Versão Do Arquivo Anexado / Version of Attached File: Versão Do Editor / Published Version

UNIVERSIDADE ESTADUAL DE CAMPINAS SISTEMA DE BIBLIOTECAS DA UNICAMP REPOSITÓRIO DA PRODUÇÃO CIENTIFICA E INTELECTUAL DA UNICAMP

Versão do arquivo anexado / Version of attached file: Versão do Editor / Published Version

Mais informações no site da editora / Further information on publisher's website: https://www.tandfonline.com/doi/full/10.1080/08912963.2017.1335311

DOI: 10.1080/08912963.2017.1335311

DIRETORIA DE TRATAMENTO DA INFORMAÇÃO

Cidade Universitária Zeferino Vaz Barão Geraldo CEP 13083-970 – Campinas SP Fone: (19) 3521-6493 http://www.repositorio.unicamp.br Historical Biology, 2018 VOL. 30, NO. 8, 1084–1093 https://doi.org/10.1080/08912963.2017.1335311

Early twentieth-century paleontological research of Freidrich von Huene: contributions to the knowledge of Late Cretaceous vertebrates of Central Brazil

Carlos Roberto A. Candeiroa,b and Silvia Fernanda M. Figueirôac

aLaboratório de Paleontologia e Evolução, Curso de Geologia, Universidade Federal de Goiás, Campus, Aparecida de Goiânia, Brazil; bPrograma de Pós-Graduação em Política Científica eT ecnológica, Instituto de Geociências, Universidade Estadual de Campinas, Campinas, Brazil; cPrograma de Pós- Graduação Multiunidades em Ensino de Ciências e Matemática, Faculdade de Educação, Universidade Estadual de Campinas, Campus Universitário “Zeferino Vaz”, Campinas, Brazil

ABSTRACT ARTICLE HISTORY Friedrich von Huene’s important scientific research on tetrapod fossils from Central Brazil has received little Received 26 March 2017 attention, even though it represents a significant contribution to early studies of vertebrate paleontology Accepted 24 May 2017 in the country including the first discoveries of dinosaur fossils in Brazil. von Huene described five reptile KEYWORDS taxa in two papers published in important German scientific journals and used these specimens to make von Huene; Central Brazil; paleogeographic inferences regarding the Cretaceous of the Southern Hemisphere. von Huene warrants Upper Cretaceous; vertebrate recognition as the first specialist to describe in detail the fossil reptile fauna of Central Brazil, which was paleontology made possible only by his network of contacts with important and influential naturalists of the cities of Rio de Janeiro and São Paulo.

Introduction paleontology – they were the first formal descriptions of Cretaceous taxa from the region – the records described in these The scientific studies of the Triassic by German paleontologist papers would later be assigned to different areas of Gondwana. Friedrich von Huene (1875–1969) during his passages through Furthermore, these studies remained overlooked and rarely cited the Southern Hemisphere are well known; however, his many in paleontological and geological literature, and even in the field important studies of Cretaceous reptiles from Argentina, Brazil, of history of geoscience. India, Madagascar, and Uruguay remain overlooked (Huene 1925, 1927a, 1927b, 1927c, 1928a, 1928b, 1929a, 1929b, 1929c, The prolific scientific life of von Friedrich von Huene is rep- 1929d, 1930, 1933, 1934; Huene & Matley 1933). The most crucial resented by a large legacy of scientific production, in addition to period of Huene’s copious scientific production on Cretaceous his many descriptions of dinosaur specimens. Most of his work dinosaurs was probably between 1920 and 1935, as exempli- on the Cretaceous of Central Brazil, and his research in general, fied by his well-known paper Los saurisquios y ornitisquios del consisted of osteological descriptions, stratigraphic surveys, and Cretáceo Argentino (Huene 1929a), as well as other notable works concise mappings focused exclusively on reptiles. His research (e.g. Huene 1927b, 1927c; Huene & Matley 1933). His remarkable on sauropods (herbivorous dinosaurs), theropods, and crocodyl- study of Cretaceous reptiles of Gondwana brought great visibility iform postcranial remains, projected him onto the national and to the fossil herpetofauna of Argentinian Patagonia. However, international scientific scene as the first paleontologist special- elsewhere in South America, such as his first descriptions of ising on Brazilian Cretaceous archosauromorphs. important fossils from western São Paulo State and from the There remains to date no literature available that specifically Triângulo Mineiro region, have been neglected. analyses the contributions of Friedrich von Huene’s studies of These studies, to a certain extent, followed the pattern used the Brazilian Cretaceous to paleontology and the history of by Huene to describe other specimens with the use drawings geoscience. The researchers who have investigated his scientific and detailed descriptions. His taxonomic assignments were more career have focused almost entirely on his contributions to ver- precise than those of other paleontologists of the time who were tebrate paleontology through his studies of the Cretaceous of publishing studies on Cretaceous fossils from Central Brazil [e.g. Central Brazil (e.g. Candeiro 2011; Candeiro & Figueirôa 2012, Orville Adelbert Derby (1851–1915) and Herbert Huntington 2014; Peyerl et al. 2015), which include some bibliographic sur- Smith (1851–1919)]. Huene published three scientific papers veys. These studies show that – after more than 90 years – the specifically about the Cretaceous of the states of Minas Gerais descriptions of Brazilian fossils made by Huene are still valid. and São Paulo. Despite their importance to South American His pioneering works with reptile remains from the Cretaceous

Published online 16 Jul 2017 HISTORICAL BIOLOGY 1085 of Central Brazil are of great value as part of the historical and Triassic reptiles from the Upper Germanic Trias. In 1902, he scientific legacy of Brazilian paleontology. defended his doctoral thesis, Uebersicht der Reptilien der Trias Based on the literature and sources from the fields of history (An overview of Triassic reptiles) (Huene 1902), in which he of science, geology, and paleontology, the present paper aims described several important new rauisuchid (basal archosaurian) to describe von Huene’s pioneering studies on Late Cretaceous taxa. reptiles of the Bauru Group in western São Paulo state and the The interest and knowledge acquired during his PhD allowed Triângulo Mineiro region conducted between 1920 and 1930. Huene, in his own words, ‘to organize the richness and the diversity of the reptilian fauna of the Triassic’ (1902; sic Schultz & Data and research Holz 1996; Holz & Schultz 1998) into works of description and classification. von Huene was interested in evolutionary research All information described herein is based on the studies of the and published treatments of tetrapod evolution and paleoecology German paleontologist Friedrich von Huene performed between (Huene 1956; Reif 1984). 1920 and 1930 describing and depicting the first dinosaurs and other fossil reptiles found in the Bauru Group in the regions of western São Paulo state and the Triângulo Mineiro. The analysis The research of Friedrich von Huene on Cretaceous fossils of original documents followed the hermeneutic method: inves- of South America tigation and interpretation of the information provided by the Friedrich von Huene is renowned as one of the vertebrate pale- sources (Alberti 1996; Seebohm 2004). ontologists who vigorously studied the Mesozoic of Germany, Austria, Italy, and Britain (see Sues 1997; Turner 2011). He had von Huene’s Late Cretaceous reptiles from Central developed a broad professional network among renowned pale- Brazil ontologists from Europe, the United States, and Argentina, which is well evidenced by his abundant correspondence (Figure 1). A short biography of von Huene The interest of von Huene turned to the Late Triassic period Friederich von Hoyningen (original name of the family that was of Europe, especially of Bavaria and Baden-Württemberg. He later altered to Hüene or Huene; Schultz & Holz 1996) was born greatly expanded previous studies, primarily with records of in the city of Tübingen, Germany, on 22 March 1875. His family Plateosaurus, a prosauropod found in southern Germany. He was from Estonia and, according to Schultz and Holz (1996) and also reviewed all previous classifications of dinosaurs, that col- Turner (2009), is traceable back to the 1600s in this region. His laborated for reassertion and in 1914, based on many osteological mother, Alexandra Baronesse Stackelberg, was from an Estonian characteristics, published a classification system of Saurischia noble family, which enabled Huene to have a good education. and Ornithischia for the dinosaurs that is still widely used today His father, Heinrich A.N. von Hoyningen, was professor of eccle- (see Huene 1908; Sues 1997). siastical history at the Imperial University of Dorpat (current The many descriptions Huene produced of reptile species Tartu, second largest city of Estonia) and, later, was professor from the Mesozoic of Africa, India, Argentina, Brazil, Uruguay, at the University of Basel, Switzerland (Rieth 1977; Reif & Lux and Australia illustratedhis great interest in the fauna of ancient 1987; Turner 2009). Heinrich received the nobility title ‘Freiherr’ Gondwana (see Huene 1927b, 1927c, 1929c, 1929d, 1930, 1931, (equivalent to Baron) due to his significant ecclesiastical work 1933; Mohabey 2011; Holz 2012) – although now many of the in the academy, as well as for coming from a German family of taxa described by him are no longer valid. von Huene and Matley high society. Consequently, Huene inherited his father’s noble (1933) published an extensive monograph on the records of title. According Reif and Lux (1987), as a theologian, Huene’s saurischian and ornithischian dinosaurs from the province of father influenced his education at an early age, which remained Chhota Simila, and recently Wilson et al. (2009) and Carrano with him throughout his life. et al. (2010) added new information on the dinosaurs and the The Huene family moved from Tübingen to Basel, where history of geosciences of the region. Friederich spent his childhood and made friends that he would His interests in biology and paleobiogeography are explicit meet years later in Santa Maria city (Rio Grande do Sul State, in some of his studies that are presently rarely cited (e.g. Huene Brazil) when they were working as Lutheran priests in the 1929a, 1933). Other paleontologists, especially mammal spe- up-country of Rio Grande do Sul State (Schultz & Holz 1996). cialists (e.g. Henry Fairfield Osborn in 1931; Figure 2), showed His studies at the University of Lausanne in Switzerland interest in the titanosaurid fauna Huene had described, and focused on Theology and Natural Sciences – areas of knowledge exchanged remarks and correspondences with him. His work that Huene was always enthusiastic about. At the age of 23, his corroborated the hypothesis that the fauna and flora of the academic interests became concentrated on geology and paleon- landmasses of the Southern Hemisphere were endemic, and tology at the University of Tübingen where he was supervised by had considerable taxonomic importance to the Gondwana the renowned geology professor Ernst Hermann Friedrich Koken supercontinent. (Gross 1969; Schultz & Holz 1996; Maisch 1999; Turner 2009). Of the countries of the Southern Hemisphere, Argentina was Huene was granted the title of Doctor in these fields of expertise. the one most visited by Huene since he had a close professional His studies initially focused on invertebrates and resulted in a relationship with the paleontologists Florentino Ameghino and monograph on Silurian brachiopods. He then enrolled in a PhD Rodolfo M. Casamiquela (Figure 1(a)). Huene’s description of program at the same university, and was invited by his supervi- several reptile specimens from the Cretaceous of North Patagonia sor to occupy the official position of Assistant in Paleontology is still of great scientific importance. In this work he described (Maisch 1999). During his PhD, von Huene started to study and named the titanosaurid sauropod species Antarctosaurus 1086 C. R. A. CANDEIRO AND S. F. M. FIGUEIRÔA

Figure 1. Stamps from Friedrich von Huene’s worldwide correspondence with important paleontologists from Argentina (A), Canada (B), China (C), France (D), Union of Soviet Socialist Republics (E/F), and India (H). (These documents are housed in Mathematisch-Naturwissenschaftliche Fakultät bietet Studierenden und/Universität Tübingen and here were kindly assigned to the present research.) wichmannianus (Huene 1929a) as a tribute to its discoverer, of Rio de Janeiro on 2 April 1924, on his way to South Africa Richard Wichmann, a German who had settled in Argentina. (see Huene 1944). Until that time, little was known about dinosaurs of Argentina. Around 12 June 1928, Huene returned to Brazil. His access to This species is still relevant to contemporary paleontology, and South American fossils was facilitated by his influential contacts was considered for many years one of the world’s largest dino- in various countries: Florentino Ameghino (1854–1911), who saurs. A. wichmannianus has proven to be an important species was the director of the Museo de Ciencias Naturales in Buenos for understanding titanosaurid evolution, and is still used in Aires, Argentina; and Hermann Friedricht Albrecht von Ihering comparisons with other sauropods and in phylogenetic analy- (1850–1830), who directed the Museu Paulista in São Paulo, ses (e.g. Upchurch 1999; Wilson 2002, 2005; Curry-Rogers 2009; Brazil. Huene’s visit was reported in the official record of the Wilson et al. 2009; Martínez et al. 2016). Huene’s monograph on Brazilian Academy of Sciences by its president, Eusébio Paulo de A. wichmannianus was a turning point in understanding dino- Oliveira. In this official communication, Huene was presented as saurs of the Southern Hemisphere, and it remains one of his most ‘the Prof. Frederico von Hulisco, eminent Hungarian paleontol- cited studies of herbivorous dinosaurs of Gondwana. ogist …’ (Acta Academia Brasileira de Ciências (ABDC) 1928, p. The reptiles from Northern Patagonia (mosasaurids, croc- 58; Figure 3), a presentation that was not only inaccurate in the odyliforms, and dinosaurs) were of great interest to Huene; spelling of his name, but also in his nationality. After his passage however, the description of those specimens were not based on through Rio de Janeiro, he traveled to the city of São Paulo, where specimens from Argentina, but from Uruguay (3 taxa) and Brazil he met with Ihering. (5 taxa; in São Paulo and Minas Gerais states), for a total of eight Huene’s stay in São Paulo allowed him to access material col- taxa. This work stemmed from his second visit to Brazil, since lected from the rocks of the Bauru Group in western São Paulo his first visit to the country was fleeting, passing through the city State, as well as materials from the Triângulo Mineiro region. HISTORICAL BIOLOGY 1087

Figure 2. Osborn letter to von Huene early 1930. (This document is housed in Mathematisch-Naturwissenschaftliche Fakultät bietet Studierenden und/Universität Tübingen and here was kindly assigned to the present research.)

These rocks had yielded fossilised teeth, long bones, and even region. At that time there were no paleontologists or natural- scutes from small-to-large turtles. Many specimens of reptiles ists in Brazil interested in describing Cretaceous vertebrates. were housed at the Museu do Ypiranga (see Huene 1933) and Hence, the paleontological knowledge of this young German were briefly described by him. However, the most detailed naturalist gave him an advantage in working on the descrip- description was that of what he formerly considered a carnivo- tion of these materials. The financial aid used by Huene to travel rous dinosaur, Brasileosaurus pachecoi, which he reassigned as to Brazil was granted by the Notgemeinschaft der Deutshchen a crocodyliform. Wissenshaft, the Bayerische Akademie der Wissenschaften, and Around 28 July 1928, Huene started to explore Brazil. He trav- the Paläontologische Institut in München. eled to the states of São Paulo, Paraná, Santa Catarina, and Rio Friedrich von Huene stayed in São Paulo working at the Museu Grande do Sul, and returned to Germany via Uruguay (Schultz Paulista, where he produced five descriptions and depictions of & Holz 1996). When in São Paulo, he described material housed reptiles from the Upper Cretaceous of western São Paulo State, at the Museu Paulista, and published studies about vertebrate some of which were published in German scientific journals. fossils from western São Paulo State and the Triângulo Mineiro In the early decades of the last century, Huene published over 1088 C. R. A. CANDEIRO AND S. F. M. FIGUEIRÔA

Figure 3. Official communication from Acta Academia Brasileira de Ciências announcing the presence of von Huene. (This document is housed in Academia Brasileira de Ciências and here was kindly assigned to the present research.) eight papers, among which were some remarkable studies on José do Rio Preto city (western São Paulo State), which were Cretaceous reptiles from Central Brazil. The familiarity of Huene dinosaurs. Ihering was also the first to taxonomically assign and with reptiles, especially from Argentina, allowed him to perform date the Cretaceous vertebrates from Central Brazil in his classic morphological comparisons among many reptile specimens from publications (Huene 1931, 1933). Thenceforth, the reptile fossils Central Brazil that were also described by him. from this region started to be intensely studied (see Staesche The German paleontologist Ihering (1911) briefly described 1937; Price 1945, 1950a, 1950b, 1951, 1953, 1955, 1961) until the first vertebrate fossils from the Bauru Group, found in São 1978 (Candeiro et al. 2006). From the 1990s until today, the HISTORICAL BIOLOGY 1089 reptiles from the Bauru Group have been the objects of many Testudines Batsch 1788 extensive studies. Chelonia indet. Original assignment: Chelonia. Huene’s Cretaceous vertebrates from Brazil Locality: São José do Rio Preto. Geological unit and age: Adamantina Formation, Turonian- Holz and Schultz (1998) published the most complete work Santonian (Dias-Brito et al. 2001). about von Huene’s stay in Brazil, where this paleontologist Repository: Museu Paulista (incorporated by the Instituto became interested in Triassic fossils from Rio Grande do Sul Geológico de São Paulo). State. The paper presents von Huene’s biography and describes Material: one scute. his prolific research as a vertebrate paleontologist. At the begin- Remarks: Huene (1933) makes a brief mention of a small turtle ning of the last century, the municipalities of Santa Maria and scute encrusted in a sandstone fragment. São Pedro do Sul were his main reasons for his visiting Brazil, Crocodylomorpha Walker 1970 since he was in search of fossil reptiles, which are still found Crocodyliformes Benton and Clark 1988 today in the outcrops of these regions. According to Kellner Crocodylomporpha indet. and Campos (1999), between the years of 1928 and 1929, von Brasileosaurus pachecoi Huene 1931 Huene performed many fossil collections, on which he pub- (Figure 4) lished during Second World War. This study (Huene 1942) is Locality: Guajuçara, São Paulo State. one of the most classic publications about Triassic reptiles from Geological unit and age: Adamantina Formation, Turonian- Rio Grande do Sul and it became the main reference for South Campanian (Dias-Brito et al. 2001). American Triassic reptiles. Material: left humeral diaphysis, a right femoral diaphysis, and The Cretaceous vertebrates from Central Brazil were a right quadrate. already known from descriptions of dinosaurian remains from Repository: Comissão Geographica e Geológica de São Paulo south-central Mato Grosso State. These fossil reptiles were dis- and Museu Paulista (incorporated by the Instituto Geológico de covered during the first prospecting trips for mineral resources São Paulo), and Serviço Geológico do Brasil (incorporated by the in Brazil in 1884 by the Comissão Geológica do Império do Departamento Nacional da Produção Mineral). Brasil. The North American naturalist Herbert Huntington Smith Remarks: During his stay at the Museu do Ypiranga, in São mentioned the remains of Cretaceous vertebrates from western Paulo, Huene described this material, which was characterised São Paulo State (Woodward 1910). Later, the geologist Orville as the first reported specimen of a Cretaceous reptile from Adelbert Derby (1851/Kellogsville, NY – 1915/Rio de Janeiro, RJ) the state of São Paulo. This taxon was named Brasileosaurus wrote an academic report about those specimens, describing ‘a pachecoi, and was the first dinosaur species described from the fragment of a turtle’s shell and a reptile vertebra’. Cretaceous of Brazil. Later, Huene (1933) assigned this spe- At the beginning of the last century, western São Paulo State cies to Crocodyliformes. However, some authors (e.g. Carvalho was considered one of the most important regions for the dis- et al. 2004, 2011; Candeiro & Martinelli 2006) consider the covery of Cretaceous fossils in South America. At that time rail- species to be a nomem dubium or a problematic taxon, since ways were being constructed in the region and it was usual to the specimens have no diagnostic features supporting a dis- find fossil remains during the drilling of water wells. These first tinct species. specimens were sent to the Serviço Geológico e Mineralógico Crocodylomporpha indet. do Brasil and Museu Paulista. The visit of von Huene to Brazil (Figure 3) in 1928 made it possible for him to access these materials and to Locality: Collina/Barretos, São Paulo State. perform the first detailed descriptions of fossils from the Bauru Geological unit and age: Adamantina Formation, Turonian- Group. Santonian (Dias-Brito et al. 2001). In the following section we present a summary of the fos- Material: a rib and teeth. sils Huene studied from the Bauru Group of western São Paulo Repository: Comissão Geographica e Geológica de São Paulo State, as well as his preliminary assignments. These are known as and Museu Paulista (incorporated by the Instituto Geológico de the first formal records of vertebrate fossils from the Triângulo São Paulo), and Serviço Geológico do Brasil (incorporated by the Mineiro region. Departamento Nacional da Produção Mineral). Remarks: Crocodyliform remains were the most commonly Systematic paleontology described materials in Huene’s works from 1931 to 1933. The teeth from Collina were originally assigned to Crocodilia (Huene The specimens ascribed to reptiles by Huene (1931, 1934) were 1931, p. 189). They are characterised by being 23 mm in diameter collected in the municipalities of Barretos, Presidente Prudente, in a circular transverse section, and for being highly curved. The and São José do Rio Preto, located in western São Paulo State, external surfaces are smooth, with large longitudinal grooves, and in the municipality of Monte Alegre de Minas, Minas Gerais typical of the Crocodyliformes from the Bauru Group. The teeth State. In the original studies (Huene 1931, 1933), the specimens are 8 mm in height, and roots are absent. were described as belonging to a variety of reptiles (crocodyli- There are two other fragmentary teeth in this collection forms to sauropods and theropod dinosaurs). There are no other with only the distal portions preserved. They are 17 mm long descriptions of this material, and the little information available and 13 mm in height, with the apex showing fine longitudinal is in the studies made by Huene. A revision is needed for these grooves, and the borders of the carinae have asymmetric den- taxa, but this material appears to be missing at the present time. ticles; the latter, according to Huene (1925), are similar to the 1090 C. R. A. CANDEIRO AND S. F. M. FIGUEIRÔA

Figure 4. Brasileosaurus pachecoi nomem dubium remains from western São Paulo State; (A–D) femur, (E–K) humerus (modified from Huene 1931; not in scale). denticles of the thalattosuchian Machimosaurus. Nevertheless, first formal records of dinosaur remains from these regions (e.g. these characteristics are widely observed in many mesoeucroc- Candeiro 2011; Peyerl et al. 2015). odylian groups of the Bauru Group (e.g. Baurusuchidae). Theropoda Marsh 1881 Numerous bones and isolated teeth from the Guarucaia Theropoda indet. region (near the Paranapanema river, currently the municipal- (Figure 5) ity of Presidente Prudente) were also housed at the Comissão Original assignment: Carnosauria (Thecodontosaurus-like). Geográfica e Geológica de São Paulo. However, according to Locality: Collina (Barretos), São José do Rio Preto, São Paulo Huene, the material was not available to be examined and, there- State. fore, it was not possible to describe it. Geological unit and age: Adamantina Formation, Turonian- Dinosauria Owen 1842 Santonian (Dias-Brito et al. 2001). Saurischia Seely 1888 Repository: Comissão Geographica e Geológica de São Paulo Sauropoda Marsh 1878 (the current repository is unknown). Titanosauria Bonaparte and Coria 1993 Material: teeth. Titanosauria indet. Remarks: Huene (1931, p. 189) originally assigned the material Locality: Collina (São Paulo) and Monte Alegre de Minas (Minas to Carnosauria. The teeth vary in size with crown heights of Gerais). 4–19 cm, and the crowns are broken at the bases. They are lat- Geological unit and age: Adamantina Formation, Turonian- erally compressed, highly curved on the posterior carinae, and Campanian (Dias-Brito et al. 2001). have finely serrated carinae with around five denticles per 2 mm. Repository: Comissão Geographica e Geológica de São Paulo Huene (1931) described three teeth from the region of São and Museu Paulista (incorporated by the Instituto Geológico de José do Rio Preto that were part of the collection of the Museu São Paulo), and Serviço Geológico do Brasil (incorporated by the Paulista. According to him (op. cit.), these specimens were sent Departamento Nacional da Produção Mineral). as a donation to the museum in May, 1909, and were received Material: femur, humerus, and isolated vertebrae. by Hermann von Ihering. The material was discovered during Remarks: Little is known about the material from the original an excavation in the rocks of the Adamantina Formation. One description of these specimens. The importance of these fos- of these teeth can be considered the first fossil discovered of the sils has been mentioned in many papers, as they represent the Bauru Group, and was briefly mentioned by Woodward (1910), HISTORICAL BIOLOGY 1091

Figure 5. Theropod dinosaur crown remains from western São Paulo State (modified from Huene 1931; (A), (C), (D), (F), (H), (J), (M) crowns, (B), (E) (G), (I), (K), (L) basal cross-section, n possible anterior denticles; not scaled).

who described it as ‘a dinosaur tooth from the Triassic of São Mineiro region. From these descriptions, five taxa were iden- Paulo’. The specimen is 15 m in length, laterally compressed, and tified. The described specimens were later used to endorse the apex and both carinae are preserved and show little curva- his paleogeographic observations of the Cretaceous areas of ture, which differs from Thecodontosaurus (Woodward 1910). Gondwana. During the first two decades of the last century, the rocks of The study of reptiles from the Upper Cretaceous of São Paulo the Bauru Group were considered by some authors (e.g. Huene State and of the Triângulo Mineiro region is a relatively short 1933) to be of Jurassic age. It was only by 1925, when the rocks chapter in the story of research from the 1920s to 1930s. During of the Bauru Group were fully explored, that this geologic unit these decades, paleontologists in these regions restricted their was referred to the Cretaceous, especially on the basis of dino- studies to the collections made by the naturalists of the Serviço saur remains. Many studies consider the first formal report of Mineralógico do Brasil - collections made mainly by Euzébio de vertebrates from the Bauru Group to have been made by Ihering Oliveira. Thus, it is clear that Huene’s descriptions of specimens (1911); however, the first description was made two years prior from Central Brazil were made possible thanks to his easy access by Woodward (1910). to the collections of the SMB and the Museu Paulista, given by its One tooth from this region, possibly from the municipality of respective directors. Thanks to his efforts, the number of formal Presidente Prudente, was assigned to Carnosauria. The specimen reptile taxa identified from the Upper Cretaceous went from one is 20 mm in height and the base of the crown is 17 mm wide to five (Huene 1931, 1933). anteroposteriorly. The posterior carina is curved; it is laterally Huene’s studies on reptiles from western São Paulo State and compressed but the denticles are difficult to observe. from the Triângulo Mineiro region are representative when com- pared to other groups (e.g. invertebrates and fish), thanks to the Discussion and conclusions size of specimens and the abundance of reference material. Since the beginning of the last century, these facts caught the attention Considering what it is known about the life of Friedrich von of collectors, and intrigued researchers, such as Huene. These Huene, and what can be identified from his work, he was clearly fossils were usually found in uninhabited areas where railway a paleontologist who made a great contribution to the scien- constructions were being made, and in extremely hard rocks tific knowledge of the Brazilian Cretaceous. He was a classic that were difficult to prepare. After the 1930s, other paleontolo- academic researcher, and his primarily interest in Brazil was gists (e.g. Llewelyn Ivor Price) conducted new field studies and studying reptiles from the Triassic of Rio Grande do Sul. systematic descriptions of the fossils from these regions. Over However, during his first stay in the city of Rio de Janeiro, he the last 25 years, these areas have become two of the most impor- was received at the Brazilian Academy of Sciences and formally tant regions bearing fossil records from the Cretaceous of the introduced to the scientific community by Euzébio de Oliveira. Southern Hemisphere. Besides consolidating his professional relationship with Euzébio Retrospectively, the substantial research conducted by Huene de Oliveira (see also Huene 1939), Huene’s stay in São Paulo on fossil specimens from Central Brazil represents a remarkable allowed him to study the fossil specimens shared with him by work, especially considering that in this country Cretaceous rep- his friend, Hermann Ihering. The significant funding provided tiles were not the main focus of his career. Huene’s main research by the Munich Academy of Sciences, allowed Huene to describe was on material from the Triassic of Rio Grande do Sul State, several turtle, crocodyliform, and dinosaur specimens from the and from the Cretaceous of Argentina. Yet he described five Cretaceous of western São Paulo State, and of the Triângulo Cretaceous taxa of the Bauru Group from four localities in two 1092 C. R. A. CANDEIRO AND S. F. M. FIGUEIRÔA published papers. His achievements were greatly facilitated by Carvalho IS, Ribeiro LCB, Avilla LS. 2004. Uberabasuchus terrificus sp. nov., his professional networks with colleagues in the cities of Rio de a new crocodylomorpha from the Bauru Basin (Upper Cretaceous), Janeiro and São Paulo, as well as with other important contacts Brazil. Gondwana Res. 7:975–1002. Carvalho IS, Teixeira VPA, Ferraz MLF, Ribeiro LCB, Martinelli AG, in South America. To this day, Frederich von Huene remains an Macedo Neto F, Sertich JJW, Cunha GC, Cunha IC, Ferraz PF. influential figure among researchers who study reptiles of the 2011. Campinasuchus dinizi gen. et sp. nov., a new Late Cretaceous Cretaceous of Brazil, and he continues to be cited in important baurusuchid (Crocodyliformes) from the Bauru Basin, Brazil. Zootaxa. studies in the fields of paleontology and geology. 2871:19–42. Curry-Rogers K. 2009. The postcranial osteology ofRapetosaurus krausei (Sauropoda: Titanosauria) from the Late Cretaceous of Madagascar. Acknowledgments J Vertebr Paleontol. 29:1046–1086. Dias-Brito D, Musacchio EA, Castro JC, Maranhão MSAS, Suazez This work is a major part of Postoctoral research presented to the JM, Rodrigues R. 2001. Grupo Bauru: uma unidade continental Programa de Pós-Graduação em Política Científica e Tecnológica, Instituto do Cretáceo no Brasil - concepções baseadas em dados de Geociências, Universidade Estadual de Campinas. The authors were micropaleontológicos, isotópicos e estratigráficos [Grupo Bauru: uma given access by Dr. Rainer Schoch (Staatliche Museum für Naturkunde unidade continental do Cretáceo no Brasil – concepções baseadas em Stuttgart/Germany), Mathematisch-Naturwissenschaftliche Fakultät bie- dados micropaleontológicos, isotópicos e estratigráficos]. Revue de tet Studierenden und/Universität Tübingen (Staff) and Dr. Rainer Radtke Paléobiologie. 20:245–304. Portuguese. (Brazilien-Zentrum, Universität Tübingen/Germany), who also provided Gross W. 1969. Friedrich Freiherr von Huene. Palaeontologische assistance in studying the bibliographical collections of their institutions. Zeitschrift. 43:111–112. We also thank Dr. Drielli Peyrell (University of Campinas) for supply- Holz M. 2012. Estratigrafia de Seqüências - Histórico, princípios e aplicações ing references to the senior author during the development of this the- [Sequence Stratigraphy – History, principles and applications]. 1st ed. sis. R. Candeiro and S. Figueirôa were supported by Conselho Nacional Rio de Janeiro: Interciencia, 1; 258 p. Portuguese. de Desenvolvimento Científico e Tecnológico (CNPq)/ Bolsista de Holz M, Schultz CL. 1998. Taphonomy of the south Brazilian Triassic Produtividade em Pesquisa fellow. herpetofauna: fossilization mode and implications for morphological studies. Lethaia. 31:335–345. Huene F. 1902. Übersicht über die Reptilien der Trias [Overview of Disclosure statement the reptiles of the Triassic]. Geologische und Paläontologische Abhandlungen. 6:1–84. German. No potential conflict of interest was reported by the authors. Huene F. 1908. Die Dinosaurier der Europäischen Triasformation mit berücksichtigung der Ausseuropäischen vorkommnisse [Dinosaurs of References the European Triassic formation, with consideration of the European approach]. Geologische und Palaeontologische Abhandlungen Suppl. ABDC. 1928. Atas das Sessões Ordinárias 1926–1929 [Actas of Ordinay 1:1–419. German. Session 1926–1919]. Academia Brasileira de Ciências (Brazil). p. 58. Huene F. 1925. Ichthyosaurier aus der Kreide Argentiniens [Ichtiosaurs Alberti V. 1996. A existência na história: revelações e riscos da hermenêutica from Argentinean Cretaceous]. Revista del Museu de La Plata. 28:234– [The existence in history: revelations and risks of hermeneutics]. 242. German. Estudos Históricos/Historiografia. 9:31–57. Portuguese. Huene F. 1927a. Beitrag zur Kenntnis mariner mesozoischer Wirbeltiere Azevedo RP, Simbras FM, Furtado MR, Candeiro CRA, Bergqvist LP. in Argentinien [Contribution to the knowledge of marine mesozoic 2013. First Brazilian carcharodontosaurid and other new theropod vertebrates in Argentina]. Centralblatt für Mineralogie, Geologie und dinosaur fossils from the Campanian Maastrichtian Presidente Paläontologie B:22–29. German. Prudente formation, São Paulo state, southeastern Brazil. Cretaceous Huene F. 1927b. Gondwana-Reptilien in Südamerika [Gondwanan Res. 40:131–142. repitilian in South America]. Palaeonlogica Hungarica II. 1:1–108. Candeiro CRA. 2011. Notes concerning Friedrich von Huene and his German. studies on Late Cretaceous fossils in the states of Minas Gerais and Huene F. 1927c. Contribucion a la Palaeogeografia de Su-America São Paulo. In: Anais do V Simpósio Nacional de Ensino e História de [Contribution to the Paleogeography of South America]. Boletin de la Ciências da Terra; Nova Friburgo, 1; p. 260–264. Academia Nacional de Ciencias. 30:231–292. Spanish. Candeiro CRA, Figueirôa SFM. 2012. El legado del paleontólogo alemán Huene F. 1928a. Aphorismen über die Stratigraphie des brasilianischen Friedrich von Huene con el estudio de los reptiles fósiles del Grupo Staates São Paulo [Aphorisms about the stratigraphy of the Brazilian Bauru (estados de Minas Gerais y São Paulo, Brasil) en los años 1920– state São Paulo]. Separat-Abdruck aus dem Centralblatt für Mineralogie, 1930 [The legacy of the German paleontologist Friedrich von Huene Geologie und Paläontologie. B10:524–531. German. with the study of the fossil reptiles of the Bauru Group (states of Minas Huene F. 1928b. Überblick über die Schichtenfolge der jügeren Kreide Gerais and São Paulo, Brazil) in the years 1920–1930]. In: Boletin de und ihre Verbreitung in Patagonien [Overview of the layer sequence Resumen XXVI Jornadas Argentinas de Paleontologia de Vertebrados; of the younger chalk and its spread in Patagonia]. Centralblatt für Buenos Aires, 1; 5–5. Spanish. Mineralogie, Geologie und Paläontologie. B1:103–110. German. Candeiro CRA, Figueirôa SFM. 2014. Friderich von Huene’s research on Huene F. 1929a. Los Saurisquios y Ornitisquios del Cretaceo Argentino Late Cretaceous Central Brazil early last century. In: Gondwana 15, [The saurischian and ornithischians of the Argentine Cretaceous]. 2014. Madrid: Abstract Book Gondwana 15, 2014. 1; p. 27–27. Annales del Museo de La Plata Serie II, T, III:1–196. Spanish. Candeiro CRA, Martinelli AG. 2006. A review of paleogeographical and Huene F. 1929b. Terrestriche Oberkreide in Uruguay [Terrestrial chalk in chronostratigraphical distribution of mesoeucrocodylian species from Uruguay]. Centralblatt für Mineralogie, Geologie und Paläontologie. the upper Cretaceous beds from the Bauru (Brazil) and Neuquén B1:107–112. German. (Argentina) groups, southern South America. J South Am Earth Sci. Huene F. 1929c. Versuch einer Skizze der paläogeographischen Beziehun- 22:116–129. gen Südamerikas [An attempt at a sketch of the palaeogeographical rela- Candeiro CRA, Martinelli AG, Avilla LS, Rich T. 2006. Tetrapods from the tions of South America]. Geologischen Rundschau. 20:81–96. German. Upper Cretaceous (Turonian?Maastrichtian) Bauru Group of Brazil: a Huene F. 1929d. Versuch einer Skizze der paläogeographischen Beziehungen reappraisal. Cretaceous Res. 27:923–946. Südamerikas [An attempt at a sketch of the palaeogeographical relations Carrano MT, Wilson JA, Barrett PM. 2010. The history of dinosaur of South America]. Geologischen Rundschau. 20:81–96. German. collecting in central India, 1828–1947. In: Moody RTJ, Buffetaut E, Huene F. 1930. Terrestrische Oberkreide im Centralen Südamerika Naish D, Martill DM, editors. Dinosaurs and other extinct saurians: a [Terrestrische Oberkreide im Centralen]. Centralblatt für Mineralogie, historical perspective. Geological Society; p. 161–173. Geologie und Paläontologie. B1:177–181. German. HISTORICAL BIOLOGY 1093

Huene FV. 1931. Verschiedene mesozoische Wirbeltierreste aus Sudamerika Price L. 1953. Quelônios da Formação Bauru, Cretáceo Terrestre do Brasil [Various mesozoic vertebrate remains from South America]. Neues Meridional [Quelonian of the Bauru Formation, Cretaceous terrestrial Jahrbuch fur Geologie und Palaontologie, Monastshefte. 66(B):181– of Southern Brazil]. Boletim da Divisão de Geologia e Mineralogia. 198. German. 147:1–39. Portuguese. Huene F. 1933. Die südamerikanische Gondwanafauna [The South Price L. 1955. Novos crocodilideos dos Arenitos da Série Bauru, Cretáceo American Gondwanafauna]. Forsh. U. Fortschr. 9:129–130. German. do estado de Minas Gerais [New crocodilides of the Bauru, Cretaceous Huene F. 1934. Neue Saurier-Zähne aus der Kreide von Uruguay [New series sandstones of the state of Minas Gerais]. An Acad Bras Cienc. dinosaur teeth from the chalk of Uruguay]. Centralblatt für Mineralogie, 27:487–498. Portuguese. Geologie und Paläontologie. B1:179–182. German. Price L. 1961. Sobre os dinossáurios do Brasil [About Brazilian dinosaurs]. Huene F. 1939. Carta de F von Huene ao bio de Oliveira [von Huene An Acad Bras Cienc. 33:3–4. Portuguese. letter to Dr. Euzébio de Oliveira]. Mineracão e Metalurgia. 4:190. Reif WE. 1984. Paleoecology and evolution in the work of Friedrich von Portuguese. Huene. In: Reif WE, Westphal F, editors. Third symposium on Mesozoic Huene F. 1942. Ergebnisse der Sauriergrabungen in Südbrasilien 1928/29 terrestrial ecosystems, short papers. Tübingen: Attempto; p. 193–197. [Results of Saurian excavations in South Brazil 1928/29]. Die fossilen Reif WE, Lux W. 1987. Evolutionstheorie und reliöses Konzept im Werk Reptilien des südamerikanischen Gondwanalandes. München: C. H. der Wirbltierpalaeontogen Friedrich Freiherr von Huene (1875–1969) Beck; 332 p. German. [Evolution theory and a religious concept in the work of the vertebral Huene F. 1944. Arbeitserinnerungen – Selbstbiographien von palaeontologists Friedrich Freiherr von Huene (1875–1969)]. Bausteine Naturforschern [Work memoirs - biographies of naturalists]. Nummer zur Tübinger Universitätsgeschichte, Folge. 3:91–140. German. 2. Halle (Saale): Leopoldina, 52 p. Rieth GA. 1977. Der Mann, der unter die Drachen ging [The man who Huene F. 1956. Grundsätzliches zur Stammesgeschichte der Tetrapoden went under the kites. Memories of The Tübinger paleontologist, [Basics of the taxa history of the tetrapods]. Paläont Z. 30:26–29. Friedrich Freiherr von Hoyningen]. Erinnerungen an den Tu¨binger German. Paläontologen Friedrich Freiherr von Hoyningen gen. Huene. Tübinger Huene F, Matley GA. 1933. The cretaceous Saurischia of the central Blätter. 64:35–36. German. provinces of India. Memoirs Geol Surv India. 21:1–74. Schultz CL, Holz M. 1996. Friedrich Von Huene - Um Paleontólogo Entre Ihering H. 1911. Fósseis de São José de Rio Preto [Fossilis from São José do A Ciência e A Fé [A paleontologist between Science and faith]. Ciências Rio Preto]. Revista do Museu Paulista. 8:141–146. Portuguese. Biológicas e do Ambiente/Santa Maria. 13:105–118. Kellner AWA, Campos DA. 1999. Vertebrate paleontology in Brazil – a Seebohm TM. 2004. Hermeneutics. Method and methodology. Springer; review. Episodes. 22:238–251. 276 p. Maisch MW. 1999. Friedrich von Huene (1875-1969) – Der Tübinger – Staesche K. 1937. Podocnemis brasiliensis n. sp. aus der oberen Kreide Saurierjäger. Trias eine ganz andere welt. 7:606–610. Brasiliens [Podocnemis brasiliensis n. sp. from the Cretaceous of Brazil]. Martínez RDF, Lamanna MC, Novas FE, Ridgely YRC, Casal GA, Martínez Neues Jahrbuch für Geologie und Paläontologie. 77:291–309. German. JE. 2016. A basal lithostrotian titanosaur (Dinosauria: Sauropoda) with Sues HD. 1997. European dinosaur hunters: Friedrich von Huene. a complete skull: implications for the evolution and paleobiology of In: Farlow JO, Brett-Surman MK, editors. The complete dinosaur. titanosauria. PLoS ONE. 11:e0151661. Bloomington: Indiana University Press; p. 17–18. Mohabey DM. 2011. History of Late Cretaceous dinosaur finds in India Turner S. 2009. Reverent and exemplary: ‘dinosaur man’ Friedrich von and current status of their study. Journal Palaeontological Soc India. Huene (1875–1969). In: Kölbl-Ebert M, editor. Geology and religion: 56:127–135. A history of harmony and hostility. The geological society. London: Peyerl D, Candeiro CRA, Figueirôa SFM. 2015. Trajectory and contribution Special Publications, 310; p. 223–243. of geoscientists (1906–1961) to dinosaur research in the Bauru Group Turner S. 2011. Dinosaurs and lost dreams: the von Huene–Longman (Cretaceous) in the Triângulo Mineiro region of Brazil. J South Am story. Hogg Dinosaurs, their Kith and Kin: a historical perspective. 32. Earth Sci. 61:64–70. Upchurch P. 1999. The phylogenetic relationships of the Nemegtosauridae Price L. 1945. A new reptile from the Late Cretaceous of Brazil. Serviço (Saurischia, Sauropoda). J Vertebr Paleontol. 19:106–125. Geológico Mineralogia. 25:1–8. Wilson JA, D’Emic MD, Curry-Rogers KA, Mohabey DM, Sem S. 2009. Price L. 1950a. On a new crocodilian, Sphagesaurus, from the Cretaceous Reassessment of the sauropod dinosaur Jainosaurus (= ‘Antarctosaurus’) of the state of São Paulo, Brazil. An Acad Bras Cienc. 22:77–85. septentrionalis from the Upper Cretaceous of India. Contributions from Price L. 1950b. Os crocodilideos da fauna da formacão Baurú, do Cretáceo the Museum of Paleontology of the Univeristy of Michigan. 32:17–40. terrestre do Brasil Meridional [The crocodilians of the Bauru Formation Wilson JA. 2002. Sauropod dinosaur phylogeny: critique and cladistic fauna, from the terrestrial Cretaceous of Southern Brazil]. An Acad Bras analysis. Zool J Linn Soc-London. 136:217–276. Cienc. 22:473–490. Portuguese. Wilson JA. 2005. Redescription of the mongolian sauropod nemegtosaurus Price L. 1951. Um ovo de dinossaurio na formação Baurú, do Cretácico do mongoliensis nowinski (Dinosauria: Saurischia) and comments on late estado de Minas Gerais [A dinosaur egg in the Bauru Formation, from cretaceous sauropod diversity. J Syst Palaeontol. 3:283–318. the Cretaceous of the state of Minas Gerais]. Notas Preliminar Divisão Woodward AS. 1910. On a tooth of Triassic dinosaur from S. Paulo, Brazil. de Geologia e Mineralogia. 53:1–7. Portuguese. Rep Assoc Adv Sci. 79:483.