Flávio Ferreira Pinto de Resende (1907–1967): A portuguese scientist and hero‐botanist, and his forays into systematics

Abstract

Flávio Ferreira Pinto de Resende (Fig. 1) was born on 28 February 1907 in the small village of Cinfães in northern Portugal, and died in Lisbon on 1 January 1967, shortly before his 60th birthday. In 1924, after finishing high school in Porto, he enrolled at the University of Porto to study medicine but later switched to natural sciences, being conferred his first degree in 1928. From 1928 to 1930 he attended the Escola Normal Superior, in Coimbra, and established a friendship with his contemporary Abílio Fernandes (1906–1994); the two were born only one year apart. Fernandes was then working for his doctorate under the supervision of Aurélio Quintanilha (1906–1987). Resende had the chance to visit Quintanilha's laboratory and observe its intense activity (Quintanilha, 1967–1968, 1975). As Resende needed a job, he accepted an appointment as a high school teacher in Guarda but soon realised that he desired further training in science. He wrote directly to the Minister of Education, whom he did not even know personally, explaining his problem and requesting a scholarship to support postgraduate studies at the University of Coimbra (Quintanilha, 1975). This unusual request was granted and he obtained a scholarship to continue with his studies at Coimbra under the supervision of Quintanilha, from 1932 to 1933 (Fernandes, 1967). It was at this time that he initiated his research in cytogenetics in Quintanilha's laboratory, having Fernandes as one of his colleagues. Both Resende and Fernandes became accomplished cytogeneticists (karyology and cytotaxonomy). However, while Resende had more eclectic research interests, and his subjects were mostly succulent plants, especially the southern African rosulate, succulent-leaved alooids and Madagascan Crassulaceae, Fernandes, who was eventually to become the director of the Instituto Botânico at the University of Coimbra from 1942–1974, focussed on Narcissus L., the well-known daffodils of the flori- and horticultural trades (Fernandes, 1968; Figueiredo & al., 2018). In 1933 Quintanilha secured a German scholarship from the Alexander von Humboldt-Stiftung for a member of his staff. He appointed Resende as the recipient and instructed him to attend the Institute of Plant Physiology, Dahlem, where Quintanilha himself had done postgraduate work. Resende went to Berlin at the end of 1933, and after studying German and making inquiries on the state of the art in his chosen field of research, he eventually moved to Hamburg to work at the Botanical Institute with Emil Heitz (1892–1965) (Linskens, 1967b; Diogo & al., 2015). Heitz, who was then at the summit of his career, would be dismissed in 1937 after his Jewish ancestry was pointed out to his superiors by a colleague, the botanist Edgar Irmscher (1887–1968) (Zacharias, 1995). Ten years later, in 1947, Resende would also be (albeit briefly) dismissed for political reasons. Resende who, for obvious reasons, was the only person Heitz could confide in at the Institute in Hamburg where both were working from 1934 to 1937 (Linskens, 1967b), remained friends with Heitz. One of the volumes of the periodical Portugaliae Acta Biologica that Resende later founded was dedicated to Heitz (Resende, 1962). In 1937, the same year that Heitz was dismissed, Resende was awarded his doctoral degree by the University of Hamburg (Linskens, 1967b; Diogo & al., 2015), and he published some of the results from his thesis in the journal Planta (Resende, 1937). In a footnote to this paper, Resende stated “ 1 Dissertation der Mathematisch-Naturwissenschaftlichen Fakultät der Hansischen Universität zu Hamburg. Der Aufenthalt des Verfassers in Hamburg und damit die Ausführung der Arbeit unter der Leitung von Prof. Dr. Heitz wurde durch das Instituto para a Alta Cultura, Lisboa, und zum Teil durch die Alexander von Humboldt-Stiftung, Berlin, ermöglicht, wofür beiden Stiftungen der Dank ausgesprochen sei.” [“Dissertation of the Faculty of Mathematics and Natural Sciences of the Hansische Universität zu Hamburg. The residence of the author in Hamburg and thus the execution of the work under the direction of Prof. Dr. Heitz was made possible by the Instituto para Alta Cultura, Lisboa, and partly by the Alexander von Humboldt Foundation, Berlin, for which I express gratitude.”] After obtaining his doctorate, Resende returned to Portugal but continued to spend research periods in Germany, first at Hamburg (1938–1939), then, from 1938 to 1941, i.e. during WWII, at the Kaiser Wilhelm Institut für Biologie in Berlin (Quintanilha 1967; Catarino & Simões, 2011). Upon his return to Portugal, Resende joined the University of Porto in northwestern Portugal, where he obtained the equivalence for his German doctoral degree and became a lecturer. In Porto the conditions were not well suited to his research interests so when a position opened for the chair of botany at the University of Lisbon he applied and was selected. From 1943 to 1967 Resende was a professor at the University of Lisbon and also director of the University's Botanical Garden (Catarino, 2001; Catarino & Simões, 2011; Smith & Figueiredo, 2016). In 1910 changes were introduced in higher education in Portugal that inter alia saw the establishment of the University of Lisbon, which had a Faculty of Science into which the Escola Politécnica de Lisboa was absorbed. The Politécnica dated from 1837 and its Herbarium (LISU) from 1839 (Figueiredo & al., 2018). In 1943 Flávio Resende became only the third director of the Instituto Botânico de Lisboa, an institution of the University of Lisbon. The Instituto also had a Botanical Garden, the Jardim Botânico de Lisboa, which was created in 1878 (Smith & Figueiredo, 2016). During Resende's directorship of the Instituto in Lisbon, including its Botanical Garden, the institution suffered from infrastructural and staffing difficulties (Resende, 1959b; Tavares, 1967–1968). Having conducted research at the University of Hamburg, one of the top European research institutions, which also conferred his doctorate, Resende had been exposed to a highly efficient and productive scientific environment in his early career in the mid-1930s. He was therefore able to fashion a clear vision of how the University of Lisbon should change to stay abreast of scientific progress and strongly advocated his opinions about the status quo, which he found unacceptable (Resende, 1945). This created discomfort for most of his colleagues at the University, as they were content with merely mastering and transmitting existing knowledge to their students and did not aspire to make any scientific discoveries (Resende, 1945; Linskens, 1967b). Together with many other professors and lecturers of the University of Lisbon, Resende was dismissed in 1947, allegedly for having contributed to political unrest (Catarino & Simões, 2011). He was reinstated shortly afterwards, as a result of a successful appeal to the then prime minister António de Oliveira Salazar (1889–1970) (Linskens, 1967b). After Resende was dismissed he received invitations to work abroad but as he was soon reinstated he did not accept any of the invitations from foreign shores (Simões, 2011). Had he accepted one of those offers and moved to another country his life and career would likely have been quite different, as he was well aware. Resende himself admitted that he did not rejoice in his readmission to the University of Lisbon (Simões, 2011). Under his directorship the Instituto at the University of Lisbon flourished, several of his students developed research careers and he established mutually beneficial working collaborations with many of his international connections (Quintanilha, 1975). Scientists from abroad were frequently invited to the University of Lisbon by Resende, who provided lodging for them in an apartment at the Botanical Garden (Linskens, 1967b). Even though Resende was essentially a cytogeneticist and physiologist, his scientific interests had a broad range and he was also active in plant taxonomy, describing new vascular plant taxa, an activity that dwindled at the Instituto in Lisbon in the mid-1960s after Resende died. During the mid-20th century Resende was a well-known student of the alooids, conducting his pioneering cytological research on these plants and their relatives, especially haworthias (see for example Resende, 1943) in Lisbon. According to Quintanilha (1975), Resende started out by working on matters pertaining to pure karyology, such as number and shape of chromosomes and their satellites in relation to the nucleoli, using techniques that he had mastered in Coimbra. For these investigations he preferred to use representatives of the Asphodelaceae subfam. Alooideae, which led him afterwards to develop systematic and phylogenetic studies in this group of plants: “Começou por trabalhar em assuntos de pura cariologia, número e forma dos cromossomas e seus satélites, nas suas relações com os nucléolos, cujas técnicas tinha aprendido em Coimbra. Para estes estudos serviu-se de preferência de Aloíneas, o que o levou depois a estudos de sistemática e de filogenia neste grupo de plantas.” The reason for Resende's research interest in the alooids was on one hand because the group is acknowledged as being in a process of active speciation, inter alia because of in situ hybridisation (Resende & Viveiros, 1948a,b; Fonseca, 2015) and, more pragmatically, because when he took up a position at the University of Lisbon, he encountered dilapidated greenhouses and a lack of adequate research facilities that only allowed the use as study subjects those plants with minimal growing requirements, i.e., “desert plants” (Resende, 1959b). Although very few alooids would qualify as true desert plants, many species will survive, if not thrive, while subject to considerable neglect in cultivation. During an 18-year period (1938–1956) Resende published ca. 50 new plant names and new combinations, not always validly. Most of these were published for taxa in the Asphodelaceae. As far as we could determine, Resende did not describe any new plant taxa from Portugal, with all of what he regarded as novel taxa originating from southern and south-tropical Africa and Madagascar. Given his background as a cytogeneticist, Resende relied heavily on giving recognition to chromosomal aberrations and minor morphological variations, mostly at infraspecific ranks. He paid special attention to two species of Haworthiopsis G.D.Rowley, H. tessellata (Haw.) G.D.Rowley and H. limifolia (Marloth) G.D.Rowley, and in both published numerous new varieties and forms. Both species are known to be remarkably variable, and today are interpreted as polymorphic entities spread over very wide distribution ranges. The only two entities that Resende described that are upheld today, albeit in a different genus [Haworthia Duval], are the nothospecies H. ×lisbonensis (Resende) Gildenh. & Klopper and H. ×sampaiana (Resende) Gildenh. & Klopper. By the early-1960s Resende's research output in systematics was somewhat diminished, and with co-authors, he published only a few additional papers in this area of botanical endeavour, mostly on cytogenetics in general. By then his interest had shifted to genetic factors in plant development (Lima-de-Faria, 1967). In 1959, as far as we were able to ascertain, Resende published a final systematics paper, on the cytotaxonomy of Haworthia. His interest in succulents, however, remained throughout his career, and (always the scientist) any new observation would stimulate his mind for further research and collaborations. For example, after observing different colour pigmentations in a hybrid aloe growing in the greenhouse of the Botanical Garden of Lisbon, which was attended by its head-gardener, A. Gomes Amaral, he initiated a study of the inheritance of floral pigments in Aloe. This resulted in a paper where, in a typical manifestation of his correctness in acknowledging the input of others, Resende included Amaral as co-author (Resende & Amaral, 1956). Although he travelled widely, in Europe, North America and South America (as a visiting professor in Brazil in 1955), as far as we could ascertain, it was only briefly that Resende visited South Africa, the country where the Asphodelaceae subfam. Alooideae that he had a strong research interest in is by far the most diverse. We are aware though that Resende visited Luanda (Angola) and Maputo (Mozambique), then Portuguese colonies in south-tropical Africa in 1960 as a professor of the Cursos de Férias no Ultramar (overseas summer courses) (Tavares, 1967–1968; Linskens, 1967b). During the visit to Angola he also went to the Museum of Dundo, in Lunda (Fig. 2) which was under the directorship of his friend António Barros Machado (1912–2002) (Machado, 1960). He returned to Mozambique in 1961 and then also visited Rhodesia (Zambia/Zimbabwe) and South Africa (Tavares, 1967). Resende was a highly productive scientist, with a wide range of interests and a long and diverse list of about 120 publications, many co-authored with his collaborators and students, such as José Pinto-Lopes (1915–1981). In addition, with other colleagues, he founded two new periodicals, Portugaliae Acta Biologica in 1944 and Revista de Biologia in 1956, both of which he supported with his own submissions. He is commemorated in Haworthia resendeana Poelln., Polygala resendeana Paiva, Dryopteris resendeana Res.-Pinto, Placodiscus resendeanus Exell & Mendonça and Bulbochaete resendeana Lacerda. With a highly scientific mind, and being inquisitive, observant, imaginative, and always constructing and testing hypotheses to explain the processes of nature, he was well-ahead of most of his contemporaries who dabbled in the field of botany in Portugal at the time. Electing to remain in Lisbon, where he lacked proper and essential research facilities, Resende did not realise his full potential. Nevertheless, his vision for research would eventually bear fruit. In 1956 the Armenian millionaire Calouste Gulbenkian (1869–1955) bequeathed his fortune to a foundation, the Fundação Calouste Gulbenkian. Within this foundation, and acting on Resende's advice and following his views, a Centre for Biology was created at Oeiras, near Lisbon. It would later become the Instituto Gulbenkian de Ciência. The Centre was built between 1964 and 1966 but Resende, who was appointed as the director of the centre (Tavares, 1967), did not live long enough to see its official opening in July 1967 (Linskens, 1967b). In one of the documents in which he expressed his opinions and views, he argued that scientific ability did not differ among people; rather it differed according to the environment in which they operated. In environments not conducive to research, difficulties had to be endured, often to the detriment of the researcher's family life. It was in those environments that heroes appeared, and not in developed societies where the environment was not adverse to their accomplishments (Resende, 1959a). The unusual number of tributes that were published in the local press when he died, by friends (Lang, 1967; Linskens, 1967a; Machado, 1967), colleagues (Catarino, 1967; Contreiras, 1966; Viana, 1967), students (Oliveira, 1967), and others, reveal in addition to his outstanding scientific ability and his love for science over any financial gain, his intrinsic qualities as a human being: honesty, integrity, loyalty, modesty, and kindness. He was also good humoured, irreverent and easy-going, a “pure soul” (“um puro”, Machado, 1967). He richly deserves to be included among the few botanical heroes of Portugal. Ana Rodrigues (Brotéria Library, Lisbon) and Branca Moriés (Biblioteca e Arquivo Histórico, Museu Nacional de História Natural e da Ciência, Lisbon) are thanked for facilitating access to literature.