des roches man paleontology. - J. Hist. BioI. (in press). eruptives. - C. R. Congr. Geol. Internatl. Part IV, 1381, Ma- RICOLES,A. DE: Tissue structures of dinosaur bones: functional drid 1928(d). significance and possible relation to dinosaur physiology. - NOPCSA,F.: Sexual differences in ornithopodous dinosaurs. - In: R. D. K. THOMAS& E. C. OLSON(Eds.): A Cold Look at Palaeobiologica, 2, 187-201, Wien 1929(a). the Warmblooded Dinosaurs, 103-139, Boulder, Colorado NOPCSA,F.: Noch einmal Proavis. - Anat. Anz., 67, 265-300, (Westview Press) 1980. Jena 1929(b). RUSSEL,L. S.: Musculature and functions in the Ceratopsia. - NOPCSA,F.: Dinosaurierreste aus Siebenbürgen. V. Struthiosau- Bull. Natl. Mus. Canada Dept. Mines, 77 (Geol. Sero 52), rus transsylvanicus. - Geol. Hungarica Sero Palaeont., 31, 39-48, Ottawa 1935. 247-279, Budapest 1929(c). RUSSEL,L. S.: The crest of the dinosaur Parasaurolophus. - Con- NOPCSA,F.: Notizen über Macrochemus Bassani novo gen. et spec. trib. Roy. Ontario Mus. Palaeont., 11, 1- 5, Toronto 1946. - Centralbl. Min. Geol. Paläont., 1930, 252-253, Stuttgart 1930. SANDBERG,C. G. S.: Geodynamische Probleme I. Isostasie NOPCSA, F.: Osteologia reptilium recentium et fossilium. - und die ursächliche Einheit von Gebirgsbildung und Vulk- Fossilium Catalogus I, 27, 1-391, s'Gravenhage (W. Junk) anismus. - 69 pp., Berlin (Gebrüder Borntraeger) 1924(a). 1931. SANDBERG,C. G. S.: Geodynamische Probleme II. A. Tektonik NOPCSA,F.: Glossen zu E. HAARMANN'SOszillations-Theorie. und Metamorphose. B. Die Widersprüche in der Kontrak- - Centralblatt Min. Geol. Paläont. 1932, Abt. B, (5), tionstheroie. - 51 pp., Berlin (Gebrüder Borntraeger) 265-268, Stuttgart 1932(a). 1924(b). NOPCSA,F.: Zur Geschichte der Adria. Eine tektonische Stu- SCHMIDT,K.: "Subfluenz" und "Subduktion" in den Alpen. - die. - Zeitsehr. deutsch. geol. Ges., 84, 280-316, Berlin Zeitsehr. deutsch. geol. Ges., 127, 53-72, Stuttgart 1976. 1932(b). SCHMIDT-THOME,P.: Gebirgsbildung. - In: MURAWSKI,H. (ed.): NOPCSA, F.: Zur systematischen Stellung von Shinisaurus. - Vom Erdkern zur Magnetosphäre, 85-112, Frankfurt/Main Zoo I. Anz., 97, 185-187, Leipzig 1932(c). (Umschau Verlag) 1968.

SCHMIDT-THOM!:,P.: Tektonik. - In: BRINKMANN,R. (ed): Lehr- VANTYNE, J. &BERGER,A. J.: Fundamentals of Ornithology. - buch der Allgemeinen Geologie. - 579 pp., Stuttgart (Ferdi- 624 pp., New York (Wiley) 1959. nand Enke) 1972. VAUGHN,P.: Seymouria from the lower Permian of southeastern SCHWARZBACH,M.: ALFREDWEGENERund die Drift der Konti- Utah, and possible sexual dimorphism in that genus. - J. nente. - 160 pp., Stuttgart (Wissensch. Verlagsgesellsch.) Paleont., 40, 603-612, Tulsa 1966. 1980. VERSLUYS,J.: Streptostylie bei Dinosauriern nebst Bemerkun- SCHWINNER,R.: Vulkanismus und Gebirgsbildung. Ein Versuch. gen über die Verwandschaft der Vögel und Dinosaurier. - - Z. Vulkanologie, V/4, 175-230, Berlin 1920. Zool. Jahrb. Abt. Anat., 30, 175-260, Jena 1910. VERSLUYS,J.: Das Streptostylie-Problem und die Bewegung im SEELEY,H. G.: The reptile fauna of the Gosau Formation pre- Schädel bei Sauropsida. - Zool. Jahrb. Suppl., 15, served in the Geological Museum of the University of Vien- 545-716, Jena 1912. na. - Quart. J. Geol. Soc. Lond., 37, 620-707, 1881. VERSLUYS,J.: Der Schädel des Skelettes von Trachodon annectens SEELEY, H. G.: On the dinosaurs of the Maastricht beds. - im Senckenberg-Museum. - Abh. Senckenb. Naturf. Ges., Quart. Geol. Soc. Lond., 39, 246-253, 1883. 38, 1-19, Frankfurt 1923. SEELEY,H. G.: On the classification of the fossil animals commonly named Dinosauria. - Proc. Roy. Soc. Lond., 43, WEGENER,A.: Die Entstehung der Kontinente. - Petermanns 165-171, 1887. Mitteilungen, 58, 185-195, 253-256, 305-309, Gotha SEITZ,A. L.: Vergleichende Studien über den mikroskopischen 1912. Knochenbau fossiler und rezenter Reptilien. - Nova Acta WEGENER,A.: Die Entstehung der Kontinente und Ozeane. - Abh. Kaiser Leop. Carol. Deutsch. Akad. Naturforsch., 37, 94 pp., Braunschweig (Sammlung Vieweg) 1915. 230-370, Halle 1907. WEGENER,A.: Die Entstehung der Kontinente und Ozeane. - SENGÖR,A. M. C.: EDUARDSUESS' relation to the pre-1950 4. Aufl., 231 pp., Braunschweig (Vieweg) 1929 (Reprinted: schools of thought in global tectonics. - Geol. Rundschau, Vieweg, Braunschweig, 1980). 71, 381-420, Stuttgart 1982. WEISHAMPEL,D. B.: The evolution of ornithopod jaw mecha- STANNIUS,H.: Die Amphibien. - In: Handbuch der Zootomie. nics. - 226 pp., PhD dissertation. Dept. Geol., Univ. Penn- Zweiter Theil. Die Wirbelthiere (Handbuch der Anatomie der sylvania; Philadelphia, Pennsylvania 1981. Wirbelthiere), 1-270, Berlin (von Veit) 1856. WEISHAMPEL,D. B.: Jaw mechanics of hadrosaurids (Reptilia STEINER,H.: Das Problem der Diastataxie des Vogelflügels. - Ornithischia). - Acta Palaeont. Polonica, 28, (in press a). Jen. Zeitsehr. Naturwiss., 1917, 221-496, Jena. WEISHAMPEL,D. B.: The evolution of jaw mechanisms in orni- STERNBERG,C. M.: Hooded hadrosaurs of the Belly River Se- thopod dinosaurs (Reptilia: Ornithischia). - Adv. Anat. Em- ries of the Upper Cretaceous. - Bull. Natl. Mus. Canada, bryol. Cell. BioI. (in press b). 77, 1-37, Ottawa 1935. WILLISTON,S. W.: Are birds derived from dinosaurs? - Kansas STILLE, H.: Grundfragen der vergleichenden Tektonik. - 443 City Rev. Sci., 3, 457-460, 1879. pp., Berlin (Gebrüder Borntraeger) 1924. WOODWARD,A. S.: BARON FRANCISNOPCSA.- Quart. Journ. SUESS, E.: Das Antlitz der Erde. - Prag (Tempsky); Leipzig Geol. Soc., 90, 48-49, London 1934. (Freytag) 1883. WUNDERLICH,H. G.: Wesen und Ursachen der Gebirgsbildung. SUESS, F. E.: Verschiedene Theorien über die Beziehungen - 367 pp., Mannheim/Zürich (Bibliographisches Institut) der Radioaktivität zu geologischen Vorgängen. - Mitt. d. 1966. Wiener Geol. Ges. 1912, 87-105, Wien 1912. WUNDERLICH,H. G.: Einführung in die Geologie II. Endogene SUESS,F. E.: FRANZBARONNOPCSAt (3. Mai 1877 - 25 April Dynamik. - 223 pp., Mannheim/Zürich (Bibliographisches 1933). - Mitteil. Geol. Ges. Wien, 26, 215-221, Wien 1933. Institut) 1968. ZAPFE, H.: Nopcsa von Feslöszilvas. - Osterreichisches bio- TARSITANO,S. & HECHT,M. K.: A reconsideration of the repti- graphisches Lexikon 1815-1950, 7, 148, Wien (Osterr. lian relationships of Archaeopteryx. - Zool. J. Linn. Soc., 69, Akad. Wissensch.) 1978. 149-182, London 1980. ZElL, W.: BRINKMANNSAbriß der Geologie. - Band 1: Allgemei- THULBORN,R. A. & HAMLEY,T. A.: The reptilian relationships of ne Geologie, 246 pp., 11. Auflage, Stuttgart (Ferdinand En- Archaeopteryx: a reconsideration. - Australian J. Z601., 30, ke) 1975. 611-634, Sydney 1982. TORNIER,G.: Wie war der Diplodocus carnegiei wirklich gebaut? - Manuskript bei der Schriftleitung eingelangt am 10. November Sitzungsb. Ges. Naturf.Freunde Berlin 1909, 193-209. 1983.

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