Lichenotheca Veneta

Originally published in Nimis P.L., 2016. The Lichens of Italy. A Second Annotated Catalogue. EUT, Trieste, 739 pp.

Lichens were collected and illustrated by many authors during the Italian Renaissance, e.g. by Ferrante Imperato (1525-1615, Roca & Nimis 1997), Fabio Colonna (1567-1640, Roca & al. 1998), Paolo Boccone (1633-1704, Roca & Nimis 2002), and Federico Cesi (1585-1630, Nimis & Zucconi 2006). The latter, who for the first time used the microscope constructed by Galileo Galilei, published the first clear illustration of soredia, paving the way to Pier Antonio Micheli (1679-1737), who is considered by many as the founder of scientific Mycology, and perhaps the true “Father of Lichenology”. Born into a poor family in Florence, Micheli was apprenticed to a bookseller in Florence at an early age and was not able to afford a formal education, although he was able to teach himself Latin and study Botany. In spite of his lack of a degree, in 1706 he was appointed to a position as a botanist to Grand Duke Cosimo III de' Medici, Duke of Tuscany, with the responsibility for the public gardens of Florence. Micheli was influential in founding, in 1716, the Società Botanica Fiorentina, most likely the world’s first botanical society. His major work, Nova Plantarum Genera Juxta Tournafortii Methodum Disposita (Micheli 1729) dealt with about 1900 species, including some 900 fungi and lichens, illustrated on 73 plates. Micheli was the first to observe and describe fungal spores as reproductive bodies, to describe asci, and to culture fungi from spores.
After Micheli, there was no significant contribution to Lichenology by Italian authors until the first half of the XIX Century. However, towards the middle of the XIX century, immediately before the unification of Italy, the study of “cryptogams”, especially lichens, underwent a sudden moment of blooming. This phenomenon affects, more or less at the same time, most of the countries of northern and central Europe; in Italy, however, it took on an unusual extent. In a period of about 15 years, from 1846 (the year of publication of the Frammenti Lichenografici by G. De Notaris) to 1860 (death of A. Massalongo), Italy became the main center of Lichenology worldwide, a position perhaps never achieved by this country in the field of Botany.
After the period of the Napoleonic wars, the economic situation of most of Europe, including northern Italy, underwent a marked improvement, due to the progressive expansion of the industrial revolution. Enlightenment considered Natural Sciences as an indispensable element of the culture of any person: many encyclopedists cultivated botanical studies as part of their cultural interests, and the scientific culture was expanding under the impetus of the political and economic upheavals of the French Revolution and the subsequent Napoleonic campaigns. The revolutions of 1848 were the expression of the inadequacy of the old political-economic system compared to the new needs of the rising bourgeoisie. In Italy these developments were hampered by the persistence of the old humanistic literary tradition: they were felt mainly in the North of the country, that was closest to the political and cultural developments in the rest of Europe (Poelt 1991). Moreover, in the first half of the XIX century Botany was an integral part of the curriculum of studies of physicians and pharmacists, by themselves emblematic representatives of the increasingly prosperous middle class. No wonder that in this period many of the greatest botanists were physicians, pharmacists, priests, or offsprings from noble families. However, the sudden bloom of cryptogamic studies cannot be attributed only to cultural or economic causes. In particular, it is difficult to explain on this basis alone the leading position assumed by Italy, an area that, in both economic and cultural terms, was lagging behind other European countries. In fact, the main reason lies elsewhere: the sudden flourishing of cryptogamic studies in Italy around the middle of the XIX Century is mainly due to technical developments.
The first important lichenological system, that of the Swedish lichenologist E. Acharius (1757-1819), was mainly based on macroscopic characters, and with hindsight very artificial. It was only in the 1840s that microscopical characters, especially those regarding spore colour and septation, were increasingly adopted as paramountly important taxonomic criteria; the new emphasis on spores resulted in a revolution of previous taxonomic schemes, exactly as it is happening today with molecular data. The technical development which revealed a wealth of new characters for defining more natural groups was the invention of a new microscope with acromatic lenses by Giovanni Battista Amici (1786-1862), which allowed a much more detailed investigation of microscopical characters (Nimis 1988, 1993, Nimis & Bartoli 1992). Amici was the foremost Italian optical scientific instrument maker of the 19th century and one of the leading figures of this period at the international level. He applied the hemispherical front lens to the microscope objectglass (1838), and introduced the technique of immersion in water (1847) and in various types of oil (1855). Between 1857 and 1860 he invented the direct vision prism which continues to be used in spectroscopy and still bears his name. A very first version of the new microscope was produced in 1827, and the instrument was available on the Italian market between 1830 and 1840. Italian botanists were the first to have the opportunity to acquire it, which opened a new world ripe for exploration by the astute observer.
That all species of a natural genus should have the same type of spores had already been stated in 1837 by the eminent French cryptogamist A.L.P. Feé (1789-1874). Many of Feé's contemporaries in lichenology, however, objected to this thesis as with the microscopes then generally available the observation of spore characters was considered too difficult for practical use. Feé soon abandoned lichenology for pteridology. Starting from 1846, however, there was a true explosion of lichenological studies by Italian botanists, where the use of the microscope played a major role. The Italians G. De Notaris (1805-1877), and A.B. Massalongo (1824-1860), both now recognised as of world stature in lichenology, worked with Amici's microscope. In conclusion, the prominent position briefly taken by Italy was due to the fact that fundamental technical progress was first achieved in this country. This explanation does not want to detract from the merit of Italian scholars of that time, but emphasizes the fact that the history of science cannot be reduced to a mere sequence of individual stories.
The main protagonists of the “Golden Period” of Italian Lichenology were Giuseppe De Notaris (1805-1877), Abramo Bartolomeo Massalongo (1824-1860), Martino Anzi (1812-1881), Vittore Trevisan di San Leon (1818-1897), and Francesco Baglietto (1826-1916). De Notaris, Massalongo and Trevisan were primarily interested in Systematics: the old classification schemes dating back to Acharius, based on macroscopical characters, were completely revolutionised by the use of microscopical characters, such as shape, colour and size of the spores, and the microstructure of ascocarps. The international importance of these studies was remarkable, and caused a series of often fierce discussions, which involved the major lichenologist of the time.
The figure of De Notaris has a clear position as a pioneer and forerunner: already in 1867 in his History of Lichenology, Krempelhuber (1867) subdivided it into six major periods, of which the fifth (1801-1845) was called from Acharius to De Notaris, thereby stressing the revolutionary character of the work of the great Italian botanist. De Notaris can be considered as the founder of a new period in the history of ascomycete classification as a whole, and not only of the lichenforming species. In his vast scientific production, articles on lichens are a numerically small portion. The same De Notaris said, with his usual modesty, that he used to deal with Lichenology “in the hours of leisure” (Nimis & Bartoli 1992). His lichenological work consists in a dozen publications, only one of which (De Notaris 1846) would have sufficed to grant him a key place in the development of lichenology. Referring to the statements of Fée on the importance of sporological characters for a natural classification of lichens, De Notaris analysed and accurately described the anatomy of sixty species. Starting from the observation that similar species are found in most genera which appear clearly distinguished on the basis of macroscopic characters, he came to the conclusion that those genera which are macroscopically similar, but substantially different in sporological characters are not natural. Therefore, he suggested the possibility of creating a much more natural classificatory system by utilizing, in order of importance: (a) spore characters; (b) structure of the ascomata; and (c) thallus morphology. De Notaris' papers had an enormous influence throughout Europe, and his basic ideas were applied and developed with extraordinary intensity by A. Massalongo, certainly the most outstanding of all Italian lichenologists.
In just eleven years, Massalongo produced an impressive series of papers, some issued posthumously, where the taxonomy of lichens was drastically altered on the basis of microscopical characters, chiefly, but not only, those of the spores (Poelt 1991). A typical example is his interest in the so-called “blasteniospore lichens”, i.e. those with widely different growth-forms and appearance, which share the typical polar-diblastic spores of what is today recognised as the family Teloschistaceae. The Synopsis Lichenum Blasteniosporum (Massalongo 1852) was a bold attempt to recognise the affinity of these lichens and to arrange them into more natural genera, most of which were almost completely forgotten after Massalongo’s death, when hundreds of species were placed into three main, very artificial genera, mainly defined by growth-form: Caloplaca (crustose), Xanthoria (foliose) and Teloschistes (fruticose). Today the molecular taxonomy of Teloschistaceae is in full swing, and the recent treatment by Arup & al. (2013), where 39 genera are recognised, has resurrected from oblivion some Massalongian generic names, such as Blastenia, Gyalolechia, Pyrenodesmia, and Xanthocarpia.
During his short life, Massalongo had to fight to defend his ideas, especially against Nylander, but also against other Italian lichenologists - including Vittore Trevisan di San Leon. While Trevisan accepted the taxonomic importance of spore characters, he was often in conflict with Massalongo in the application of such principles and simultaneously investigated the taxonomical arrangement of several groups. The greatest part of the lichenological papers of Trevisan was published between 1853 and 1869. The publication of Massalongo's fundamental Ricerche sull'Autonomia dei Licheni Crostosi (Massalongo 1852) was probably the main stimulus to Trevisan's concentration on lichenological papers in the early 1850s. In the following months, Trevisan hastily published 7 lichenological papers. It is difficult to understand the effect that Massalongo's papers produced on Trevisan, without knowing that in the previous years he had intensively worked on a new synopsis of lichenised genera, in which the new sporological ideas were taken up.
The publication of Massalongo's work, whose importance he could not deny, anticipated some of the new genera he wanted to describe, and compelled him to revise his previous ideas, to adopt a critical position against several of Massalongo's concepts, and above all to publish as soon as possible what he had worked out until that time, without having the possibility of rounding up the whole, as he probably had wished. This situation led to serious misunderstandings between the two lichenologists (Nimis & Hawksworth 1995).
During 1853 and 1854, Massalongo's lichenological activity exploded in a series of important papers which brought about a true revolution in the generic arrangement of lichenised fungi. In the introduction to one of these fundamental contributions, the Memorie Lichenografiche, Massalongo (1853) provided a detailed response to Trevisan's former criticism. First, he expressed his disagreement on the relative importance of characters for taxonomic purposes: according to Massalongo, Trevisan underestimated the importance of thalline characters, the size of spores, and the structure and genesis of the apothecia. These considerations were illustrated by means of a decided defence of some Massalongian genera that had not been accepted by Trevisan. Finally, Massalongo tried to demolish many genera proposed by Trevisan, either because they were very poorly characterised, or because they were too heterogeneous. It must be recognised that much of Massalongo's criticism seems to be fully justified today. The Veronese lichenologist was a much more acute scientist than his Paduan colleague; Trevisan continuously strove towards a synthesis, but had the misfortune to live in a period in which analytical work was much more important and productive.
In 1860, the year of Massalongo's demise, Trevisan published what is perhaps the most important of his works today, a general conspectus of pyrenocarpic lichens, which also deals rather fully with the lichenicolous species known at that time. The Conspectus Verrucarinarum (Trevisan 1860) is a typical example of Trevisan's style: the text is extremely concise, being limited to the presentation of a taxonomic conspectus with the main characters of the accepted taxa, the main synonymies, nomenclatural information, and numerous telegraphically presented new combinations. Hidden in the dense smaller-typed text are nomenclatural details all too frequently overlooked.
Massalongo and Trevisan followed similar principles and were members of the same school. However, their scientific attitudes were quite different. Massalongo was a powerful analytical spirit, whereas Trevisan had a clear tendency towards synthesis and the correction of the historical record. Almost all his lichenological papers show a continuous effort to bring about clarity in a period characterised by a confusing flow of new information deriving from the developments of the sporological school. From carefully examining his lichenological papers one has the impression that his contribution to Lichenology would have been much greater if he could have published his ideas a few years before the “Massalongian” period of 1852-1860, and if he had not become so preoccupied with what he perceived as putting the past into order (Nimis & Hawksworth 1995). Unfortunately for him, the activity of Massalongo thwarted his plans, and his concept of an all-embracing classificatory system was reduced to a scattered series of hastily published fragments in need of continuous re-building and adjusting after the appearance of every Massalongian paper. Nevertheless, Trevisan's system, although published in a fragmentary form, constitutes one of the last examples of a general taxonomic arrangement of lichenised and lichenicolous fungi based on microscopical characters which appeared in the last century.
After the death of Massalongo, the interests of the main Italian lichenologists moved toward the floristic study of the territory, with the important studies of M. Anzi, F. Baglietto and A. Carestia (1825-1908). The excellent work of these lichenologists aroused some international attention mainly because of the distribution, in exsiccata, of the many new species that were gradually described, but this was not comparable to that caused by the publications of De Notaris, Massalongo and Trevisan. During the second half of the Century, the crisis sharpened quickly: already at the turn of the XX century Lichenology in Italy was virtually extinct (Nimis 1988). The life of the Società Crittogamologica Italiana was short-lived: the publication of the Atti ceased in 1868, while in 1872 the distribution of the Erbario Crittogamico ceased as well. The attempt to revive the Association, in 1878, failed, and in 1885 it was again virtually extinguished. Towards the end of the XIX century Italian Lichenology was represented mainly by Antonio Jatta (1853-1912), a wealthy landowner from southern Italy who began a meritorious work of synthesis that culminated in the publication of the part devoted to lichens in Flora Italica Cryptogama (Jatta 1900-1909). This work is undoubtedly laudable, but would have required lasting improvement by a new generation of lichenologist. Unfortunately, at that time, Lichenology could be considered as extinct in Italian universities. The rapid decline of Lichenology in Italy cannot be attributed solely to the disappearance of three outstanding personalities such as De Notaris, Massalongo and Trevisan. It is evident that it was decisively influenced by the unification of Italy, and the resulting profound changes in university policy of the new Governments (Nimis 1993). The new State had to face a series of difficult economic problems, including the restructuring of the agricultural system. Frequent outbreaks of pathogenic fungi in the second half of the XIX century further aggravated the situation. Botany was increasingly seen as an applied science, following the developments of late XIX Century positivism, which was increasingly influenced by the impressive progress of the industrial sector. Taxonomy, in particular, started to be seen as a “science of the second category”, something comparable to the activity of petulant stamp collectors, and appeared as obsolete and of little use when compared to the progress of plant physiology and the need to acquire detailed information on the biology of pathogens. After the unification of Italy, the university system underwent drastic reform. Botany, in particular, previously included in the Faculty of Medicine, was generally transferred to the Faculty of Sciences, with the creation of several new positions of full professor (Nimis 1988). The results of the new policy were disastrous for the Italian lichenological school: only De Notaris managed to become full professor, but only at a very old age, and his last years at the University of Rome were rather bitter for him. He was honoured as a great Master of Botany, but remained substantially isolated from the scientific world, and was left without means for carrying out his researches (Nimis & Bartoli 1992).
Very different was the fate of another prominent Italian cryptogamologist, a contemporary of De Notaris, Santo Garovaglio (1805-1882). He worked thoroughly in Lichenology before the publication of the works of De Notaris and Massalongo, but after the unification of the country, in 1869, he launched the idea of establishing a laboratory in Pavia specialised in fighting diseases caused by parasitic fungi. This captivated the confidence of the Ministry of Agriculture and of the administrative authorities of Pavia, and the Laboratory, which had a long period of deserved glory, was founded in 1871 (Nimis 1993). The last important work by Garovaglio devoted to lichens, the distribution of the Lichenes Langobardiae Exsiccati, dates back to 1864. In Rome, something similar happened a few years after the death of De Notaris: his student Giuseppe Cuboni (1852-1920), in the new cultural atmosphere, was appointed as director of the Royal Experimental Station of Plant Pathology of Rome, with the creation of a large experimental field, while the new Botanical Garden of Panisperna, promised to poor De Notaris for years, failed to see the light due to some gardeners that the authorities were unable, or unwilling, to dislodge from the ground that should host it (Graniti 1989). The political misfortunes of Taxonomy meant that none of the great Italian lichenologists honorably managed to fit in the new university system: some of them, being nobles or priests, were entirely unrelated to the academic environment, while those who had already entered into universities, as F. Zanfrognini in Modena and F. Baglietto, who was assistant to De Notaris in Genoa, were unable to advance in their careers, leaving no school. The Flora Italica Cryptogama (Jatta 1909-1911) appears today not as a new starting point, but as a conclusive work, a sort of gravestone lying on the “Golden Period” of Italian Lichenology, which was brought to almost complete extinction over a very short time as a result of a changed political, economic and cultural climate.
From a screening of the lichenological literature of the first half of the XX century, one is impressed by the high number of authors who published a few articles on lichens at the beginning of their careers, and suddenly abandoned this field. This is probably a consequence of the disappearance of a true lichenological school, and of the difficulties found by young botanists to pursue their lichenological studies. The decadence of Italian Lichenology is evident in the scarcity of really important figures throughout this period. Four main lichenologists can be mentioned: Eva Mameli Calvino (1886-1978), Maria Cengia-Sambo (1888-1939), Camillo Sbarbaro (1888-1967) and Ruggero Tommaselli (1920-1982). The only threads connecting the old lichenological tradition with the more recent years can be identified in the fact that Sbarbaro was introduced to lichenology by G. Gresino (1859-1946), a priest who was himself in contact with Baglietto, who died in 1916 at the age of 90 years, while Eva Mameli-Calvino, in the first part of her career, worked at the Laboratorio Crittogamico of the University of Pavia founded in 1871 by Garovaglio.
Eva Mameli Calvino, who incidentally was the mother of the famous writer Italo Calvino, was the first Italian woman to become university professor, first at the University of Catania then at that of Cagliari in her native Sardegna. She was introduced to Lichenology at the Cryptogamic Laboratory of the University of Pavia, where she worked as assistant to the Director Giovanni Briosi (1846-1919). In the first part of her career she published a dozen papers devoted to lichens, not only from various parts of Italy, but also from the new Italian colonies in northern Africa and Eritrea, after which she switched her interests to genetics and phytopathology applied to ornamental plants.
Maria Cengia-Sambo was a school teacher who soon became involved in the study of lichens: she was working first at Urbino, later in Florence, and collected extensively, especially in the Italian Alps; in the last years she also published some papers on extra-European lichens. The main weak point of Cengia-Sambo was probably her isolation: she was completely alone, as a lichenologist, within the Italian academic world, and had the misfortune to produce much of her work in the very difficult period following the I World War; she also never succeeded in obtaining a position within a university, which made still more difficult the contact with prominent foreign lichenologists. She had a keen interest in lichen ecology, and many of her numerous papers contain interesting observations; some of her lichen records, however, appear rather dubious. Camillo Sbarbaro as well was completely isolated from the academic world, and, as with Cengia-Sambo, he left no school. Sbarbaro was a very interesting personality, and presently is considered among the classics of modern Italian poetry; his interest in lichens was mainly aesthetic, and some of his prose writings devoted to these organisms would be worthy of being translated into other languages (see e.g. Knowles 2000). Although not a specialist, he had a very keen eye, and assembled an important herbarium, which, for lack of money, he was forced to sell, at least in part, to foreign institutions. Unlike Cengia-Sambo, however, Sbarbaro was in close contact with several foreign lichenologists, to whom he sent most of his material for identification.
For this reason, the lichens collected by him are among the few Italian collections of the first half of the XX Century which are cited in modern monographs. The few scientific papers published by Sbarbaro, which summarize the results of his investigations, mainly in Liguria and in northern Tuscany, are the best floristic contributions concerning Italy which appeared in this period.
Until the ‘80s, the only lichenological activity carried out within a university was that of Ruggero Tomaselli, an eminent, very versatile botanist, who for a period was also President of the Italian Botanical Society. The lichenological production of Tomaselli, however, cannot be considered as important for the progress of this discipline. He published very little on lichen floristics, and left no relevant school in Lichenology; during the last years of his life, Lichenology was virtually extinguished in the Italian universities.
Abroad, however, the situation was very different, and, starting from the end of the II World War, Lichenology experienced an extraordinarily intense growth in several European countries. The effects of this situation were also felt in Italian Universities, and in the 70's several young Italian botanists started to get involved in the study of lichens. A first course of Lichenology was organised at Trieste in 1986, attended by 30 persons from all parts of Italy. On that occasion, it was decided to found an Italian Lichen Society (Società Lichenologica ltaliana), whose first meeting was held at Trieste in 1987. A great interest was aroused, at the national level, by the activities of the Society in the fields of air pollution monitoring with lichens, and of lichens and monuments (see e.g. Nimis 1991). Presently, the Society continues an intense activity including courses, excursions, and the publication of a Notiziario. The latest developments are too recent to be treated here under a historical perspective. They are somehow summarised in the list of references of this book, which mostly covers the period between 1993 and 2016. More than 1000 papers were produced in this period by Italian lichenologists, devoted to widely different aspects of lichen biology, with a high number of papers in international journals, especially in the fields of lichen ecology and physiology.