Edvard August Vainio

Short description: Finnish lichenologist

Edvard August Vainio

Vainio in 1924 at age 71

Born| (1853-08-05)August 5, 1853

Pieksämäki, Finland

Died| May 14, 1929(1929-05-14) (aged 75)

Turku, Finland

Nationality| Finnish
Alma mater| University of Helsinki
Scientific career
Fields| Lichenology
Institutions| University of Helsinki; University of Turku
Author abbrev. (botany)| Vain.[1]

Edvard August Vainio (né Lang until 1877; until 1919 Wainio)[2] (5 August, 1853 - 14 May, 1929) was a Finnish lichenologist. His works on the cryptogam flora of Lapland, his three-volume monograph on the genus Cladonia, and in particular his study of the natural classification and morphology of lichens in Brazil made Vainio renowned internationally.

Young Vainio's friendship with the ten-years older university student Johan Petter Norrlin (fi) helped him develop an impressive knowledge of local cryptogam flora, and allowed him ample opportunity to hone his collection and identification techniques at an early age. It was through this association that Vainio also met Norrlin's teacher, the prominent lichenologist William Nylander, who supported his early botanical efforts. Vainio's earliest publications dealt with phytogeography or elucidating and enumerating the local flora. Already in these early publications he demonstrated an attention to detail and thoroughness that would later become characteristic of his later work.

Although Vainio was an associate professor of botany at the University of Helsinki during the period 1880–1906, and despite his scientific success and the international recognition he gained through his research, he never obtained a permanent university position with this institution. This was a result, he claimed, of his intense Finnish nationalism and desire to promote the use of the Finnish language in academia during a time of language strife, when Latin dominated the scientific literature, and Swedish was the predominant language of administration and education. Subsequently, he was forced to earn his living, among other things, by working for the Russian censorship authority, which became a heavy burden for him in his academic circles.

Vainio made significant scientific collections of lichens himself, and, as a result of his work as herbarium curator at both the University of Helsinki, and later the University of Turku, he catalogued and processed other collections from all continents, including the Arctic and Antartica. Because of the significance of his works on lichens in the tropics and in general, he has been called the "Father of Brazilian lichenology" and the "Grand Old Man of lichenology".

## Contents

* 1 Early life
* 2 Later life
* 3 Education
* 4 Career
* 4.1 Work abroad
* 4.2 Application for professorship
* 4.3 University of Turku (1919–1928)
* 5 Legacy
* 6 Recognition
* 6.1 Eponymy
* 7 Selected publications
* 8 References
* 8.1 Cited literature

## Early life

Johan Norrlin was Edvard Lang's neighbour, early mentor, and later became his brother-in-law.

Edvard Lang was born on August 5, 1853 in Pieksämäki at a time when Finland was under Russian rule. Edvard was one of several children of parents Carl Johan Lang and Adolfina Polén, both of whom were bailiffs.[3] Young Edvard's early interest in natural history was manifested in his early interest in flowers and his mineral collection;[4] his favourite flower was the marsh willowherb (Epilobium palustre). His eldest brother, Joel Napoleon Lang (fi), was also an avid naturalist; he would later go on to become a well-known legal scholar.[5] In the early 1860s, Carl Johan was transferred to Hollola near Lake Vesijärvi in southern Finland, and the family settled into a farm near Laitila. It was here where Edvard met the phytogeographer Johan Petter Norrlin (fi), who was the neighbour's son. At the time, Norlinn, who was 10 years his senior, was a university student. Norlinn would later marry Vainio's sister in 1873.[4]

Norlinn had become interested in cryptogams after hearing university lectures given by the well-known lichenologist William Nylander at the Imperial Alexander University (later the University of Helsinki), and he later became Nylander's student. He had developed an expertise in the local cryptogam flora, particularly the lichens, which are quite diverse in Finland. Edvard Lang accompanied and assisted him during field trips in the summers of 1868 and 1869, eagerly absorbing and accumulating knowledge.[6] When Norlinn published Beiträge zur Flora des südöstlichen Tavastlands ("Writings on the flora of south-eastern Tavastia Province") in 1870, he credited public school pupil E. Lang for his numerous and valuable contributions to his work.[7]

## Later life

Vainio married Marie Louise Scolastique (née Pérottin),[8] the daughter of a French official, in 1891. They had five children together, although one daughter died soon after birth.[9] His eldest son, with whom he had a close relationship, was the well-known Scout leader and painter Charles Edouard Ilmari (fi) (1892–1955).[10] The walls of the elder Vainio's Turku University office were adorned with portraits of prominent lichenologists that were painted by his son.[11] His other children were Marie Marcienne Alice (1894–1979); Louise (born and died in 1896); Irja Louise Mercedes (1899–1976); and Ahti Victor August (1902–1958).[12] The elder Vainio was described as "a person of retiring habits contented with the bare necessities of life."[13] Adolf Hugo Magnusson, in his obituary of Vainio, recalled the occasion of his 70th birthday party, where Vainio was visited in his home by a group of colleagues from the University of Turku. Although Vainio seemed to be uncomfortable with the attention, he was always willing to draw on his knowledge and extensive memory to give advice and information to inquiring lichenologists.[14]

Regarding his character, his colleague Kaarlo Linkola noted that "he appeared extremely friendly and helpful, though reserved old man, and also a very eccentric personality, with many peculiar features, some of which greatly contributed to his difficult, even tragic life",[8] further noting that "he was extremely obstinate, and he was absolutely unwilling to withdraw from a step which he had once taken."[15] Vainio was dedicated to his research, and could be found working at all hours, even on public holidays.[16] Linkola indicates that he had not taken a day of rest for decades, even when sick.[11] Other biologists in Turku referred to "Vainio's lighthouse", as light would often be seen emerging, often well past midnight, from the lamp coming from the windows of his small room in Turku in the old university building.[17]

Vainio was a patriot and proponent of Finnish nationalism. All his life he had made every effort to support Finnish interests, language and culture to emerge from under the pressure of either the long traditional Swedishness or the attempted Russification of his country by the Russian rulers, who applied every means of pressure to suppress Finnish nationalism.[18] In the 1870s he was involved with pro-Finnish student activism.[8] He was one of the first to replace his non-Finnish name with a Finnish one,[19] Wainio. The name – which means "field"[20] – was taken from a village in Hollola of the same name.[21] He later changed this to the modern Finnish spelling "Vainio" in 1921,[22] in accordance with contemporary changes in Finnish orthography.[23]

Vainio was generally healthy for most of his life, but suffered from severe nephralgia (pain in the kidney) and spent the final three weeks of his life in the hospital of Turku. He died on May 14, 1929, at age 76.[24] He is said to have expressed two great regrets before his death: his uncompleted Lichenographia fennica manuscript, and the infrequency with which he saw his children after his move to Turku.[10]

## Education

Vainio described many new Cladonia species, including C. sobolescens (top), C. subradiata (middle), and C. transcendens (bottom)

Vainio graduated from the Jyväskylä primary college (Jyväskylän Lyseon lukio (fi); now the Jyväskylä Lyceum) in 1870.[25] He began study at the Imperial Alexander University (now the University of Helsinki) the same year, and under the guidance of Norrlin studied phytogeography and lichenology.[26] As a young student, in 1871 Vainio was granted membership in the Societas pro Fauna et Flora Fennica, which is still the oldest scientific society in Finland.[27] Vainio was particularly skilled at identifying and collecting specimens in the field. During the summers of 1873 and 1874 he collected 472 different lichen species from Luhanka and Korpilahti parishes in Central Finland, and in spring of the following year, he discovered 324 species in the vicinity of Vyborg.[4] In one of Nylander's publications,[28] eleven new species were described based on the collections of "E. Lang". A grateful Nylander ordered and sent Vainio a microscope in the summer of 1874 to help him with his botanical studies.[29] In letters between Norrlin and Nylander, the latter praised the young Lang's collecting ability, writing "He is a sharp and fit collector of lichens. With a little work and the help of a decent microscope, he will probably soon surpass everyone else in the North, where no one is better than he in this respect."[30] Lang received his Candidate of Philosophy in 1874.[31]

During his time as a graduate student, Vainio published two works on the cryptogams of Finland: Lichenes in viciniis Viburgi observati ("Lichens observed in the vicinity of Viburg")(1878)[32] and Florula Tavastiae orientalis ("Flora of east Tavastia")(1878),[33] which dealt with the results of his collecting excursions from the years previous.[34] In these publications, Vainio determined the lichen material he collected from the Vyborg region, including new species observations, without assistance from Norrlin or Nylander.[35] Another early publication (Adjumenta ad Lichenographiam Lapponiae fennicae atque Fenniae borealis, published in two parts in 1881 and 1883) was based on material he had collected in 1875 and 1877 in desolate locations near the border between northern Finland and the Russian Karelia, and in Lapland.[34] In these works – considered the earliest publications on phytogeography in the Finnish language – Vainio meticulously catalogued the moisture, light, and soil conditions of the places where he collected, and defined terms that would eventually become standard terminology in the field.[36][37] Because he not only described plant communities but also identified ecological factors that increased or decreased the dominance of different kinds of vegetation and distributional limits for different species, Vainio's work has been described as "ahead of its time".[38] Already in these early publications, the characteristics that would represent his later work were evident:

> "the keen observations, the detailed descriptions and the careful study of the specimens in question. He was never superficial in his work nor prone to hasty inferences however numerous and extensive were the collections submitted to him for examination and for determination. An extreme trustworthiness, thorough investigation and unyielding consistency distinguish the whole of his work".[39]

Nylander, however, was not impressed with Vainio's choice of language for his publications, and this marked the start of a downward turn in their professional relationship.[40] In a letter to Norrlin (dated 20 March 1876), he wrote

> "It is sad for the science as also for Mr Candidate Lang that he has written in Finnish the mentioned work. If he does not want to accommodate Latin he is lost for the intelligent world, and it really would be a great misfortune because he has an excellent talent. But it is true that among the characteristics of childhood and youth, often generously granted by nature, the most common is obstinacy, which has a damaging and destroying direction, destroying both for the individual and his neighborhood. To write special botany in Finnish goes out as if a Frenchman would deliver such a work in Breton or Basque or another of the dialects of the 12 tribes, which jointly constitute the French nation."[20]

In 1880, Vainio defended his dissertation for his licentiate, which, according to the practice of the time, gave him the qualification of a Docent and his teaching rights at the University of Helsinki.[40] His thesis was a study of the phylogeny of Cladonia, a large and widespread genus of fruticose lichens that includes the reindeer lichen and British soldiers lichen species. Titled Tutkimus Cladoniain phylogenetillisestä kehityksestä ("An Investigation on the Phylogenetic Development of Cladonias"), this work was the first dissertation on natural science that was published in the Finnish language.[38] According to Linkola, "this paper of 62 printed pages was sensational on account of its modern theme, as well as its youthful freshness and its originality."[41] Vainio supported the theory of evolution in his work, and proposed that the science of systematics required an examination of phylogeny, rather than mechanical categorization based on sometimes superficial characters.[42] At the same time, Vainio's research contradicted some of Nylander's previous work by presenting shortcomings in his species definitions in Cladonia.[43]

## Career

During his undergraduate days, Vainio took on several temporary positions to support himself. These included work as a translator of Swedish and Finnish for the Uusimaa Provincial Government in 1874; teaching natural history, physics, and gymnastics at a school (Viipurin Realikoulu) in Vyborg in 1875; teaching at the Jyyäskylän seminary (fi) from 1879 to 1881. Starting in 1880, Vainio lectured about botany at the University of Helsinki, where he worked as a docent.[39] His courses consisted of lessons in microscopy, which were mostly given in his home, or in field trips to hunt for cryptogams.[44] Even during his docentship, Vainio continued to work additional modest jobs: teaching botany at the Leppäsuo (fi) horticultural school 1878–1882; natural sciences at the Swedish Private Lyceum 1879–82; the Swedish Real Lyceum 1881–84; the Finnish Primary School 1882–84; the Finnish Girls' School 1882–84, and the Finnish Graduate School 1882–84.[25] Despite all of his experience teaching, he did not enjoy this work, and is said to have had difficulties in maintaining discipline in his classrooms.[15]

### Work abroad

Vainio's research was focused on lichens, and he collected them from all over Finland and abroad. With the help of grants from the university, Vainio made several scientific expeditions abroad around this time.[34] In 1880, accompanying Swedish professor of linguistics August Ahlqvist, he explored the eastern slopes of the Middle Urals in western Siberia.[45] This included the Konda River area extending from the river Irtysh to lake Satyga. The results of this botanical excursion were not published until almost 50 years later.[42] In 1882, he took trips to Berlin and Rostock to botanical museums and herbaria to study the Cladonia specimens located there; in 1884–85 to botanical museums in Moscow, Vienna, Geneva, Paris, and London.[22] It was during a second trip to Paris in 1889–1890 that he would meet his future wife.[46]

During his time in the Caraça Mountains, Vainio stayed at the Santuário do Caraça, shown here. The Pico do Sol is the highest peak in the upper right.

Vainio was one of the first European lichenologists to perform field work in the tropics.[23] After being granted a stipend from the University, in 1885 Vainio undertook a year-long expedition to Brazil, collecting primarily lichens in the vicinity of Rio de Janeiro and in Minas Gerais. He spent some time initially in Sítio (now known as Antônio Carlos), and then in Lafayette (now Conselheiro Lafaiete). Many of his type specimens were collected from these locations.[47] He wrote favourably of the conditions there: "Sítio was a very convenient place for my work: it offered opportunities to study the plant life in the forests as well as in the grasslands. The dryness of the air was also favorable for getting my specimens properly (pressed) and dried."[48] In Rio de Janeiro, Vainio met the French botanist and later landscape designer for Brazilian royalty, Auguste François Marie Glaziou, who advised him on possible travel routes.[49] It was also during this initial part of the trip that he met the French naturalist Bernard Germain de Lacépède, with whom he had several collecting excursions. Lacépède advised Vainio against traveling via his originally intended route, and convinced him instead to visit the biodiverse Caraça Mountains (pt), north of Ouro Branco. It was here that the Caraça sanctuary (pt) was located, a monastery where Lacépède himself had stayed, and which welcomed scientists as guests.[50] Some of the monks residing there were interested in science and collected insects and plants.[51] The monastery had a large library, including works on the local flora, such as Carl Friedrich Philipp von Martius' influential work Flora Brasiliensis. During his stay here Vainio had several collecting excursions with the French entomologist Pierre-Émile Gounelle.[50]

Vainio's tools for fieldwork in Brazil included a knife, hammer, chisel, paper and a bag, and a shotgun for protection against jaguars.[47] In one of his later collecting trips in the Caraça Mountains, Vainio ventured out alone to the highest peak of the eastern mountain ranges, the Pico do Sol (pt)–2,107 m (6,913 ft).[52] Because of his relatively poor knowledge of the terrain, he misjudged the distances involved as well as the amount of available daylight. He ended up spending a night in a wet cave without food, water, or a way to make fire, and suffered a sandfly infestation.[50] It was only the next morning he was able to find a stream to quench his extreme thirst, and not until the afternoon, when, exhausted, he eventually found his way back to the monastery. During his week-long recovery, one of the monks had to extract fly larvae from large bulges on the back of his neck.[52] Nonetheless, by the end of his time in Caraça, he had collected a large volume of specimens.[53] Vainio next went on to Rio de Janeiro, making excursions in coastal areas such as Niterói, Tijuca mountains and Sepetiba region. With the permission of museum director Ladislau de Souza Mello Netto, Vainio performed his studies at the National Museum of Brazil.[53] Vainio returned from Brazil with about 1600 collections packed in five large crates. Vainio worked with this material in Helsinki for the next few years; the material he collected was so abundant that during a few months of study in Paris during 1889–90, he issued "Lichenes brasilienses exsiccati", a set of 1593 exsiccata (dried herbarium specimens) distributed in eight copies.[54]

Cover of Vainio's 1888 popular travel account Matkustus Brasiliassa. Kuvaus luonnostaja kansoista Brasiliassa

In addition to his scholarly work published later, Vainio published in Finnish a popular account of his travels in Brazil, Matkustus Brasiliassa. Kuvaus luonnostaja kansoista Brasiliassa ("Travels in Brazil. A Description of Nature and Travels in Brazil") (1888).[23] This book combines a description of his travel adventures with a folkloric account of Brazil, its flora and fauna and its inhabitants.[55] Vainio does not indicate in this book nor in his later scholarly work the reason for visiting Brazil in the first place. German botanist Fritz Mattick (de) suggests that the idea may have originated from the fact that a number of botanists from the Nordic countries had lived in the interior of Minas Gerais. Examples include Danish naturalist Peter Wilhelm Lund, who lived in Lagoa Santa and made palaeontological discoveries in the nearby limestone caves; and Danish botanists Peter Clausen and his assistant Eugenius Warming. Cladonia specimens that were collected by Warming are mentioned in Vainio's monograph. Swedish physician and botanist Anders Fredrik Regnell was another Nordic scientist who settled in Brazil and made several geological and meteorological discoveries.[55]

In 1887, Vainio published the first of his three-volume monograph on Cladonia; the final volume was published in 1897.[39] This was an extensive work, totalling 1277 pages, on all aspects of this group of taxa. It included descriptions of species old and new, analysis of species synonymy, distribution records, and detailed analysis of the structure and development of the Cladoniae. The publication of just the first volume had already secured Vainio's reputation as a prominent lichenologist.[56] This major work was later judged to be the best work during this era in the field of lichen research.[57] As an indication of the accuracy and reliability of Vainio's work, modern-day examination showed that of the 18 new Cladonia species he described from Brazil a century before, 16 are still considered valid species.[58]

Vainio also published several works based on analysis of collections made by others, for example, those of Erik Acharius, William Nylander,[22] and the late Hungarian botanist Hugó Lojka.[59] In 1899, after the death of Nylander, his numerous collections were transferred from Paris to the University of Helsinki, where it was Vainio's responsibility to arrange and catalog it – a total of 51,066 specimens.[56] Although his relationship with the university was strained at the time, there was no-one else qualified for the job.[60] Similarly, Vainio published works based on collections he was sent from Puerto Rico,[61] Japan,[62][63] Thailand,[64] Tahiti,[65] and Trinidad,[66] among other locations. In some instances, his studies of collections sent to him greatly advanced the knowledge of the local flora from which they were sent. For example, Vainio was sent for identification the collections of Portuguese botanist and army doctor Américo Pires de Lima (pt), who made the collections as part of a military campaign in Mozambique during 1916–1917. Vainio's results were published posthumously;[67] of the 138 taxa he identified, about half were described as new to science.[68]

### Application for professorship

As the culmination of his studies in Brazil, Vainio published Etude sur la classification naturelle et la morphologie des lichens du Brésil ("Study on the natural classification and the morphology of the lichens of Brazil") in 1890. This 526-page work dealt with 516 species, of which 240 were new to science.[56] The Brazilian taxa were distributed amongst 78 genera (12 of which were described as new), the most well-represented of which included Lecidea (68 species), Graphis (43), Parmelia (39), Lecanora (33), Arthonia (25) and Buellia (19). The genus Cladonia was not included in this work, as he reserved it for his Cladonia monograph.[69] Vainio also discussed the general theory of lichens, supporting Simon Schwendener's then-controversial theory that lichens were the result of a symbiotic union between fungus and alga.[56] Vainio advocated including the lichens in the general classification of the fungi. He argued that lichens are a polyphyletic group, with only one uniting characteristic—the symbiosis—distinguishing them from the ascomycetes and other fungi.[70]

Vainio's work was intended to be a thesis submission for the Chair of botany at the University of Helsinki, to replace the position held by the eminent bryologist Sextus Otto Lindberg.[22] Norrlin had obtained a similar position in 1878, which perhaps inspired Vainio to make the application.[71] Two other docents competed for the position: Fredrik Elfving (fi) and Oswald Kairamo.[70] Because Lindberg did not trust his Finnish language skills sufficiently to be able to judge the merits of Vainio's work, other opinions were sought, and so in addition to Nylander, Theodor Magnus Fries, and Johann Müller were recruited.[72]

Most prominent contemporary lichenologists, including Gustav Wilhelm Körber, Arthur Minks, Johann Müller, and William Nylander, disagreed with the so-called "Schwendenerian hypothesis" and the dual nature of lichens.[73] They thought that Vainio's proposal to classify lichens with fungi was ridiculous, as they considered lichens a plant group.[74] Müller in particular published two articles that were highly critical of Vainio's conclusions in Études Brésil.[75][76] Vainio's relationship with Nylander had become strained since their successful collaborations years before. Nylander, in previous correspondence with Norrlin, expressed doubts about Vainio's decision to publish his early scientific works in Finnish instead of Latin, which was the norm in the international scientific community. He also questioned Lang's decision to change his name, writing "A most curious matter is also the disappearance of Mr Lang and the birth of Mr Wainio instead. This is a matter that may be possible and explainable in Finland (and unfortunate is that such is the situation) but in the common practical world, here in the logical humanity, such a thing is impossible even to mention without incurably injuring the person concerned."[20] Vainio, in correspondence with Johann Müller in 1889, wrote "it is perhaps necessary that the knowledge of my thesis remains between us, because there are people who lay very peculiar intrigues to prevent me from the professorship. Nylander has taken a stand as a very unscrupulous enemy against me and has taken up a very scandalous intrigue".[72]

Johannes Müller, William Nylander, and Theodor Magnus Fries were prominent botanists invited to comments on the merits of Vainio's research for his dissertation.

Nylander criticized and dismissed Vainio's work, claiming it had little scientific value. In contrast, Fries praised Vainio's work, and described him as one of the most competent contemporary lichenologists.[72] Johann Müller disagreed with most of Vainio's general conclusions, and thought that chemical reactions, a characteristic that Vainio emphasized, have only a physiological, not taxonomical value.[77] Although Müller was public about his criticisms to Vainio's work, he did acknowledge his careful working method and anticipated that Vainio in future research, "after return from wrong paths", would use his excellent observational skills in a systematically correct way.[77] German lichenologist Ferdinand Christian Gustav Arnold, who was also present at Vainio's public thesis defence, introduced himself as a supporter of Schwenderer's theory, and indicated that Vainio's work was the first to create a consistent system for classification.[77] The idea of merging the classification of non-lichenized fungi and lichenized fungi did not gain general support until the mid-20th century.[78]

Vainio did not receive the professorship for which he applied. Elfving was given the position; he later became known infamously for his erroneous views on the nature of photobionts.[79] Possible reasons could be Vainio's narrow, mainly lichen-focused field of expertise, lack of teaching skills, and the personal resentments developed between Vainio and Nylander, his former teacher, as well as issues of language policy. Vainio championed Finnish interests and was a strong supporter of the Finnish language, but at that time Finland was still part of the Russian Empire and the position of the Finnish language in teaching was weak.[74] Suspecting that he had been discriminated against in the selection of a professor, Vainio appealed the decision, claiming that the expert opinions came from representatives of an "openly hostile" school that discriminated against him, and further, that he was the only one of the applicants with the ability to lecture fluently in both Finnish and Swedish. He concluded that he had been rejected on political rather than scientific grounds. The university's position was that the more general studies on topics provided better guarantees for the successful management of the professorship.[80]

Vainio in his private study at the University of Turku Plant Museum, 1925

Vainio's failure to secure a professorship must have been a further blow to him personally in the sense that, like Norrlin, his big brother Joel Napoleon Lang had successfully pursued a university career as a professor in the Faculty of Law.[81] Unsuccessful in his bid for professorship, and convinced that a supporter of an independent Finland like him would never be elected to university duties, Vainio began working as a censor in the press service of Helsinki in 1891, a position in which he was appointed superintendent in 1901.[22] It was during this time that the Russian Empire pursued the policy of Russification, a mandate carried out by the polarizing figure Governor-General Nikolay Bobrikov.[81] Vainio suffered deeply from the consequences caused by this action, although he defiantly hid his distress.[82] His decision to work for the loathed Board of Press Censorship led to him becoming a pariah amongst his colleagues and compatriots.[44] For example, despite their innovativeness and importance, Vainio's early publications on phytogeography in the border regions of northeast Finland and Russian Karelia were rarely cited by his Finnish colleagues, largely for political reasons.[38] Another source suggests that additional resentment amongst his colleagues was stoked by his publication of the first Finnish-language dissertation.[74] Shortly after the turn of the century, Vainio was forced to suspend his teaching position as he had so few students.[44] After Finland gained independence in 1917, he was left without work and without a pension at the age of 64. Forced to live on modest savings, Vainio continued his lichenological studies. He transferred his microscope and part of his library to the botanical institution of the university, where he spent much of his time for the next couple of years.[44]

### University of Turku (1919–1928)

Vainio's fortunes improved in 1918, as the Turku Finnish University Society bought his herbarium collection of about 22,000 specimens for 60,000 FIM. The society was organizing a new university at Turku, which was then the second largest city in Finland after Helsinki. The university was intended to be entirely Finnish, in contrast with the University of Helsinki, which taught both in Swedish and Finnish.[83] The transaction was also subject to the condition that Vainio himself would be responsible for organizing and increasing the collection in museum condition, and would also participate in teaching if necessary.[84][83] As an eager Finnish nationalist, Vainio was pleased with the arrangement and started on the payroll of the Turku University Society under the title of custodian of the collections of the Department of Botany in 1920, two years before the start of the university's teaching activities and the transfer of his collection to Turku.[85] He moved to Turku and the university's main building on the edge of the market square in the former Phoenix Hotel when teaching began in 1922.[84] Although only offered a modest yearly salary to organize the specimens, he carried out this task with great devotion.[86] He obtained this job—his only permanent teaching position—at 69 years of age, and held it until his death.[87] His living conditions, however, remained so modest that his wife and family were unable to visit him in Turku, and their visits were limited to his vacations in Helsinki.[54] In order to optimize his productivity of his holiday time, he would take the evening train from Turku to Helsinki, and could be found the next morning in the lichen department of the Helsinki Plant Museum.[88]

The University of Turku as it appeared during its grand opening in 1922.

In 1921, at the initiative of Alvar Palmgren,[89] Vainio was commissioned by the Societas pro Fauna et Flora Fennica to continue work on Lichenographia fennica, a seven-part book series about Finnish lichens.[90] Knowing he had only limited time left to complete a multi-volume series because of his age, he started work on the harder groups, confident that in the event of his death the easier groups could be handled by other researchers.[74] This book series became an important resource for the study of the lichen flora of all Northern Europe.[91]

Starting 1922, Vainio taught as an assistant professor at the University of Turku, and headed the cryptogamic herbarium at the university.[86] His teaching consisted of courses in plant systematics, and organized field trips with students.[91] This field work he continued until 1927, leading a class expedition to a small island in Lake Ladoga.[17] During his time at the University of Turku, the collections expanded to 35,000 samples, a result of additions from local excursions, and collections sent from abroad.[86] Vainio also advised younger colleagues Kari Linkola and Veli Räsänen (fi).[17] He was granted a state pension while on his deathbed.[86]

Vainio's final work, the fourth volume of the Lichenographia fennica, was left uncompleted on his worktable because of his death. His last entry was to name and describe Lecidea keimioeënsis (collected by Linkola in Keimiötunturi (fi)) as a new species, when his illness suddenly forced him to stop work and hurry to the hospital.[92] Started by Vainio in 1924, the fourth volume was completed posthumously by Norwegian lichenologist Bernt Lynge in 1934.[3]

## Legacy

Vainio described about 1700 taxa, circumscribed several new genera, and emended several existing ones.[22] He had 87 publications in his career. Most of his work dealt with lichens, although he occasionally published about other related topics.[46] Examples include the plant and cryptogam floras of Tavastehus and the northern Finland and Russian Karelia border area, a discussion of Salix hybrids, a listing of phanerogams in Finnish Lapland, and a list of the vascular cryptogams and mosses from the area of the Konda river in Western Siberia. Vainio described and catalogued collections from all continents, including the Arctic (Greenland) and Antarctic.[46] Finnish botanist Reino Alava, who was a curator of the University of Turku herbarium, compiled a comprehensive listing of the location of all of Vainio's type specimens in a 1988 publication,[93] and twenty years later, a list of all collectors whose collections are represented in Vainio's lichen herbarium in Turku.[94] Because of Vainio's pioneering works on Brazilian lichenology and his extensive collecting in Caraça, it has since become an international hub for lichenology and a destination for pilgrimages for lichenologists.[95] Vainio is generally considered to have made the most important contributions to the study of foliose lichens in the neotropics prior to the work of Rolf Santesson.[96]

Vainio's idea of integrating the classification of lichens and fungi represented a criticism of the prevailing ideas of 19th-century lichenology. These outdated ideas would persist into the first half of the 20th century, largely due to the publication of Zahlbruckner's influential Catalogus series, issued in ten volumes from 1922 up until 1940, which was based on these old views.[97] Although Vainio understood that the ideal classification scheme would have position lichen genera close to their nearest non-lichenized relatives, all he could realistically do with the information he had available was assign lichens and ascomycetes to one group, although the lichens were placed in separate classes, the Discolichenes and Pyrenolichenes.[98] It was at the International Botanical Congress in Stockholm in 1950 that Rolf Santesson advocated for Vainio's ideas and presented an integrated classification for fungi and lichens based on an updated system developed by John Axel Nannfeldt.[99] This initiated discussions and an eventual consensus for an integrated classification system.[100]

Some of the species Vainio described as new to science in the "script lichen" genus Graphis include G. crebra (top), G. leptospora (middle), and G. plumierae (bottom).

Vainio's infrageneric subdivision of the genus Parmelia laid the nomenclatural cornerstone to two later recognized genera, Hypotrachyna and Xanthoparmelia (later raised to generic status by Mason Hale[101]), as well as for Allantoparmelia, which was promoted to genus by Theodore Esslinger.[102] By describing the section Amphigymnia of the genus Parmelia, Vainio, in his treatment of Brazilian lichens (1890), had an essential role in the separation of species which are now part of the genus Parmotrema.[103] In the family Lobariaceae, Vainio segregated the genus Pseudocyphellaria for species having pseudocyphellae and not true cyphellae on the lower surface of the thallus. This was a radical idea at the time, as the presence or absence of cyphellae and pseudocyphellae were not considered to be taxonomic and generic characters. Although some other influential lichenologists took a conservative view and lumped Pseudocyphellaria with Sticta (such as Zahlbrucker in his influential series Catalogus Lichenum Universalis, Vainio's concept of the genus prevailed and has been used extensively for over a century. Later work has shown the presence of pseudocyphellae to correlate strongly with a diverse secondary chemistry consisting of orcinol derivatives, beta-orcinol derivatives, triterpenoids, terphenylquinones and 4-ylidenetetronic acids; the genus Sticta, in contrast, does not produce these compounds.[104] Vainio also introduced in this same work the current concept for the genus Lobaria, which at the time was broadly used for foliose lichens.[105]

## Recognition

In his memorial address, Alvar Palmgren, then President of Societas pro Fauna et Flora Fennica, recalled that a considerable part of Vainio's scientific papers appeared in the Society's publications and were among the best of them.[17] Vainio's travels in Brazil were recounted in Reinio Alava's 1986 book Edvard August Vainio's Journey to Brazil in 1885 and his Lichenes Brasilienses Exsiccati. Based on Vainio's diaries, it describes the difficulties he experienced in collecting in a tropical foreign country.[106]

Vainio's 3-volume Cladonia monograph was reprinted in 1978.[107] Although at the time of reprinting some parts of the book were quite outdated, a review noted "[I]t is no ordinary monograph, but one which has a long-standing value as a taxonomic, floristic, and bibliographic source. One of its outstanding features is its almost infallible reliability as a nomenclatural source", and that "[F]or many significant details on the world's Cladonia's, Vainio still gives the freshest information!".[108]

In 1997, a symposium on Vainio and his work was organized in Brazil by the Grupo Latino-Americano de Liquenólogos and the International Association for Lichenology. Held at the Caraça Monastery he stayed at during his collecting trip there over a century prior (now a hotel), Vainio was declared the "Father of Brazilian lichenology" by the participants. A portrait of Vainio, donated by the University of Turku, was mounted in one of the main corridors.[109] A book containing the proceedings of symposium was issued in 1998, Recollecting Edvard August Vainio. Written by several specialists on various lichen groups, it reviews his contributions to tropical lichenography, and biographic details about him and his travels, publications, and collections.[16] He is also known as the "Grand Old Man of lichenology",[107] a sobriquet originally given to him by Bernt Lynge: "Through all of his papers Dr. Vainio has acquired an uncontested position as the Grand Old Man of Lichenology. He is an ornament to his science and an honour to his country."[110][111] Because of his significant contributions to the knowledge of the family Graphidaceae in the Philippines, he has also been called the "Father of Philippine lichenology".[112]

### Eponymy

Five genera are named after Vainio, although most of these names are now obsolete and have been sunk into synonymy with other genera:[2]

* Vainiona Werner (1943) (= Cercidospora[113])
* Vainionia Räsänen (1943) (= Calicium[114])
* Vainionora Kalb (1991)[115]
* Wainioa Nieuwl. (1916) (= Byssoloma[116])
* Wainiocora Tomas. (1950) (= Cora[117])

Many species have also been named to honour Vainio. These include:[19] Teichospora wainioi P.Karst. (1884); Nectriella vainioi P.Karst. (1889); Meliola wainioi Pat. (1890); Filaspora wainionis Kuntze (1898); Clathroporina wainiana Zahlbr. (1902); Cladonia wainioi Savicz (1914); Physcia wainioi Räsänen (1921); Opegrapha wainioi Zahlbr. (1923); Pannaria wainioi Zahlbr. (1925); Rhizocarpon vainioense Lynge (1926); Peltigera vainioi Gyeln. (1929); Pannaria vainioi C.W.Dodge (1933); Usnea vainioi Motyka (1936); Nesolechia vainioana Räsänen (1939); Melanotheca vainioensis Werner (1944); Lecidea vainioi H.Magn. (1949); Tricharia vainioi R.Sant. (1952);[118] Candelariella vainioana Hakul. (1954); Caloplaca vainioi Hafellner & Poelt (1979); Lecanora vainioi Vänskä (1986); Bulbothrix vainioi Jungbluth, Marcelli & Elix (2008);[119] Gyalideopsis vainioi Kalb & Vězda (1988);[120] Hypotrachyna vainioi Sipman, Elix & T.H.Nash (2009); and Coppinsidea vainioana S.Y.Kondr., E.Farkas & L.Lőkös (2019).[121]

## Selected publications

A complete listing of Vainio's scientific publications is given in Schulz-Korth's 1930 Hedwigia  obituary,[122] and also on the University of Turku's Museum of Natural Science webpage.[123] Vainio's major works include:

* Wainio, Edvard August (1887). Monographia Cladoniarum universalis: I.". Acta Societatis pro Fauna et Flora Fennica. 4. pp. 1–509. https://www.biodiversitylibrary.org/page/5587642.
* ~~ ~~ (1890) (in fr, la). Étude sur la classification naturelle et la morphologie des Lichens du Brésil, I–II. Acta Societatis pro Fauna et Flora Fennica. 7. Helsinki: J. Simelius. pp. 1–247; 1–256. https://www.biodiversitylibrary.org/page/4458421.
* ~~ ~~ (1894). Monographia Cladoniarum universalis: II. Acta Societatis pro Fauna et Flora Fennica. 10. pp. 1–499. https://www.biodiversitylibrary.org/page/5583213.
* ~~ ~~ (1897). Monographia Cladoniarum universalis: III. Acta Societatis pro Fauna et Flora Fennica. 14. pp. 1–268. https://www.biodiversitylibrary.org/page/5634825.
* Vainio, E. (1909) (in Latin). Lichenes in viciniis stationis hibernae expeditionis Vegae prope pagum Pitlekai in Sibiria septentrionali a D:re E. Almquist collecti. 8. 11–175. https://www.biodiversitylibrary.org/page/29155446.
* ~~ ~~ (1909). Lichenes insularum Philippinarum. I.. 4. 651–662. https://www.biodiversitylibrary.org/page/699093.
* ~~ ~~ (1913). Lichenes insularum Philippinarum. II.. 8. 99–137. https://www.biodiversitylibrary.org/page/51382087.
* ~~ ~~ (1921). "Lichenes insularum Philippinarum. III.". Annales Academiae Scientiarum Fennicae Series A 15 (6): 1–368.
* ~~ ~~ (1921). Lichenographia Fennica I. Pyrenolichenes iisque proximi Pyrenomycetes et Lichenes imperfecti. Acta Societatis pro Fauna et Flora Fennica. 49. pp. 1–274. https://www.biodiversitylibrary.org/page/5595130.
* ~~ ~~ (1922). Lichenographia Fennica II.. Acta Societatis pro Fauna et Flora Fennica. 51. pp. 1–340.
* ~~ ~~ (1923). "Lichenes insularum Philippinarum. IV.". Annales Academiae Scientiarum Fennicae Series A 19 (15): 1–84.
* ~~ ~~ (1927). Lichenographia Fennica III. Coniocarpaceae. Acta Societatis pro Fauna et Flora Fennica. 57. pp. 1–138. https://helda.helsinki.fi/bitstream/handle/10138/15818/1927ActasocFFF571.pdf?sequence=1.
* ~~ ~~ (1934). Lichenographia Fennia IV. Lecideales 2. Acta Societatis pro Fauna et Flora Fennica. 57. pp. 1–531. https://helda.helsinki.fi/bitstream/handle/10138/15819/1934ActasocFFF572.pdf?sequence=1.

## References

1. ↑ "Vainio, Edvard (Edward) August (1853–1929)". International Plant Names Index. https://www.ipni.org/a/12959-1. Retrieved 15 July 2020.
2. ↑ 2.0 2.1 Stafleu, Frans A.; Cowan, Richard S. (1986). Taxonomic literature: A selective guide to botanical publications and collections with dates, commentaries and types. Volume VI: Sti–Vuy. 6 (2nd ed.). Utrecht: Bohn, Scheltema & Holkema. pp. 636–637. https://www.biodiversitylibrary.org/page/33212678.
3. ↑ 3.0 3.1 Linkola 1934, p. 3.
4. ↑ 4.0 4.1 4.2 Linkola 1934, p. 5.
5. ↑ Linkola 1929, p. 155.
6. ↑ Linkola 1934, p. 4.
7. ↑ Magnusson 1930, p. 5.
8. ↑ 8.0 8.1 8.2 Linkola 1934, p. 18.
9. ↑ Vitikainen 1998, p. 24.
10. ↑ 10.0 10.1 Linkola 1929, p. 167.
11. ↑ 11.0 11.1 Linkola 1929, p. 165.
12. ↑ "Helsingin yliopiston dosenttiyhdistys: WAINIO (→1872 Lang), Edvard August" (in Finnish). University of Helsinki. https://www.helsinki.fi/sites/default/files/atoms/files/p-o.pdf. Retrieved 15 July 2020.
13. ↑ Magnusson 1930, p. 8.
14. ↑ Magnusson, A.H. (1929). "In memoriam. E. Vainio" (in Swedish). Botanisker Notiser 4: 278–280. https://journals.lub.lu.se/bn/article/view/11043/9846.
15. ↑ 15.0 15.1 Linkola 1934, p. 14.
16. ↑ 16.0 16.1 Hawksworth, David L. (2007). "Book Reviews. Recollecting Edvard August Vainio". The Lichenologist 31 (4): 405–406. doi:10.1017/S0024282999000535.
17. ↑ 17.0 17.1 17.2 17.3 Vitikainen 1998, p. 26.
18. ↑ Alava 1998, p. 9.
19. ↑ 19.0 19.1 Hertel, Gärtner & Lőkös 2017, pp. 156–157.
20. ↑ 20.0 20.1 20.2 Vitikainen 1998, p. 18.
21. ↑ Linkola 1929, p. 166.
22. ↑ 22.0 22.1 22.2 22.3 22.4 22.5 Magnusson 1930, p. 7.
23. ↑ 23.0 23.1 23.2 Buck, William R. (1994). "Vainio's Brazilian bryophytes at TUR". The Bryologist 97 (4): 420–423. doi:10.2307/3243910.
24. ↑ Linkola 1934, p. 17.
25. ↑ 25.0 25.1 Linkola 1929, p. 154.
26. ↑ Magnusson 1930, pp. 5–6.
27. ↑ Vitikainen 1998, p. 16.
28. ↑ Nylander, W. (1874). "Addenda nova ad lichenographiam europaeam. Contin. XVIII" (in Latin). Flora (Regensburg) 57: 305–318. https://www.biodiversitylibrary.org/page/62230.
29. ↑ Vitikainen 1998, p. 17.
30. ↑ Tarmio 2000, p. 159: "Hän on terävä ja kelpo jäkälien kerääjä. Hieman työskentelemällä ja kelvollisen mikroskoopin avulla hän varmaan pian ylittää kaikki muut Pohjolassa, jossa ei näy ketään häntä parempaa kykyä tässä suhteessa."
31. ↑ Alava 1998, p. 2.
32. ↑ Wainio, E.A. (1878). "Lichenes in viciniis Viburgi observati" (in Finnish). Meddelanden af Societas pro Fauna et Flora Fennica 2: 35–72. https://www.biodiversitylibrary.org/page/3735417.
33. ↑ Wainio, E.A. (1878). "Florula Tavastiae orientalis" (in Finnish). Meddelanden af Societas pro Fauna et Flora Fennica 3: 1–121. https://www.biodiversitylibrary.org/page/3844596.
34. ↑ 34.0 34.1 34.2 Linkola 1934, p. 8.
35. ↑ Tarmio 2000, p. 159.
36. ↑ Linkola 1934, p. 6.
37. ↑ Haikurainen 1961, pp. 6–7.
38. ↑ 38.0 38.1 38.2 Kaila, Annu; Vasander, Harri (2010). "Vegetation relations in the border regions of NE Finland and Russian Karelia: a review of Wainio (1878)". Scandinavian Journal of Forest Research 26 (S10): 65–71. doi:10.1080/02827581.2011.517949.
39. ↑ 39.0 39.1 39.2 Magnusson 1930, p. 6.
40. ↑ 40.0 40.1 Tarmio 2000, p. 160.
41. ↑ Linkola 1934, p. 7.
42. ↑ 42.0 42.1 Alava 1998, p. 3.
43. ↑ Tarmio 2000, p. 161.
44. ↑ 44.0 44.1 44.2 44.3 Linkola 1934, p. 15.
45. ↑ Schulz-Korth 1930, p. 1.
46. ↑ 46.0 46.1 46.2 Schulz-Korth 1930, p. 2.
47. ↑ 47.0 47.1 Ahti 1998, p. 41.
48. ↑ Galloway 1998, p. 62.
49. ↑ Mattick 1956, p. 414.
50. ↑ 50.0 50.1 50.2 Alava 1998, p. 5.
51. ↑ Mattick 1956, p. 415.
52. ↑ 52.0 52.1 Mattick 1956, p. 416.
53. ↑ 53.0 53.1 Alava 1998, p. 6.
54. ↑ 54.0 54.1 Mattick 1956, p. 405.
55. ↑ 55.0 55.1 Mattick 1956, p. 413.
56. ↑ 56.0 56.1 56.2 56.3 Linkola 1934, p. 9.
57. ↑ Tarmio 2000, p. 164.
58. ↑ Ahti 1998, p. 43.
59. ↑ Gyelnik, Vilmos (1930). "Meghalt. – Gestorben. E.A. Vaino" (in Hungarian). Magyar Botanikai Lapok 28: 176–178. http://real-j.mtak.hu/9912/1/MTA_MagyarBotanikaiLapok_28.pdf.
60. ↑ Tarmio 2000, p. 167.
61. ↑ Vainio, E.A. (1929). "New species of lichens from Porto Rico. II". Mycologia 21 (1): 33–40. http://www.cybertruffle.org.uk/cyberliber/59350/0021/001/0033.htm.
62. ↑ Wainio, Dr. Edv. A. (1918). "Lichenes ab A. Yasuda in Japonia collecti" (in Latin). Botanical Magazine, Tokyo 32: 154–163. https://www.biodiversitylibrary.org/page/41605467.
63. ↑ Wainio, Dr. Edv. A. (1921). "Lichenes ab A. Yasuda in Japonia collecti. Continuatio 1." (in Latin). Botanical Magazine Tokyo 35: 45–79. https://www.biodiversitylibrary.org/page/5431838.
64. ↑ Vainio, E.A. (1921). "Lichenes Siamenses". Annales Zoologici Societatis Zoologicae Botanicae Fennicae 1 (3): 33–55.
65. ↑ Vainio, E.A. (1924). "Lichenes a W.A. Setcheli and K.E. Parks in insula Tahiti a. 1922 collecti". University of California Publications in Botany 12: 1–16.
66. ↑ Vainio, Edward A. (1923). "Lichenes in insula Trinidad a Professore R. Thaxter collecti". Proceedings of the American Academy of Arts and Sciences 58 (3): 131–147. doi:10.2307/20025979.
67. ↑ Vainio, E.A. (1929). "Lichens Mozambici". Boletim da Sociedade Broteriana. Vol. II (II Série) 6: 144–179. https://bibdigital.rjb.csic.es/viewer/10658/?offset=#page=163&viewer=picture&o=bookmark&n=0&q=.
68. ↑ Paz-Bermúdez, Graciela (2004). "The new taxa of lichens and other ascomycetes from Mozambique collected by Pires de Lima and described by E. A. Vainio". Taxon 53 (2): 511–519. doi:10.2307/4135630.
69. ↑ Mattick 1956, p. 406.
70. ↑ 70.0 70.1 Vitikainen 1998, p. 21.
71. ↑ Tarmio 2000, p. 165.
72. ↑ 72.0 72.1 72.2 Vitikainen 1998, p. 20.
73. ↑ Mattick 1956, p. 407.
74. ↑ 74.0 74.1 74.2 74.3 "Edvard August Vainio (1853–1929)" (in Finnish). University of Turku, Museum of Natural Sciences. https://collections.utu.fi/kasvimuseo-tur-2/tur-v/edvard-august-vainio-1853-1929/. Retrieved 15 July 2020.
75. ↑ Müller, J. (1891). "Kritik über Dr. Wainio's Étude" (in German). Flora 74: 383–389. https://www.biodiversitylibrary.org/page/58054897.
76. ↑ Müller, J. (1892). "Critique de ”l'Étude” du Docteur Wainio" (in French). Revue de Mycologie 13: 33–38. https://www.biodiversitylibrary.org/page/11342839.
77. ↑ 77.0 77.1 77.2 Vitikainen 1998, p. 22.
78. ↑ Tibell 1998, pp. 96–97.
79. ↑ Vitikainen 1998, p. 23.
80. ↑ Tarmio 2000, pp. 165–166.
81. ↑ 81.0 81.1 Tarmio 2000, p. 166.
82. ↑ Linkola 1929, p. 162.
83. ↑ 83.0 83.1 Stenroos 1998, p. 29.
84. ↑ 84.0 84.1 Tarmio 2000, p. 158.
85. ↑ Tarmio 2000, p. 168.
86. ↑ 86.0 86.1 86.2 86.3 Linkola 1934, p. 16.
87. ↑ Ahti 1998, p. 39.
88. ↑ Linkola 1929, p. 164.
89. ↑ Linkola 1934, p. 11.
90. ↑ Vitikainen 1998, p. 25.
91. ↑ 91.0 91.1 Alava 1998, p. 10.
92. ↑ Linkola 1929, p. 160.
93. ↑ Alava, R. (1988). Edvard August Vainio’s Types in TUR-V and Other Herbaria. Publications from the Herbarium of the University of Turku. 2. Turku: University of Turku. pp. 1–513. ISBN 978-951-88-0200-9.
94. ↑ Alava, R. (2008). Index of Collectors whose Specimens are part of Edv. Aug. Vainio's Lichen Herbarium. Publications from the Herbarium of the University of Turku. 12. Turku: University of Turku. pp. 1–123. ISBN 978-951-29-3369-3.
95. ↑ Ahti 1998, p. 44.
96. ↑ Lücking 1997, p. 89.
97. ↑ Tibell 1998, p. 96.
98. ↑ Mitchell 2015, p. 90.
99. ↑ Santesson, R. (1950). "The new systematics of lichenized fungi". Proceedings of the Seventh International Botanical Congress. Stockholm: Almqvist & Wiksell. pp. 809–810. https://archive.org/details/in.ernet.dli.2015.351670/page/n825/mode/2up.
100. ↑ Tibell 1998, p. 97.
101. ↑ Hale, Mason E. (1974). "New combinations in the lichen genus Parmotrema Massalongo.". Phytologia 28 (4): 334–339. https://www.biodiversitylibrary.org/page/13046332#page/348/mode/1up.
102. ↑ Esslinger, Theodore L. (1978). "A new status for brown Parmeliae". Mycotaxon 7 (1): 45–54. http://www.cybertruffle.org.uk/cyberliber/59575/0007/001/0045.htm.
103. ↑ Feuerer 1998, p. 47.
104. ↑ Galloway 1998, p. 75.
105. ↑ Yoshimura 1998, p. 86.
106. ↑ "Briefly Noted – Edvard August Vainio's Journey to Brazil in 1885 and his Lichenes Brasilienses Exsiccati. By Reino Alava. [Publications from the Herbarium, University of Turku, no. 1.] Turku: University of Turku. 1986. Pp. 174, 5 figures. ISBN 951 642 878 9". The Lichenologist 21 (1): 97–98. 2007. doi:10.1017/S0024282989000198.
107. ↑ 107.0 107.1 "Vainio ['Wainio'"]. Dictionary of the Fungi (10th ed.). Wallingford, UK: CAB International. 2008. p. 719. ISBN 978-0-85199-826-8. https://archive.org/details/dictionaryfungit00kirk.
108. ↑ Ahti, T. (2007). "Books Reviews. Monographia Cladoniarum Universalis I–III. By E. A. Wainio [= Vainio]. [Reprinted from Acta Soc. Fauna Fl. fenn. 4: 1–509 (1887), 9: 1–268 (1897), 10: 1–499 (1894).] Koenigstein: Otto Koeltz Science Publishers. 1978. 3 vols. Pp. 509 + 268 + 499. Price DM 400.-.". The Lichenologist 12 (1): 155–156. doi:10.1017/S0024282980000102.
109. ↑ Marcelli & Ahti 1998, preface.
110. ↑ Alava 1998, p. 11.
111. ↑ Linkola 1929, p. 161: "Kaikilla näillä teoksillaan on toht. Vainio hankkinut itselleen jäkälätieteen Grand old man'in riidattoman aseman. Hän on tieteensä kaunistus ja kunniaksi maallensa."
112. ↑ Fajardo, Weenalei T.; Bawingan, Paulina A. (2019). "Taxonomy and new records of Graphidaceae lichens in Western Pangasinan, Northern Philippines". Philippine Journal of Systematic Biology 13 (2): 40–54. doi:10.26757/pjsb2019b13006. http://asbp.org.ph/wp-content/uploads/2020/03/PJSB_S01-2019-006.pdf.
113. ↑ "Record details: Vainiona Werner". Index Fungorum. http://www.indexfungorum.org/Names/NamesRecord.asp?RecordID=5697. Retrieved 15 July 2020.
114. ↑ "Record details: Vainionia Räsänen". Index Fungorum. http://www.indexfungorum.org/Names/NamesRecord.asp?RecordID=5698. Retrieved 15 July 2020.
115. ↑ Kalb, Klaus (1991) (in German). Lichenes Neotropici ausgegeben von Klaus Kalb. Fascikel XII (No. 476–525). Neumarkt in der Oberpfalz: Self-published. OCLC 163537210.
116. ↑ "Record details: Wainioa Nieuwl.". Index Fungorum. http://www.indexfungorum.org/Names/NamesRecord.asp?RecordID=5757. Retrieved 15 July 2020.
117. ↑ "Record details: Wainiocora Tomas.". Index Fungorum. http://www.indexfungorum.org/Names/NamesRecord.asp?RecordID=18744. Retrieved 15 July 2020.
118. ↑ Santesson, Rolf (1952). Foliicolous lichens. I. A revision of the taxonomy of the obligately foliicolous, lichenized fungi. Symbolae Botanicae Upsalienses. 12. p. 382.
119. ↑ Jungbluth, Marcelli & Elix 2008, p. 62.
120. ↑ Kalb, Klaus; Vězda, Antonin (1988). "Neue oder bemerkenswerte Arten der Flechten-familie Gomphillaceae in der Neotropis" (in German). Bibliotheca Lichenologica (Lubrecht & Cramer) 29: 51. ISBN 978-3-443-58008-7.
121. ↑ Kondratyuk et al., p. 302.
122. ↑ Schulz-Korth 1930, pp. 5–9.
123. ↑ "Vainion julkaisut [Vainio's publications"] (in Finnish). University of Turku, Museum of Natural Sciences. https://collections.utu.fi/kasvimuseo-tur-2/tur-v/vainion-julkaisuja/. Retrieved 15 July 2020.

### Cited literature

* Haikurainen, Leo (1961). "Swamp forestry research in Finland". Silva Fennica 108: 1–20. https://silvafennica.fi/pdf/article4696.pdf.
* Hertel, Hannes; Gärtner, Georg; Lőkös, László (2017). "Forscher an Österreichs Flechtenflora" (in German). Stapfia 104 (2): 1–211. https://www.zobodat.at/biografien/Flechtenforschung_Austria_STAPFIA_0104_2_0001-0211.pdf.
* Jungbluth, Patricia; Marcelli, Marcelo Pinto; Elix, John A. (2008). "Five new species of Bulbothrix (Parmeliaceae) from cerrado vegetation in São Paulo State, Brazil". Mycotaxon 104: 51–63. http://www.cybertruffle.org.uk/cyberliber/59575/0104/0062.htm.
* Kondratyuk, S.Y.; Lőkös, L.; Farkas, E.; Jang, S.-H.; Liu, D.; Halda, J.; Persson, P.-E.; Hansson, M. et al. (2019). "Three new genera of the Ramalinaceae (lichen-forming Ascomycota) and the phenomenon of presence of ‘extraneous mycobiont DNA’ in lichen associations". Acta Botanica Hungarica 61: 275–323. doi:10.1556/034.61.2019.3-4.5. https://www.researchgate.net/profile/Halda_J_P/publication/336562131.
* Linkola, K. (1929). "Edvard August Vainio" (in Finnish). Luonnon Ystävä 33 (5): 1–16.
* Linkola, K. (1934). "Edvard August Vainio 1853–1929". Acta Societas pro Fauna et Flora Fennica 57 (3): 1–26. https://helda.helsinki.fi/bitstream/handle/10138/15820/1934ActasocFFF573.pdf?sequence=1.
* Lücking, Robert (1997). "Estado actual de las investigaciones sobre líquenes foliícolas en la región Neotrópica, con un análisis biogeográfico preliminar" (in Spanish). Bryophyte Diversity and Evolution 13 (1): 87–114. doi:10.11646/bde.13.1.10. https://pdfs.semanticscholar.org/cf44/f06592bf3d9abf6f777fb017a6947e41a5e9.pdf?_ga=2.48592678.1858997801.1594701123-1996427379.1585886003.
* Magnusson, A.H. (1930). "Edvard August Vainio (1853–1929)". Annales de Cryptogamie exotique 3: 5–12. https://bibdigital.rjb.csic.es/viewer/11227/?offset=#page=6&viewer=picture&o=bookmark&n=0&q=.
* Recollecting Edvard August Vainio. São Paulo: CETESB – Companhia Ambiental do Estado de São Paulo. 1998. pp. 1–188. ISBN 951-642-878-9. OCLC 42766193.

Alava, Reino (1998). "Edvard August Vainio (1853–1929)". Recollecting Edvard August Vainio. São Paulo: CETESB – Companhia Ambiental do Estado de São Paulo. pp. 1–14.
Vitikainen, Orvo (1998). "E. A. Vainio – life and lichenological significance". Recollecting Edvard August Vainio. São Paulo: CETESB – Companhia Ambiental do Estado de São Paulo. pp. 15–28.
Stenroos, Soili (1998). "Vainio collections – TUR-V". Recollecting Edvard August Vainio. São Paulo: CETESB – Companhia Ambiental do Estado de São Paulo. pp. 29–31.
Marcelli, M. P. (1998). "The Caraça History and Importance". Recollecting Edvard August Vainio. São Paulo: CETESB – Companhia Ambiental do Estado de São Paulo. pp. 33–36.
Ahti, Teuvo (1998). "E. A. Vainio and his journey to Brazil, with notes on the Cladoniaceae". Recollecting Edvard August Vainio. São Paulo: CETESB – Companhia Ambiental do Estado de São Paulo. pp. 37–46.
Feuerer, Tassilo (1998). "E.A. Vainio's contribution to the knowledge of the Parmeliaceae". Recollecting Edvard August Vainio. São Paulo: CETESB – Companhia Ambiental do Estado de São Paulo. pp. 47–60.
Galloway, David J. (1998). "Edvard Vainio and the family Lobariaceae, with special reference to the taxonomic history of Sticta". Recollecting Edvard August Vainio. São Paulo: CETESB – Companhia Ambiental do Estado de São Paulo. pp. 61–84.
Yoshimura, Isao (1998). "Vainio and Lobaria, old and modern concepts". Recollecting Edvard August Vainio. São Paulo: CETESB – Companhia Ambiental do Estado de São Paulo. pp. 85–94.
Tibell, Leif (1998). "Vainio's ideas on the classification of calicioid lichens". Recollecting Edvard August Vainio. São Paulo: CETESB – Companhia Ambiental do Estado de São Paulo. pp. 95–112.

* Mattick, Fritz (1956). "Auf den Spuren des Lichenologen Wainio in Brasilien: Das Carassa-Gebirge" (in German). Willdenowia 1 (3): 404–432. https://www.jstor.org/stable/pdf/3995199.pdf.
* Mitchell, M. E. (2015). "Autonomy’s long shadow: A report on issues concerning lichen classification, 1870 to 1981". Huntia 15 (2): 87–104. https://www.huntbotanical.org/admin/uploads/03hibd-huntia-15-2-pp87-104.pdf.
* Schulz-Korth, Karl (1930). "Nachruf auf E. A. Vainio" (in German). Hedwigia 70: 1–9. https://www.zobodat.at/pdf/Hedwigia_70_1930_0001-0009.pdf.
* Tarmio, Timo (2000). "Edvard Vainio – Kohtalona Jäkälät" (in Finnish). Päin nousevan Suomen rantaa. Tutkijaprofiileja Turun yliopistosta. Turku: Kirja-Aurora; University of Turku. pp. 155–177.