Mushrooms: a very special place… in the plant kingdom

1Before the development of mycology as a recognized science in the 19th century, mushrooms were most frequently studied with plants, though naturalists since Antiquity had noticed and emphasized their very unique nature.

2The discourse found in the voluminous herbariums of the Renaissance remained heavily influenced by the writings of naturalists from Antiquity. Theophraste’s (ca 372ca 288 B.C.) opinion that mushrooms are plants devoid of their main parts (roots, leaves, seeds, etc.) was usually borrowed and expanded with various theories about the growth of these organisms. The main theory was that mushrooms were born spontaneously from humidity, by effect of putrefaction of various organic matters, especially vegetal. They were, as the neo-platonic Porphyre would call them, “the children of the Gods”. The mushroom was considered for a long time as either the most mysterious production of nature or as a seedless plant.

3Thus, according to Gaspard Bauhin (1560-1624), who restated almost word for word Jerome Bock’s opinion (1498-1554), “mushrooms are neither plants, nor roots, flowers, or seeds, but only humidity coming from the soil, the trees, dead wood and other rotten elements; their appearance and disappearance last approximately seven days, and they appear usually before thunder and rain.” The theory of the thunderstorm as generator of mushrooms (truffles in particular) is a classical theory of Antiquity (Pliny, Plutarch). This theory based on mystery, the secret forces of the Cosmos and of the “ex nihilo” shaped a real trend of thought in the history of mycology. Until the 18th century, the idea of the spontaneous production of mushrooms was largely regarded as true. In 1665, Robert Hooke (1635-1703) who had made many microscopic discoveries supported however this theory. In the 18th century, Carl von Linné (1707-1778) accepted Otto von Münchhausen’s (1716-1744) theories related to the spontaneous production of mushrooms, and Friedrich Kasimir Medicus (1736-1808) noted in 1788 that mushrooms are “the products of the jelly that results from the organic decay of leaves under the effect of heat and humidity.” The idea was in the line of thought of the ancient roman naturalists. But Johann Samuel Traugott Frenzel (1746-1807) used a style that was rather pre-Socratic when he wrote that “the shooting stars pull mushrooms from the earth” (Wit 1994, p. 131). Meanwhile, Luigi Ferdinando Marsigli (1658-1730) studied in 1714 the question of reproduction in mushrooms (Dissertatio de generatione fungorum) and came to the conclusion that the mycelium (the “white thread-like vegetative part of the fungus”) developed spontaneously on decayed matters, and that the mushroom itself (“carpophore” in modern language, which means “fruit bearer” in Greek) was the result of a metamorphic process of the mycelium. The main theory of spontaneous generation of organisms would be rejected only in the 19th century by Louis Pasteur. During this century, two authors fought this theory that was “rooted”, as we saw it, in Antiquity. These two authors were Christiaan Hendrik Persoon (1761-1836) and Elias Magnus Fries (17941878), who were the two most significant mycologists of all times. But their contribution to the knowledge of mushrooms resides more in their systematic and taxonomic work rather than in their thoughts on generation and reproduction of mushrooms.

4Bock and G. Bauhin excluded mushrooms from the plant kingdom in theory; however, they included them as part of their work that was devoted to plants. The presence of mushrooms in De plantis (1583) by Andrea Cesalpino (1519-1603) makes more sense. The author considered them as organisms between inanimate objects and plants, yet he defined them as “plants without seeds”. A few years later, at the beginning of the 17th century, Johannes Heckius (1576-ca1618) and Federico Cesi (1585-1630) referred to “imperfect plants” (Plantae imperfectae), somewhat in the same line of thought, to talk about all things of nature for which they had no knowledge about their reproduction system. These beings, as well as the ferns and fossils, were then considered as “deficient” (deficientes), “mutilated” (mutilae) and “inordinate” (inordinatae)31. This traditional view that we could call “Theophrastic” of mushrooms as “plants that are not really plants” lasted until the 18th century. In his Methodus plantarum of 1682, John Ray (1627-1705) classified mushrooms in the “plantae imperfectae” group, which he defined as follows: “I call imperfect plants those that have neither flower nor seeds, or at least that seem to lack of them, since until now, neither flower nor seeds have been observed in them, which leads us to believe that they grow spontaneously.” In 1694, in his Eléments de botanique, Joseph Pitton de Tournefort included mushrooms in Group XVII of “plants for which we know of neither flowers nor seeds” (Pitton de Tournefort 1694, vol. 1, p. 438). His definition meets Ray’s definition: flowers and seeds are not known, which does not imply that they don’t exist…

5The history of a science, as we know it, rarely follows a constant pattern of continuity. Mycology illustrates very well the difficulty there is into chronologically outlining scientific progress. The main crossroad in the study of mushrooms was the discovery of spores by Pier Antonio Micheli in 1729 (Nova plantarum genera), which he considered similar to seeds in plants, as the fundamental element of reproduction. But starting in 1588, Giambattista della Porta (1535-1615) used the very first magnifying lenses to observe spores in mushrooms and started to sense that these organisms developed similarly to other plants, from a seed. He wrote in Phytognomonica: “We picked up very thin seeds, small and black, hidden in the elongated veins that go from the base to the cap. The seeds that fall are always fertile and germinate” (Porta 1588, p. 412)32. In 1601, Charles de l’Ecluse (1526-1609), also named Clusius, asserted that mushrooms developed from “seeds”. At the Academy of the Lynx-Eyed in Rome, Cesi extensively used the microscope that Galileo offered him in 1624. With this instrument, he hoped that he would be able to solve the mysteries of reproduction in mushrooms. Cesi caught sight of the spores and painted many microscopic elements that we can still admire in the extraordinary compendium of mycological plates that he left, in a “codex” which is preserved at the library of the Institute of France (Freedberg & Pegler 2005, vol. 1, pp. 26-27).

6The cases of Bock and G. Bauhin, who did not consider mushrooms as plants, remained isolated. Starting in the 16th century, most authors included mushrooms in the kingdom of plants. Thus, we can find in Matthias de l’Obel’s work (1538-1616), also named Lobelius (De Lobel 1581), clustered mushrooms, others with almost swiveling roots, or others with bulbs at the base of their stem. In 1674, Martin Lister compared very naturally the tubes of what we commonly call today “bolete” and “polypores” to the tubular inflorescence of some compositae (Asteraceae) and concluded that the tubes of mushrooms were both flowers and seeds. Micheli (1729) finally looked in the fertile parts of mushrooms (in particular on the crest of the gills) for the elements that were well known for the fructification of plants: stamen, pistils, anthers, stigmas (Lamy 2008, p. 148). However, the suspicion that mushrooms represented a distinctive group within the plant kingdom started to get stronger. Johann Jakob Dillen, more often called Dillenius (1684-1747), was the first naturalist to create, as early as 1719 (Catalogus plantarum sponte circa Gissam nascentium), a class specific to mushrooms, the “fungi”. It seems that at the end of the 18th century, some confusion remained as to what legitimate place to give to mushrooms. Opinions differed and almost all the traditions of thoughts were intertwined. Yet in 1783, Noël Joseph de Necker (1729-1793), in his Traité sur la mycitologie, asserted that mushrooms formed a separate kingdom, a fourth kingdom that he named “regnum mesymale” (Necker 1783, pp. 103-104). After the assertions of Necker, Pierre Bulliard (1742-1793) suggested that “mushrooms differ from the staminiferous plants” in spite of undeniable “similarities”. At the same time, Jean-Jacque Paulet (17401826) maintained that mushrooms belonged to the plant kingdom and that they reproduced through “seeds”, as with any other plants33. However, 19th century advances in biology, technology (improvement of microscopes, which became tools of systematic use), and scientific terminology ensured the promotion of “mycology” as a full-fledged science (Lamy 2008, p. 148). Joseph-Henri Léveillé’s research (1796-1870) highly contributed to the advent of “mycology”, a terminology that was actually proposed by Paulet as early as 1795.

7Until the 18th century, the status of mushrooms remained quite problematic and often paradoxical vis-a-vis plants. The difference in nature was usually not noticed. Yet this difference existed in practical terms in two areas.

• Before the modern era of microscope investigation, the herbarium of mushrooms did not seem very useful. While a dried plant presented all characters necessary to its determination, a dehydrated mushroom did no seem to give much information because it lost, in particular, most of its original shape. Some attempts at preserving it in alcohol or formalin were done but with not much success; thus the plate, preferably in color, became the herbarium specimen of choice.
• Furthermore, though plants had been since Antiquity an object of interest for health purposes (research of their medicinal virtues), mushrooms were also sought for both sensual appetite (consumption) and intellectual appetite (by the fascination mushrooms exerted on people, which still applies today). It is true that very early on mushrooms were considered for many practical applications, such as what was done with the extraction of dyes in plants. Jerome Bock repeated Dioscoride’s discoveries (ca 40- ca 90 A. C.), which included the preparation of tinder from a specific polypore, and also presented some of Bock’s own fabrication (Dörfelt & Heklau 1988, p. 35, 43).

8But what fascinated people the most was the toxicity of mushrooms. The possible healing effects of fungi were set aside and the fungi ranked with plants, for according to Paracelsus (1493-1541), mushrooms had no healing power because they were nothing more than degeneration of nature; “it must be” Paracelsus wrote, “a plant that is void of anything, that has no root, no trunk, it is a mistake of nature”, and questions arose about the true nature of the only mushroom that was used at that time in pharmacopoeia, the “Agaric” (Agaricum officinale) that Micheli introduced, not with the other Fungi, but with the Larch (Larix) on which it grew and of which it was thought to be a protuberance of internal origin. However, emetics are concocted (Dioscoride, Matthiole) and even antidotes are created, such as the ones made by Jean Bauhin (1541-1612) in his Historia plantarum universalis. From Pliny the Elder (23-79 A. C.) to Dioscoride, Galien (ca 130-201 or 216 A. C.) and Sterbeeck (1675), classifications occurred generally around the dichotomy of Edibles/Poisonous (edules/noxii) because of a lack of well defined systematic characters. Romans’love for mushrooms is well documented and emperors were particularly fond of the Oronge mushroom which they called “Boletus” and that we call today, in their remembrance, “Caesar’s mushroom” (Amanita caesarea). There were also a few famous poisonings attributed to mushrooms, such as the poisoning of Emperor Claudius (10 B.C.-54 A.C.) by his niece Agrippine; and even better —if we can say so— Bouddha’s poisoning in 480 B.C. The delicacy of mushrooms as a gourmet food was known since Antiquity. Exquisite and complex recipes were created very early on, and Sterbeeck in 1675 (Theatrum fungorum) talked at length on how to prepare or preserve these strange products from nature. He even created a dressing especially reserved to the tasting of mushrooms which he named “white mustard”.

9The mushroom, then, fascinates as much because it can be delicious as because it can be lethal. Yet the number of toxic plants (such as the poison hemlock, Conium maculatum) is far greater than the number of toxic mushrooms, and the ancients were already aware of this. But, in terms of toxicity, the mushroom has always fascinated people. Why? Probably simply because its texture and the variety of its smells and flavors makes it an ideal dish for human consumption; because contrarily to plants, it rots fast while spreading fetid odors which remind of death and putrefaction; and because perhaps there is, in this radical opposition between the “goods” and the “bads” some kind of Manichaeism that brings back to the most profound and basic religious beliefs: the “good”, the “bad”, and even more loaded with pathos: “Paradise”, “Hell”, “God”, “Satan”… Maybe it is thus possible to explain a very weird conviction of the 16th and beginning of the 17th centuries that stemmed from Paracelsus’ “signature theory”: the toxic nature of mushrooms, it was believed, expressed itself in their morphological characters (external). Clusius (1601) was convinced of it (thus his classification in “esculenti” and “perniciosi”), as did Johannes Heckius, following in the footsteps of the “master”34: “For Heckius, as for many of the exponents of the traditional forms of natural history, it was precisely the external signs that gave clues to the inner nature, and, even more, to the inner forces —the vis as well as the virtus— of things” (Freedberg & Pegler 2005, vol. 1, pp. 40-41). The bottom line is that the bad can be seen in some external characters of mushrooms as it could be seen on some people’s face. The young amanita phalloides, or death cap, wrapped in its volva was called “devil’s egg”. Lenz, in 1831, still called “satanas” a big bolete so colorful that it reminded hell. Numerous myths and superstitions have been linked to mushrooms, far more than to plants. Ancients believed that rusted or dissolved iron spread “suspicious qualities to mushrooms” (Paulet 1793, p. 504, note 16). During the Renaissance, a mushroom of rather insignificant aspect, the Polyporus tuberaster, was called in Italy Lapis lyncurius because it was believed that it originated from lynx urine “which, when freezing, became hard and was transformed into a mushroom root” (op. cit., p. 497, note 2). This edible polypore develops indeed from some kind of hard, black root called “sclerotium”, deeply buried within the woody substrate. Other beliefs included the aphrodisiac virtues of mushrooms (inevitably) and the famous “fairy rings” formed by some species, which, as explained by Bernard Duhem, “were believed to be bewitched traps where travelers lost their way after they dared passing through them, and Middle-Age priests exorcised them.” (Duhem 1992, p. 46)… Superstitions still exist today; for example some people still believe that a mushroom that becomes blue when touched or exposed to the air is toxic… or, inversely, that a mushroom that “looks good” can only be delicious, which leads to accidents as stupid as dramatic.

Mushrooms among plants: hegemony in plants illustrations of the 16th and 17th centuries

10Starting in the 16th century, artists were able to paint plants with an amazing realism. “The Large Turf” by Albrecht Dürer (1503) is one of the most famous examples of his artwork, but there are many other similar works that are equally as superb. We can, for example, admire the purple columbine that we owe to the talent of an anonymous German painter (1526) which was reproduced in Madeleine Pinault’s work, Le Peintre et l’histoire naturelle (Pinault-Sørensen 1990, p. 16). The “herbariums” by Otto Brunfels (1488-1534), Jerome Bock and Leonhart Fuchs (1501-1566) are deservedly famous for the superior quality of their engravings. Fuchs’ New Kreüterbuch, a colorized copy of which we can consult online (SICD, University of Strasbourg-cultural heritage on line), is sometimes considered as the best illustration work of the 16th century35. But as far as mushrooms are concerned, this is far from being the case. Of course there is Frederico Cesi’s compendium, preserved by the Italian patron of the arts Cassiano dal Pozzo (1588-1657) in which mushrooms are painted with an amazing vivacity; these productions, which were not meant to be published, are isolated cases in the history of mycology and deserve a completely unique place and critical treatment. That being said, the compendium was consulted by a few botanists, in particular Micheli. His plates give evidence of this positive influence (Freedberg & Pegler 2005, vol. 1, pp. 29-30). It wasn’t until the 18th century, in particular Jacob Christian Schaffer’s plates (1762), that truly vivid and realistic illustrations (which does not imply though a superiority in terms of scientific approach) began to appear.

11Thus, between the 16th and the end of the 18th centuries, mycological illustration which was since its beginnings included in botanical books such as the large “herbariums” or the anthologies from the Renaissance, or the floras of the 17th and 18th centuries, followed a lengthy process of maturation. It is probably in the 1546 edition of Jerome Bock’s Kraütter Büch that we can find the very first representation of mushrooms. Pietro Andrea Mattioli, also called “Matthiole” (1500-1577) did not describe or represent any more than what Bock did, and it is only with the works of Clusius, Lobelius, and Franciscus van Sterbeeck (1641-1693) that these mysterious organisms became the object of thorough attention. Sterbeeck’s Theatrum fungorum (1675), as its name indicates, is entirely dedicated to the study of mushrooms. In modern language, we could call this work the first “monograph” of its kind, though Clusius’s Fungorum in Pannonniis observatorum brevis historia published in Rarorium plantarum historia (1601) could be argued to be the first.

12Next to these “bibles” are some isolated publications which present mushrooms that appear, to our modern eye, as definitely wild-looking. For example, Jean-Jacques Paulet refers to the studies of Seger, Berniz, Breyne and Welch, all published between 1671 and 1673 in the Ephémérides des curieux de la Nature (Paulet 1793 pp. 109-113) and states: “From 1662 to 1675, the least accurate observations were made because they were distorted by the love of the fantastic […]. The Philosophical Transactions suffered somewhat from this trend which was typical of the century” (op. cit., p. 108). Georgius Seger showed in 1671 a “mushroom” that would become famous, the “fungus anthropomorphos”. The plate shows six men, roughly outlined, gathered in the hollow of a giant volva. If we look a little bit closer, we can see many starred branches which remind of the earthstar (Geastrum). Sterbeeck reproduced an exact copy of this fungus anthropomorphos (1675, pl. XXIX, fig. B), which demonstrates that perhaps, what we consider today as fantasies based upon myths and superstitions were at that time very serious topics of study. However, observations at that time were also not always done rigorously. Thus, the mushroom that Jacques Breyne described and drew in the Ephemeris was called “tiger milk” because it was said to grow from the milk of a tiger. Discovered in the sandy soil of China, it is reminiscent of “a large truffle from which a stemmed and capped mushroom pops out” (op. cit., p. 111). It could be a Battarraea, a kind of gasteromycete whose figure is quite unique. It wasn’t until the beginning of the 18th century that these kinds of isolated publications began to appear to be more in accordance with our current knowledge; this new wave included pieces such as Reaumur’s article on a teratological form of a red cage (1713) and two articles written by Antoine de Jussieu, the first one on the white saddle, also named “mushroom-lichen”, the second one on a small mushroom with a strong garlic smell (1728). All of these texts were illustrated by Aubriet’s artworks and published in the Mémoires de l’Académie Royale des Sciences de Paris.

Influence of vegetal attributes in the representation of mushrooms

13The study of the mycological illustrations of the 16th and 17th centuries shows that the plates on mushrooms usually lacked the artistic quality and the scientific accurate descriptions of the plants they were supposed to represent. Why is there such a gap?

14Until the 19th century, mushrooms were equated to plants, one way or the other; whether seedless plants, whole plants, or mysterious and isolated products of nature, mushrooms were featured in botanical books. Surprisingly enough, a few plants (plates 33 to 36) also appear in Sterbeeck’s Theatrum fungorum (1675), though this work was exclusively dedicated to mushrooms, if we go by its title. The “children of God” were not orphans. They were placed in the large plant family and therefore their attributes were the same as plants: mycelia are roots, volvas are bulbs, there are even corollas in some cases...

1° Roots

15The “Fungi vulgatissimi esculenti” engraved in Lobelius’ artwork (1581), with their stems adorned with beautiful roots that emerge from all around, are one of the pioneer representations. The engraving was then duplicated by Rembert Dodoens (1517-1585) in 1583 and by Clusius in 1601. But the representation of roots or rhizomorphs is not unique to Clusius. A large lepiota engraved in 1616 in Fabio Colonna’s book (15671640) (Fabii Columnae Lyncei Minus cognitarum rariorumque nostro coelo orientum stirpium ekphrasis…), shows a network of roots that is very close, at least in the intent, to what Lobelius illustrated. Quite a large number of mushrooms represented on the beautiful plates made by Sterbeeck (1675) show a stem extending from these “pseudo-roots”. Aubriet added this plant attribute to most mushrooms he painted for Antoine de Jussieu’s collection. But these kinds of swiveling roots are no longer what surprises us today in Lobelius’ representation; what strikes us today are the thin strands that emerge from a clod of earth that looks far too thick to be a true representation from direct observation. At that time, under the impulsion of Micheli’s work (1729), mushrooms were about to be included in the plant kingdom. The microscope revealed stigmata, anthers, stamens, and pistils: the mushroom was a plant. From then on, its representation no longer needed to show floral attributes and shapes, which maybe explains the excessive realism in the engraved illustrations published in the Nova plantarum genera. The bases of the stems, in particular, were not cluttered with “extra mycological” elements36. A century earlier, Joannes Loeselius (1607-1655) represented in his posthumous book Flora Prussia, sive Plantae in regno prussiae sponte nascente (1703) a nest fungus, a kind of mushroom in the shape of a bird-nest containing small eggs called “peridioles” (“fungus pyxioides seminifer” as indicated by the author), three specimens of which were represented with very long and tufted roots. This kind of representation, which today, in relation to our knowledge and our practices in mycology, surprises us at first, disappeared progressively during the 18th century. Dillenius was the first one to represent the famous “wood hedgehog” (Hydnum repandum) in his Catalogus plantarum sponte circa Gissam nascentium (1719). On the sketch (plate 1, p. 84), the compact mass of filaments that is drawn at the base of the stem is more reminiscent of the roots of a plant rather than a fungus mycelium. In 1739, Johan Wilhem Weinmann (1683-1741) still “adorned” the foot of what looks like a grey spotted amanita (Amanita spissa) out with many filaments (Phytanthoza iconographia, vol. II, pl. 525, a), in the perfect “Aubriet’s way”.

2° Bulbs

16Though there is a real “tradition” of root representation in mycological plates, there is no such “tradition” for bulbs. The bases of stems that look like bulbs (of Liliaceae for example) have been in fact very rare in mycological literature since the 16th century. We will only mention, in Lobelius’s Plantarum seu stirpium icons, the species of ringless lepiota that shows, at the base of its stem, a bulge in the shape of a bulb that is divided into three parts (1581, vol. 2, pl. 273). The drawing looks like a gland that would contain three testicles. This mushroom was reproduced in 1601 by Clusius (pl. CCXCIII), then in 1675 by Sterbeeck. Both authors actually offer illustrations that are usually superior in terms of accuracy. In the “codex”, compendium of watercolors painted at the end of the 16th century by the French painter Esaye Le Gillon under Clusius’ supervision, mushrooms were really represented the way they were, without any artifact. This collection inspired both Cesi and Sterbeeck immensely. Sometimes artists drew the base of the stems in a strange way, yet they did not represent them as bulbs of tulips (or orchids). In those representations that simplified the form, or “made it more geometric”, some parts did not really represent reality, but they did not negate it either. Thus, in plate 167 of the posthumous Traité des fougères de l’Amérique written by Charles Plumier (1705), the bases look like saucers or hollow cylinders. Whereas Micheli (1729) outlined very elegantly the volvas of the straw mushroom (volvariella volvacea) as well as the amanitas (tab. 76-77), Aubriet, in this artwork, appeared, if not really awkward, at least confused by preconceptions that hampered his spirit. Look, for example, at his illustration of the death cap (pl. 65): doesn’t it look like a tulip bulb? In this respect, while browsing through Sterbeeck’s plates (1675), one can only be surprised by a mushroom whose stem stands on a real pedestal (pl. 21, fig. H). It is no longer reminiscent of geometry, but, rather, of architecture. It is even more surprising to find such drawings in an artwork book whose illustrations are, generally speaking, very realist.

3° General presentation

17Generally speaking, mushrooms between the 16th and the first half of the 18th century often resembled flowers. Either the representation geared the mushroom towards a floral appearance, or the authors chose to illustrate species that naturally reminded the world of plants. In 1714, the Italian doctor and naturalist Giovanni Maria Lancisi (1654-1720) pointed out that some mushrooms seemed to be “covered with flowers”. The species of the West Indies drawn by Plumier look amazingly similar to flowers (look in particular at plate 168 in the Traité des fougères de l'Amérique) or lichens, when, in fact, the much more typical “stem and cap” species are abundant in the Caribbean. Besides, the distinction between lichens and mushrooms was probably not always easy to define. Thus the white saddle (Helvella crispa) that Aubriet drew for Antoine de Jussieu in 1728 (see Ms 92, pl. 21b. and 25) is also known as “Mushroom-Lichen” (Jussieu 1728a, pp. 268-272).

Triumph of engraving: recurrent and ordinary illustrations. The question of color

18With the advent of printing in the 15th century, engraved plates benefited from the largest distribution37. Wood engraving was the oldest technique, but copper engraving soon became more competitive. It is said that copper engraving appeared circa 1550, and according to Denis Lamy, “P. di Nobili would be the first one to use it in botany in the 1580’s” (Lamy 2008, p. 151). However, the plates in the first major artwork dedicated to mushrooms, Clusius’ Fungorum in Pannonniis observatorum brevia historia (included in his Rariorum plantarum historia, 1601) were wood engravings. It wasn’t until 1675, in Sterbeeck’s Theatrum fungorum, that copper engraving was used to introduce a large number of new mushrooms. In the 16th and 17th centuries, color was quasi nonexistent in books and only a very few colored copies exist. According to Jean-Marc Chatelain and Laurent Pinon, “engravings from Fuchs’ book38, finely executed and with very few shades, were meant to be painted. And it seems that some were even colored in the very studio of the printer. We also know of the existence of a painter’s studio associated with Christophe Plantin’s printing press in Antwerp: it is said that women acquired expertise in the coloring of botanical artworks”39. Yet, we don’t know of any “colored copy” of any book with mushrooms illustrations. However, we know for sure that Clusius’s compendium of eighty seven watercolors, usually called “Clusius’Codex” (Istvánffi, 1900), created at the end of the 16th century by the French miniaturist Esaye Le Gillon (Florike, Hoftijzer & Visser 2007; Backer et al. 1993), was not only useful to Clusius himself to illustrate his Fungorum in Pannonniis…, but also to Sterbeeck, who in 1672 had the immense chance to hold in his hands this “codex” that Clusius had lost. Unfortunately Federico Cosi was not so lucky and did not have the opportunity to see it; he would have to settle for the engraved plates from the Fungorum in Pannonniis40…, which would, however, be for him a very positive source of inspiration (the Clathrus ruber from “Cesi’s Codex” is directly issued from Clusius’ engraving).

19In this survey, it is very important to distinguish the works that were published in color. In terms of colored books on mushrooms, it seems that J. W. Weinmann’s Phytanthoza iconographia (vol. 2, 1739) was the first to be published. But J.C. Schäffer’s (1718-1790) Fungorum qui in Bavaria et Palatinatu circa Ratisbonam nascuntur (in 4 volumes), that was published starting in 1762, is far more famous (it was the first major monograph in color on mushrooms). It is so famous that this artwork is sometimes cited as being the first book, in the history of mycology, to present illustrations in color. Many more artworks in color followed in the footsteps of Schäffer, such as those by Johann Georg Karl Batsch (1761-1802), Elenchus fungorum (1783-1789), or Herbier de France (17801795) by Pierre Bulliard, or Couloured Figures of English Fungi or Mushrooms (17971805) by James Sowerby (1757-1822). In the 19th century, C. H. Persoon published several watercolors (in particular from 1803 to 1808 in Icones pictae specierum rariorum fungorum) and E. M. Fries published much later in Icones selectae hymenomycetum (18611884), as a final achievement to his immense work, very meticulous, if a bit rigid, colored illustrations painted by his students. Lucien Quélet, French mycologist, who had noticed the numerous errors of interpretation because of brief or incomplete descriptions, tried to include colored drawings in the 22 supplements of Champignons du Jura et des Vosges (1875-1902). We must also mention the extraordinary Icones Selectae Hymenomycetum by the Hungarian monk Károly Kalchbrenner (1878) who for the first time, drew numerous species from the mountains in large format as well as the caricatural drawings dashed off by the Bavarian Max Britzelmayr (from 1883 to 1905) in Hymenomyzeten aus Südbayern. Yet there was still at that time mycological floras ornamented with black and white engravings, such as Selecta fungorum carpologia by Charles and Louis-René Tulasne (1861-1865). At the very end of the 19th century (in 1899), Petter Adolf Karsten (1834-1917) published Finlands Basidsyampar—in the collection Floristiska handböker för nybegynnare (which means “floristic manual for beginners”) which also included black and white engravings. In the first quarter of the 20th century, color became more common. Some of the very last artists to resist color were the Czech Joseph Velenovský, České Houbý (1920-1922), and the French scientific mycologists, such as René Maire, Roger Heim and Robert Kühner, who preferred (for lack of time?) the quality of the writing and of microscopic drawings to the drawings of carpophores.

20It should be noted that, at that time, colors had a far more significant taxonomical value in mycology than in botany. Even if the color varied within a same species of mushroom, it was never disregarded, as often happened with plants. The botanist could determine the classification of almost all plants —up to their very final taxonomical rank-by using only morphological characters. For the systematic mycologist, organoleptic and chromatic characters cannot be ignored. Yet, was this data as critical for naturalists of the past who studied mushrooms? Though Sterbeeck (1675) attached the utmost importance to color for his groupings (Paulet 1793, p. 118), most authors from the end of the 17th and from the 18th century grouped mushrooms according to strictly morphological criteria. Before Sterbeeck, the primary division (when a classification occurred, which was not always the case) separated usually the “good” mushrooms (edible) from the bad ones (toxic). Distinction by means of color was used, however, by Micheli (1729), but only at the very end of the taxonomic division. The use of almost exclusively morphological characters makes sense since naturalists considered mushrooms to be real plants. When mushrooms were finally separated from the plant kingdom, characters specific to them were sought and defined in order to create a new mycological classification. Color then became very important and, subsequently, had a direct impact on the scientific value of illustrations, whether or not they were in color. In most cases, a black and white illustration of mushroom was worth far less than a black and white illustration of plant.

21Furthermore, what makes scientific illustration of the 16th and 17th centuries unique is the duplication of figures from one book to another41. The process was almost systematic in the 16th and 17th centuries. Annie Chassagne states that in “natural history, engraved woods were used from one edition to the other, transmitted from one studio to the other, and were used again, sometimes with some modifications, for books different from those for which they were initially designed” (Chassagne 2007, pp. 122-123). The creation of new plates implied a very complex job and was extremely expensive: “Hiring an artist and an engraver,” explains A. Chassagne, “whether for wooden or copper engravings, represented a very high expense. Thus, many treatises were not illustrated, or they used already existing engravings” (op. cit., p. 123). In botany, Madeleine Pinault-Sørensen mentions The Herball or General Historie of Plantes by John Gerarde (1597), which was a reference “herbarium” that used many former illustrations, such as the ones engraved in Jacobus Theodorus Tabernaemontanus’s (1525-1590) Neuw Kreuterbuch, published in 1588. In mycology, the illustrations that were reproduced the most were those by Clusius, which we find in particular in Sterbeeck’s books (1675). Sterbeeck was criticized for not citing his iconographic sources but this practice was rather rare at that time. However, the Flemish priest and naturalist did mention Lobelius, Dodoens, Clusius and the Bauhin brothers in his text, mostly written in Dutch. Clusius himself borrowed most figures presented by Lobelius in 1581 in his Plantarum seu stirpium icones, such as the morel, whose very first representation is owed to Lobelius. But he did offer a large number of new ones. Among the compilation artworks is Historia plantarum universalis (volume 3) by Jean Bauhin, posthumously published in 1651. The engravings on wood exhibit forty-two species of mushrooms, yet most of them are reproductions, sometimes awkwardly represented, of Lobelius and Clusius’ artwork. However, Jean Bauhin gave a rather good representation of the “yellow chanterelle” (p. 832, Cap. XXVII) and of an amanita (p. 826, Cap. III), which seems to be of the white forma of the citron amanita (Amanita citrina f. alba).

22Starting at the end of the 17th century, the tradition of compiling images started to slow down. Medieval and Renaissance traditionalists started to be forgotten, which left room for new illustrations. Thus the representations from Paolo Silvio Boccone (1633-1704) in his Museo di fisica (1697), those by Pier Antonio Micheli in Nova plantarum genera (1729), and those by Johann Gottlieb Gleditsch in Methodus fungorum (1753) are all original illustrations. Furthermore, the very fine engravings in Sébastien Vaillant’s Botanicon Parisiense (1727), based on Aubriet’s drawings, presented mushroom specimens that were not represented in any previous book. However, Tournefort’s plates in his work Elémens de botanique showed in three different instances some level of “archaism”: the morel and the red cage, which appear on the same plate (pl. 329) were a return to Lobelius and Clusius’ images; the truffles (pl. 333, vol. 3) grouped together, were very reminiscent of Matthiole’s 1571 representation (the first one in history), which was later taken up by Lobelius. But in the paintings of manuscript 92, there are no illustrations that would remind us of any previous representation—either “historical” or more contemporaneous.

23Since engravings were often reproduced from one book to another, some species were represented repeatedly. And we can notice that some mushrooms were more popular than others. Some were definitely favorites: the red cage (Clathrus ruber), the stinkhorn (Phallus, various species), the bird’s nests (in shape of tiny cuplike nests), the ramarias (Ramaria) and the clavarias (Clavaria), the morels, and the truffles. The red cage, though not that common, was the most depicted mushroom. Clusius represented it for the first time in 1601. It appeared afterwards in almost every single book and was still, in the 18th century, very popular. Plumier represented a very similar species endemic to the West Indies (1705, pl. 167, fig. H, Clathrus cripus). In 1729, Micheli had some magnificent copies of it engraved (tab. 93), and later, in 1753, Johann Gottlieb Gleditsch (1714-1786) represented it as well (Methodus fungorum…) with mucus dripping at the level of the volva (tab. IV). Aubriet represented it for Tournefort’s Elémens de botanique (1694) and in 1713 for an article by Réaumur (Ferchault de Réaumur 1713, pp. 71-76); however it is clear that Aubriet did not have first-hand experience with the specimen, since the Clathrus ruber presented there had such a monstrous shape that it looked like a “new species”. Aubriet did not paint the mushroom again; it appeared neither in Vaillant’s Botanicon Parisiense (1727), nor in the series of miniatures that we are presenting here.

24It is easy to imagine the success of the ramarias and of the clavarias: these mushrooms, growing in dense clusters, branching or cylindrical, are very reminiscent of plants. Persoon eloquently named a species of Ramaria “botrytis”, which means “cauliflower” (rosso coral). It is probably in Jacques Barrelier’s (1606-1673) posthumous artwork Plantae per Galliam, Hispaniam et Italiam observatae… (1714) that we can admire, among the ever popular Clathrus and Phallus, the biggest number of ramarias and clavarias. Mushrooms that present the most analogies with either the plant or the animal kingdom (the stinkhorn, the bird’s nests, and the truffles, which people in the past confused sometimes with deer testicles), have been the most popular in the history of mycological illustration. Maybe it was more natural for scholars of the 16th, 17th, and 18th centuries to include, in their botanical works, productions that could easily be perceived as “intermediary elements” between the animal and the plant kingdoms, rather than productions whose general appearance and unique characters were completely out of “the known”, of “the familiar”. As for corals, which were considered for a long time as plants, they include a few species in the fungus world whose appearance is very close to plants: the ramarias and the clavarias are sometimes reminiscent of them, as well as the morel which is, so to speak, a “land sponge”. This delicate vernal mushroom was repeatedly represented since the 16th century (and, until Persoon, grouped with common stinkhorn of the Phallus genus).

Discrepancies in the mycological illustration: reports and causes

25Thus, until the 18th century, illustrations of plants and of mycological plates (engravings or rare existing miniatures) showed an unequal level of quality and it is difficult to establish a relationship of familiarity42 (as per Bertrand Russell’s definition) between some of these illustrations and the mushrooms. A typical example is J.W. Weinmann’s work, Phytanthoza iconographia, which illustrates the best this historical “discrepancy” of mycological illustration with regard to botanical representation. Dated 1739, this work already showed a stark contrast between plants and mushrooms plates. Species were painted with rough outlines, their characters were misrepresented (the fly agaric, or fly amanita, for example, is represented without a ring), and their colors were rough and sometimes doubtful (see in particular plate 525b). Obviously, the author did not master any mycological knowledge, which was still, for the most part, in its embryonic stages. Thus he mixed up (plate 525c) the parasol mushroom (Macrolepiota procera) with the common stinkhorn (Phallus impudicus). In contrast, his plants are painted with a delicacy, precision, and beauty that reflect a thorough scientific knowledge and a deep “familiarity” with the plant world. This difference in how to master realism in the representation of plant and mycological specimens reflects the scientific knowledge of the time: as early as the second half of the 17th century, the botanical knowledge was already advanced, in particular thanks to the work of Ray, Pierre Magnol (1638-1715), and Tournefort. But in terms of mushrooms (as well as cryptogams, mosses, algae and lichens), no real strong scientific corpus would exist until the 19th century (C. H. Persoon and later E. M. Fries’s works). As for plants, the accurate knowledge of their parts, of their physiology, and the existence of a rigorous terminology enabled the objectivity of their perception and later, the realism of their representation. As for mushrooms, illustrators still fumbled. And illustrations suffered from it, since there is obviously a correlation between scientific knowledge and an accurate representation. In his book “Mémoires pour servir à l'histoire naturelle des animaux”, Claude Perrault (1613-1688) wrote that what mattered was “not so much to represent properly what we see but to see correctly what we want to represent.” In this respect, it took a very long time to “see correctly” mushrooms, because the mechanism of their reproduction was not understood, botanical terminology was used to designate mushrooms’ parts, and because the functions and interrelations between these parts remained completely unknown.

26Thus, often when something was not understood well, nobody would dare taking any risks with an original illustration and works were duplicated from one book to another. There is an exception in regards to Frederico Cesi who had a theory about plants and mushrooms in total contradiction with his very realistic “practice”. Indeed, he wrote, “there are lines in rushes, spheres in bulbs, fruits and tubers, the circumference of an oval at the top of a cabbage, and the look of a pyramid on the pointy and flowering stem of a bellflower”, and he said of mushrooms, “if you want to use a compass, go and see mushrooms”43. However, these mushrooms, under our eyes, are alive, something from their “core essence” is often conveyed to us through their depiction. Mushrooms were painted the way they were picked up: whole, broken, deformed, half rotten, etc. They were neither geometrically represented nor simplified. As far as we know, the trend of original designs in mycology started with Frederico Cesi’s collection44, which includes an impressive number of miniatures painted with gouache that illustrate hundreds of mushroom species. Amazingly realist for this period of the beginning of the 17th century, they are also distinguished historically by their uniqueness. Indeed, they were not meant to become engravings. Thus, each of them is a unique artwork in itself, whose purpose was only to be part of Cassiano dal Pozzo’s “Paper Museum” and, thus, to participate in the knowledge of the natural world through the irreplaceable medium of image. This collection, of such quality that we still praise it today, violently broke the historical continuum of the development in mycological illustration.