Footsteps: Systematics

Understanding and Explaining Plant Diversity

It is natural for humans to make sense of the world around them, which causes me to wonder how we comprehended circumstances before language. I mean, how did we keep track of objects without names, and even once we had internally applied some symbolic association, how did we communicate thoughts, or decide that two different objects could merit the same symbol? Like, are a rock and a pebble the same as a stone? And how was it possible to pass down knowledge about the world around you when there was no language or taxonomy. What is the story we feel compelled to understand, and to convey? What words and symbols are needed?

Taxonomy – studying “beings” and “things” in order to give them appropriate names and organize them in a system that makes sense to the user.

  • Thing – an inanimate object
  • Being – a sentient, living object

But we have to discern differences and patterns, we must practice taxonomy because it’s necessary to discriminate between different kinds of beings and things. Some people are born to do this, people who sort everything around them into explicit categories, screws and bolts for example. There are the ones with points, like wood screws and machine screws. And there are the ones that marry to nuts, and don’t need pointed tips. But some of the bolts have faceted heads that fit a wrench, while others have rounded heads with slots that require a screwdriver. Or maybe the head is fashioned for a Phillips screwdriver, or even an Allen wrench. So those have to be separated. The same-looking screw or bolt will exist in different gauges, 1/4, 1/8, 1/16th of an inch. Or, as we have all learned over the past two decades, the entire range of fasteners can exist in metric sizes, 6mm, 7mm, 8mm etc. And there are different kinds of threads, such as tightly wound machine threads that provide more grip versus conventional threads, that are still pretty good.

For those of us who love to mess with gadgets, the ability to find just the right bolt or screw is highly prized. A lot of tinkerers in the Pasadena area still mourn closing of the famous Berg Hardware. You could walk up to the counter with some random mechanism, and discover the staff had the very bolt you needed. “Yeah, we should have this. Do you want it in galvanized, or stainless? How long?” Then they’d disappear into the ranks of shelving and come back with a small, torn box, match the fastener, and ask how many you need. Sometimes, it was just one screw, or one bolt with matching nut and washer; less than a dollar. And you didn’t have to purchase a dozen, or go on-line and hope that the words you are using for the screw are the same ones deployed in some massive database. I don’t know how they stayed in business, but of course, the store closed, so perhaps I know why…..

The folks at Berg Hardware were taxonomists, and they were systematists. They had names for each minorly different fastener, and they had a system by which to pigeonhole them. I wish I had begged a visit behind the counter before the store closed, just to see how the thousands and thousands of fasteners were organized. Were the stainless ones on separate shelves, or were they interspersed with the same sized gizmos of other material? Were the nuts nearby, or over in their own area? What kind of shorthand evolved so that one staff member could most efficiently help another find their prey?

Well the world of plants is a bit, just a bit, like nuts and bolts. There are thousands of kinds (hundreds of thousands), and we even have places like Berg Hardware, herbaria with metal cabinets in which specimens of different plants are sorted, organized, and shelved. The world’s larger herbaria have millions of specimens, filling cubbyholes in thousands of cabinets.

The problem with plants, what makes them so very different from nuts and bolts, is that specimens collected from different individuals are not exactly the same. The closest you’ll come to a precise match would be in an herbarium that documents clones, like rose cultivars. Every ‘Altissimo’ basically should be fairly similar to every other ‘Altissimo’ sample, though each would be a unique cutting. Even if the specimens were similar cuttings, the growing conditions, stage of growth, treatment of the collection, etc. would vary, so you’d never get the perfect replicas we expect from nuts and bolts.

In naturally-occurring populations of every kind of plant, just as in a community of people, each individual differs from the other. Even though each of us is an individual, we have no problem deciding who else is a human; you just know a person when you see one. It’s part of being human, knowing your kind. But we are not as competent when faced with other kinds of organisms. We do not, inherently, know the range of variation of one plant species as compared to another. We don’t really even know how to define the limits of a species, when two similar-seeming plants are of the same kind, or different. The plants “know” (yes, this is pushing it), or in many cases, their pollinators, but we don’t. What’s worse, we do not really agree in every case how to define what separates one species from another, because different plant groups don’t share the same growth and reproductive strategies.

Scientists have come to a general consensus that if two related plants, when grown within breeding distance, are not capable of interbreeding naturally, then those two plants probably represent different species. That “biological species” concept had interesting consequences when Russian botanists studying dandelions (which are notorious for apomixis, i.e. establishing clonal colonies through generating cloned seed) decided to assign a formal scientific name to each separate colony.

But almost any good idea can be taken too far.

A more contemporary thought would be that we should be able to study the genetics of plant populations, and then decide through genetic evidence what makes one species different from another. The problems with this are many. Who has the time, money, staff, and facilities to make all of those studies? How do you draw lines (based on genetic makeup) when you don’t understand the way those genetics are expressed in nature, and how they impact populational behavior? What sense can be made of differences when there may not be clear expression of that difference in any discernible characteristic? How would you use such finely tuned differences in a practical sense, how would you group plants into functional or useful categories? How would a gardener, or a field biologist, know plants of one species from those of another?

But we are getting ahead of the story. Having delved into this topic, I have made my own sense out of historical approaches to plants, and the way science has come to portray the vegetable kingdom. Simply outlined, the story I relate here identifies several epochs in this saga:

  1. Folk Traditions describe the most ancient, regional ways differing cultures use, understood, and named the plants around them.
  2. Herbal Traditions remind us that plants were valued as a gift of Nature to humankind, and in that anthropocentric world, the key to understanding diversity would relate to how each plant was meant to serve us.
  3. Bringing Order reflects the dawning of discovery and globalization, when it became clear there are actually thousands of kinds of plants, and there must be a scientific basis to the differences we find.
  4. A Natural System explains the realization of botanists that documenting and categorizing the world’s plants would, in turn, provide a map to the basic plan of Nature.This was a 19th century attempt to explain the reason and relationships so evident in the world’s flora.
  5. Biosystematics relates post-Darwinian and post- Mendelian efforts to learn more about how the biology, the nature of plants, impacts speciation and diversity. Lacking direct access to the genetic code, 20th century scientists explored chromosomes, chemistry, and biologies to document the basis of variability and explain the origins or plant species.
  6. Phylogenetics came into its own at the end of the 20th century, when sequencing of RNA and DNA began to provide direct evidence as to genetic codes. Combining morphological, chemical, ecological, biological, and genetic clues, botanists began assembling stick figures, “trees” (cladograms) that attempt to map evolutionary origins of plants we know today.
  7. Comprehension, a new dawning, will come at some future stage, when botanists actually understand the roles genes play in the lives of plants in nature and in the garden. At that point, biologies will re-emerge as core, and we will come to understand the multivariate relationships between phenotype and genotype, nature and nurture. Almost everyone will be seen to have been right, Linnaeus, Jussieu, Darwin, Bessey, Hennig…, will have held a different part of the elephant.

Folk Traditions….

Once humans devised languages and systems of writing, the world changed, as words became symbols to record information, establish rules, name and track things, and encapsulate concepts. Names were given to plants of value, plants worth ongoing attention, because they were sources of food, flavoring, fuel, shelter, and fiber, even medicine. Each ancient society seems to have come up with names for a few hundred plants out of the thousands that surrounded their communities. The names reflected something about culture and use, with no need to reflect much about biology.

Shreds of traditional naming systems linger throughout the world, and folk names (or “common” names) for plants survive in daily language and commerce. But of the many plant naming traditions, only one survived as the basis for a bona fide system for giving plants botanically-useful names. That Greco-Roman tradition led to the European system which evolved as the world of scientific nomenclature.

“Nomenclature is a system of names or terms, and the rules for forming these terms in a particular field of arts or sciences….” Wikipedia (adapted)

It’s undeniable that scientific nomenclature and taxonomy are based in Greco-Roman philosophical traditions of the Mediterranean Basin (including Arabic scholarship) and northern Europe, which began with practical records, ancient compendia of information on useful plants. Some people are bothered by the Eurocentric nature of plant scientific naming conventions (animal names as well), but if it brings comfort, people should also know that norms governing plant nomenclature reflect continual oversight, review, and adjustment by an internationally-based organization of scientists native to all areas of the world. This collaboration has been on-going for decades, even during the now-forgotten Cold War, Russian, Cuban, and Chinese scientists participated in reviews and rule-making. Still, the Mediterranean roots (and social dominance of the Church) mean that science, which was derivative of Greek, Arabic, and Latin texts, clings to Latin as the lingua franca for plant and animal names.

Early botany was the province of medicine and agriculture, with Western accounts of plant identification driven through Greek and Roman discourse. Thus, many recognizable and durable plant names derive from ancient texts that retained or regained currency through nearly two millennia of cultural development, based on texts ascribed to Aristotle, Theophrastus, Pliny, Dioscorides, and a bit later, Galen.

Those texts, however, are far from singular. They were not carved in stone, and no “original” copies exist. Each has been filtered and even lost at some level, passing through Greek, Latin, and Arabic versions. Theophrastus, who is sometimes called the Father of Botany, provided information on approximately 400 different kinds of plants, information that was “re-discovered” in the Renaissance and still cited as valid knowledge by writers into the 18th century. Sources for this information are of variable quality, but the impact remains in that some of the earliest names for plants in Western culture are tied to peripatetic sources, i.e. Aristotle, Theophrastus, and Pliny (whose Historia extracted liberally from Theophrastus.)

Herbal Traditions…

Dioscorides’ de Materia Medica (around 70 AD) differs from other sources, however, in that for over 15 centuries this pharmacopoeia remained an actively utilized text, adopted and annotated through translation, transliteration, transcription, and reinterpretation by practitioners around the Mediterranean basin – eventually as far afield as Germany, France, England, and the Netherlands.

Singer (1927,) and Popa (2010) (see text box) explain some of the complex history of the Dioscorides record, differing manuscript versions showing annotations in varying languages, creating a scholarly tradition of study that was still active when printing emerged. After Gutenberg, several versions of Dioscorides were mass-produced, with Aldus’s 1499 Greek edition regarded as the most authoritative. The Aldus publication gained legs when Mattioli issued his Latin translation which hit the presses in 1554.

Invoking Dioscorides tells us more about a tradition than a person, speaking to a body of knowledge that survived and morphed through centuries as a basis of the herbal movement. There are scant records of the purported author’s life, and no contemporary materials that confirm a person named Dioscorides truly lived and breathed. But we envision a famous herbalist named Dioscorides, who authored scrolls that documented a Greek pharmacopoeia, which was copied in multiple iterations in Greek, as well as translated into Arabic and Latin. Passed down through loosely connected generations of knowledgeable specialists, experiencing annotations and changes, the Materia Medica was core to medical plant traditions (pharmacognosy) into the early Renaissance.

Ioana Claudia Popa, 2010. “The Lists of Plant Synonyms in De materia medica of Dioscorides” Global Journal of Science Frontier Research, Vol. 10 Issue 3 (Ver 1.0), July 2010

Charles Singer, 1927. “The Herbal in Antiquity and Its Transmission to Later Ages” Source: The Journal of Hellenic Studies, Vol. 47, Part 1 (1927), pp. 1-52 Published by: The Society for the Promotion of Hellenic Studies

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Formal training in medicine demanded familiarity with important treatises, which meant Dioscorides was standard knowledge for classically-trained doctors. As printing revolutionized the impact of text, availability to Dioscorides grew and its influence was given a bit more life due significantly to Mattioli’s Latin version, which became the basis for many regional herbals. That means the names and properties of plants mentioned in de Materia Medica stretched into the era of discovery and colonialization.

As a Mediterranean floristic document, practitioners realized the hallowed text could not fit a more northerly climate, or a world moving toward scientific revolution and empiricism.

No place was this more obvious than in Germany, where many plants described in Dioscorides were irrelevant. Moreover, introduction of exotic plants from other continents provided increased challenges, as interest in pure botany and horticulture grew.

It should be no surprise, therefore, that the more recent Herbal tradition (which endured about 250 years) began in Germany, with the beginning of Western printing in Mainz. The new genre spread throughout Europe, and became an extensive, iterative, and somewhat baffling body of work, encompassing a rich mixture of original, copied, translated, and modified texts, illustrated by a growing album of prints that ranged from graphic and nearly abstract to representationally natural and accurate. The early use of woodblock prints led to resale and reuse, as well as outright copying. Quality and accuracy (as well as size) varied from one publication to another.

Histories focus on the “German Fathers of Botany.” We celebrate Otto Brunfels for his early German herbals (1530-1537), but that celebrity relies more on the illustrations by artist Hans Weiditz than on the writing, which is reported to draw heavily (and un-expertly) from older texts, such as Pliny and Dioscorides. The situation shifted quickly when Hieronymus Bock published original descriptions (in German) for 700 plants in his 1539 Kreutterbuch. Just a few years later, in 1542, Leonhart Fuchs published De historia stirpium, which though reliant on text of other authors also broke new ground with fresh descriptions for native plants and for newly introduced exotics such as American corn and pumpkin. Most importantly, as with Brunfels, Fuch’s Stirpium was gloriously illustrated by artists who were credited: plant depictions were created by Albrecht Meyer, transferred to wood by Heinrich Füllmaurer, and then carved to woodblocks and printed by Vitus Rudolph Speckle. The images made an impact, which meant they were copied and reused extensively, even by artist David Kandel who illustrated the 1546 re-issued edition of Bock’s herbal.

As important as Brunfels, Fuchs, and Bock were, Rembert Dodoens 1554 herbal, Cruydeboeck (which included 714 plates) was the basis for a growing asset that matured in 1583 as the much expanded Latin version, Stirpium historiae pemptades sex. Significant botanists L’Obel and Clusius seem to have contributed to that maturing text, which in turn provided the basis for Clusius’ French translation. Dodoens, we learn, was one of the most widespread and translated texts, second (perhaps) only to the Bible. Thus, Clusius’ French translation is the basis for Lyte’s Historie…(1678) in English, and an English translation of the Latin Dodoens provides much of the text for Gerard’s Herball (1597), two of the most prominent English herbals in the second half of the 16th Century.

Curiously, concurrently to Dodoens’ Cruydeboeck, Mattioli published the first edition of his translation of Dioscorides, the Commentarii, making Mattioli Dioscorides’ bona fide spokesperson. Just as Dodoens’ Cruydeboeck gave rise to significant translations and editions, Mattioli’s Commentarii took on a similar remarkable a role across Europe. Most importantly, Caspar Bauhin launched his own career through compiling plant synonymies while editing the 1598 Opera (consolidated works) of Mattioli’s publications.

Somewhere in these herbal episodes we find the inimitable alchemist Paracelsus (Theophrastus von Hohenheim, 1493-1551), the devolution of medicine as Simples, and several phytologists, such as Jakob Boëhme, Giambattista della Porta, Thomas Browne, and William Coles. These are names often tied to a Doctrine of Signatures (DOS), the idea that a plant would provide some sign or signal (such as leaf shape) to suggest value or appropriate usage, particularly as a curative. There isn’t much to relate in this regard; it isn’t even clear whether or not recognition of shapes drove any real trials, so I leave this episode aside, in that the Doctrine of Signatures was never a source of botanical understanding.

I mention the concept of Signatures because it appears in many texts and teachings. Moreover, some version of DOS has surfaced at many times in various cultures, with little evidence it was ever truly mainstream or marked any particular movement. The idea did not play significantly in herbals or materia medica, but allusions to shape and form often crop up in descriptions. One also needs to distinguish between Simples and Herbals, which overlap. Simples, however, read more like “Hints from Heloise” – compendia of suggestions and formulae for curing ills, removing stains, and cleaning house. (Bradley C. Bennett, 2007. “Doctrine of Signatures: An Explanation of Medicinal Plant Discovery or Dissemination of Knowledge?” Economic Botany 61(3) 246-255)

The latter flourishing of European herbals was informed by movement of plants, people, and pestilence around the planet. Despite concerns any of us have about the consequences of European voyages east (around Africa’s Cape to India and Asia), and west (across the Atlantic, to the “New World”) – the impacts are undeniable. European estates, gardens, farms, and cuisines changed through introduction of plants from around the world, plants Europeans had never seen or used before, and in many cases, plants of great economic potential.

Global exploitation and landed wealth led to luxury, and the herbalists were joined by other authors interested almost purely in beautiful garden plants, ushering in Europe’s age of florilegia, books overladen with lavish illustrations and information on spectacular flowers, reflecting an expanding garden flora that did not require every plant represent a cure or comestible. To gain a sense of the spectacular publications these flowers inspired, check out the 1613 publication, Hortus Eystettensis, on the web, where you will see incredible portrayals of horticultural forms of native European plants alongside remarkable introductions, such as Helianthus annus, the American sunflower and Mirabilis, Four O’Clock, the Marvel of Peru.

Bringing Order….

In searching the many herbals, florilegia, codices, etc. that were published in the 16th and early 17th century, we quickly realize that as plant names are concerned, things were coming to a head. Imagery and avarice overloaded existing systems for naming and categorizing plants. There were no standards as to how a plant name might be formulated, and the idea of “scientific” names was not imagined. And there was no system for identifying or organizing the growing wealth of global and horticultural forms.

Because Latin crossed borders as the scholastic language, it became the language of scientific letters and books. But Latin and vernacular publications were incompatible. It was a quietly growing chaos; lacking index or concordance to correlate plants discussed by different authors.

Help was imminent, through the assiduous work of brother bibliophiles, Jean and Caspar Bauhin. The earliest and most straightforward to be published was Caspar’s 1623 Pinax Theatri Botanici. This detailed work suggests the Bauhins had access to good resources, because Caspar was able to get his hands on scores of important books and take the time and effort to determine which plant names and descriptions were synonymous. For each entry in Pinax, someone had to typeset the descriptions by different authors Caspar had encountered for each different kind of plant. I can only imagine he must have acquired two sets of each book, cut them into ribbons of text, and then glued them to pages that correlated like with like. It almost seems overwhelming that someone had to handwrite and then interpret all of those crazy names and descriptions. But that is likely what happened.

Download and examine a copy of Pinax (there are free copies available over the internet.) It’s plainly obvious that Caspar Bauhin was going in the right direction. Even though he does not consistently follow the concept of a genus as the necessary way to group similar (or related) plant species, Bauhin seems fairly clear that he is dealing with plants in groups we would readily appreciate as genera. Pinax came out in 1623, more than 100 years before Linnaeus, and it was sufficiently important to hold sway for that entire century.

During that next 100 years, there were gradual advances. Jean Bauhin’s own take on a concordance (with more complete descriptions) was published posthumously. John Ray published descriptions and discussion that seem somewhat modern. Hans Sloane set a modern pattern, as an educated and productive gentleman scientist who began his career with personal field experience and published a flora of Jamaica in 1696. Works in English, such as Joseph Miller’s Botanicum Officinale (1722), read as amazingly contemporary treatments, with reference to genera and species, and casual adherence to the system on nomenclature soon to be formalized by Linnaeus.

And microscopy provided incredible insights through publications by Nehemiah Grew and Marcello Malpighi (see Cells as the Basis of Life, earlier in this Section). We learn through microscopy that the course of scientific inquiry constantly changes in response to a new technology. But inquiry also responds to new ideas. Sometimes, when a fresh, important idea surfaces, it’s as though blinders are removed and observations/data that have not made sense are instantly intelligible. Sex had that effect on systematics.

Joachim Camarerius (who followed Fuchs at Türbingen) made observations and conducted trials that convinced him plants reproduce sexually. Pollen is the male element (sperm) and the egg (in the ovule) is equivalent to the female. This was a slow-growing “aha” moment. One of the more entertaining interludes in botanical history was part of this story – the famous 1718 address by Sébastian Vaillant, his “Lecture on the structure of the flowers, their differences and the use of their parts .” Vaillant chose his moment, the opening day of classes for 200 medical students as well as gaggle of luminaries. Normally, the task would have fallen to Antoine de Jussieu, but he was out of the country. Thus Vaillant had the opportunity to stake his claim on the topic of sex in plants. He succeeded.

Vaillant’s prose was pregnant. In translation, (for the source, see 1717 in the following TimeLine) we read his discussion of unisexual flowers, in which Vaillant describes the activity of the pollen-bearing (male) anthers: “the tension or swelling of the male organs occurs so rapidly that the lips of the bud, giving way to such impetuous energy, open with astonishing speed. In that moment, these enthusiastic organs, which seem to think only about satisfying their violent desires, abruptly discharge in all directions, creating a tornado of dust which expands, carrying fecundity everywhere; and by a strange catastrophe they now find themselves so exhausted that at the very moment of giving life they bring upon themselves a sudden death. “

The sensational aspects of Vaillant’s Discours struck a chord with Carl Linnaeus, who seems to have encountered the exposé about a decade later, while he was a student at Uppsala University. The sexual nature of plants promised to be such a profound aspect of plant life that Linnaeus was able to convince himself that hard data about stamens and pistils (numbers and structure) provided inherent evidence as to how plant groups differ, most especially genera and higher groupings.

“ we see the victory of the ingenious and simple method of classification of plants developed by Linnaeus. However, classification is not yet sys tematics; a key is not a monograph.” (Staffleu, 1971)

And at a functional level, he was right. Using Linnaeus’s methods, counting stamens and examining pistils, often provides a quick and simple way to sort plants into categories, making identification much easier, and bringing order to the mental storage cabinet. In fact, Linnaeus’s system was, basically, one spectacular key, not a typical binary key, but a functional sorting system to get to a genus of plants. That order, the beauty of it all, was short-lived, in that Antoine de Jussieu and the other people who had worked for the Jardin du Roi in Paris were already imagining a way of grouping plants they believed reflects natural affinities.

Jussieu (and future generations of botanists) would remain indebted to Linnaeus however. Beginning with Species Plantarum (1753), Linnaeus had rigorously applied Latinized binomials (a generic name and a specific epithet) to plants. Due to the popularity of his works and his own fame, Linnaeus nomenclature established a new international standard. Plant names have made sense ever since. Indeed, Jussieu adopted binomial nomenclature for Jardin du Roi in 1776, which likely contributed to the universal appreciation of his contribution to systematics.

A Natural System…

Jussieu’s Paris system came into full expression in 1789 with publication of Genera Plantarum. Linnaeus’s purely mechanical system of classification, in some instances, grouped plants botanists understand to be naturally related, but just as often, plants could be strangely separated from their nearest relatives. For example, some azaleas (Rhododendron) have five stamens, while others have ten. Otherwise these plants are incredibly similar, and clearly closely related. But in Species Plantarum, Linnaeus grouped Rhododendron as the class Decandria (SP page 392) and Azalea as Pentandria (SP page 150). De Jussieu’s Genera Plantarum described Rhododendron and Azalea in the same class and order, one after the other, on GP page 158. For people who studied plants, this new organization recognized the inherent order in Nature – it is a “natural system.”

It’s critical, however, to differentiate between Jussieu’s concept of what is natural as contrasted with a post-Darwin interpretation. Jussieu, and systematists who followed in his mold (i.e., Condolle, Brown, Willdenow, Lindley, and Bentham) were, in no way, imagining that a natural system reflects evolutionary origins, or lineages. These plant experts were solidly schooled in the concept that Nature came into being fully-elaborated. All of God’s creation appeared on the 3rd day, even before the stars, which shone at the end of the 4th day.

It was a matter of doctrine, now long abandoned. But do not confuse church-based acceptance with lack of technical knowledge. Those botanists were far from ignorant, and knew more about plants than most educated people in the world today.

Clearly they had studied plant variation enough to understand one can describe each kind of plant based on its similarities to another distinct kind. It was evident such pairings could be discovered through the entire plant and animal kingdoms.

The idea emerged that one might imagine Nature as elaborations on themes, interpreted as a chain of possible types, a continuum. Botanists came to envision a plan for Creation in which there were logical combinations of characteristics, graduating from simple to complex structures. Along that chain, each different kind of plant was one link, varying in some salient way from the link before and the one that followed. That meant that each different species represented a successive, possible combination, one link in the chain of being. This could be seen as a continuity from simple to complex forms, and would be interpreted as evidence of Nature’s plan. We had our hands on the the core rational for all of Creation. In today’s street language, someone might refer to this as part of Nature’s DNA.

A recognizable segment of the chain, consisting of one to many links (one to many species), would be designated as a genus. A concatenation of genera would constitute a family. To Jussieu, this meant genera and families were convenient ways to parse out the chain. He even used the number of entities as one factor in delimiting genera and families. However, it was clear the groups reflected a kinship of similarity, based on proximity in the chain of being.

The kind of similarity Jussieu imagined, however, was somewhat mechanical, like different kinds of railcars. A box car is similar to a refrigerated box car, but fundamentally different. A passenger car is yet more distinct. And a caboose is even more specialized. Then we discover coal cars, which are similar to (yet distinguishable from) hopper cars. If you assemble a train using one of every kind of railcar, ordering those cars based on some idea of similarity in structure or function, you’ll have a train of railcars analogous to Jussieu’s chain of being. Someone made all of those railcars by exploring the many variations that would be both possible and functional. Those kinds of railcars were built and the train of cars was Created.

If you knew every possible variation in railcars, then you’d know all of the possible cars in the train, that is, you’d know all of the links in the chain, your collection would be complete.

To Jussieu, each new plant species described would flesh out one of the links in the chain of being. When botanists had discovered and described every link, the collection would be complete. Nature’s great diversity would be explained, and our description of that arrangement would reflect the natural order of things.

Jussieu’s system and philosophy quickly eclipsed Linnaeus’s classes and orders. It reflected the times and made sense to people who knew enough about plants to realize that clear and intrinsic relationships were discernible in the great range of plant diversity.

Significantly, from Jussieu through Condolle, Brown, Willdenow, Lindley, and Bentham, this concept of natural relationships, though interesting, was ultimately academic. Lacking the idea of an evolutionary origin of species, these scientists could only opine as to what nature’s larder, its chain of being, (whether linear or reticulate, continuous or gap-ridden) told us about the qualities of different plants, or nature’s plan. But these botanists were not theologians, so there was not reverse extrapolation – knowing the plan of nature would not bring us closer to the mind of God.

Perhaps that complacency, or incapacity, explains one reason systematics went a bit quiet for several decades. Or maybe there was, simply, a new game in town, as attention and activity were drawn to the astonishing revelations of newly discovered plant species in the 19th century, a richness that propelled new fields in natural history. True, plant discovery had been a rising tide since Linnaeus, whose many students and supporters were dedicated collectors, venturing to areas around the globe.

These botanists and others were collecting and returning plants to Europe’s growing herbaria, most of which were still privately owned. Increasingly those specimens resembled Linnaeus’s own collection of pressed and dried plants, mounted on individual sheets of paper rather than bound unrelentingly in unintelligible volumes.

But as Linnaeus aged and his grip faltered, botany arrived at the era of Joseph Banks, who (in 1766, at the age of 23), was elected to London’s Royal Society. In 1778, Banks was selected as President of the Royal Society, a role he would occupy until his death in 1820. It would be difficult to overstate Banks’ importance in imperializing science, eclipsing Linnaeus in political and economic spheres. With Banks, the attention of science was drawn from organizing to the glory of plunder. Describing and enumerating nature’s larder, luxuriating in the richness and beauty of diversity proved so exhilarating that decades would pass before attention returned to order, and to the nature of creation.

And it seems Banks knew everyone on the scene. His endorsement advanced discoveries and careers of many scientists. Robert Brown was his protege, and Banks was an early supporter of his friend and correspondent, Alexander von Humboldt. George III relied on Banks’ opinion regarding investment in discovery and science. Banks seemed at the center ofbotanical science at an apogee of discovery. With eventual waning of Banks’ notoriety and influence, Joseph Hooker and a host of collectors, taxonomists, horticulturists, and sponsors kept the ball rolling. Concurrently, Alexander von Humboldt earned the role of wunderkind, collecting and compiling real data about climate and geography, establishing the basis for emerging studies in biogeography and ecology. Humboldt’s methodologies and exploits in South America brought that transcendental land home for many naturalists to consider.

Climate rules. Proper study of climate would explain the world’s vegetation types, yielding eventually to an understanding of the biomes. The collection, enumeration, cataloging, and mapping of plants supported an emerging sense of bioregions and growing appreciation of the great floristic regions… Realizations such as these demanded explanation. How did all of these plants originate? How did they come to colonize the world, and what is to be made of their patterns of distribution? With Bentham and Hooker listing nearly 100,000 plant species by 1883, how many more were to be discovered? What was the nature and extent of creation? What is the plan?

Realizations in floristics and ecology increasingly had confounded the idea of a linear chain, a continuity from the simplest to the most complex plant, was clearly untenable. Early in the 19th century, botanists like Willdenow had concluded there was not a single, long chain of being. The chain became regarded as a network, with branches and inter-connections. And botanists discovered sudden significant breaks, as well as isolated segments that were not easy to tie to other linkages. Some botanists even came to explain the fossil record as documenting links that had gone extinct. Even with those realities, even knowing the chain of being was a net (a reticulum), not much very new happened.

For Systematists who had been active before 1860, Darwin’s Origin of Species did not immediately alter course; systematics continued in a similar mold. Younger researchers, however, recognized that our basis of understanding had experienced a sea change. At the age of 48, in 1893, Charles Bessey (whose career matured after publication of the Origin) wrote: “It is now a full third of a century since a great light was first turned upon all biological problems by the formulation of the doctrine of evolution by the master mind of Darwin. In its light many puzzles have been solved and many facts hitherto inexplicable have been made plain. We now know what relationship means, and we have given a fuller meaning to the natural system of classification. From the new point of view a natural classification is not merely an orderly arrangement of similar organisms It is an expression of genetic relationship. The present similarity of two organisms is not enough to determine their relationship, or place in a system. Common origin must be inferred in order that relationship shall be assumed.”


By the beginning of the 20th Century, appreciation of Darwin and then Mendel had altered the course of nearly every botanical endeavor. As Bessey predicted, a search for natural order became increasingly redirected through concern for origins, evolutionary sequence, and lineage. Lacking understanding as to the physical basis for inheritance, systematists turned their attention to karyology – the study of nuclear chromosomes (which researchers knew harbored the plant’s genetic information.) It wasn’t yet clear how genetic information is stored or activated, but it seemed obvious that studying the nature of a plant’s chromosomes could yield understanding as to the nature of species. The era of Cytotaxomy had arrived.

But chromosomes and cytology were not the only new issues for taxonomists. The development of ecology, biogeography, pollination biology, and population biology challenged botanists to consider how plant biologies impact the way we should understand or delimit species.

Mapping plants and animals, relating those distribution patterns to climate and geography yielded real information that helped define species and generated new concepts about floristics, plant migration and distribution, and historical geography. Moreover, we learned that particular circumstances, such as long-isolated areas, like islands, merited special consideration and study.

Five decades earlier, observations by young Charles Darwin on his journey to the Galapagos Islands (off the west coast of South America) led to inevitable realizations that would focus Darwin’s thoughts about both naturally-occurring and domesticated populations of plants and animals. Those ideas found expression in Darwin’s Origin of Species… (1859) in which he crystallized and documented an emerging realization that plants, animals, and other kinds of organisms are not fixed, they were not simply created in one fell swoop. Rather the myriad species on Earth emerged through a natural process that evidences success in procreation. Each organism was the most prolific fit for its current circumstances.

Species, we came to understand, have come into being over hundreds of millions, billions of years. They have evolved, through countless generations. Darwin’s consolidation of many lines of thought, of much evidence, proved compelling, but left much to the imagination. If plants vary, what is the basis of inheritance; what is the source of that variation?

In Taraxacum: “The genus consists of more than 60 sections world-wide). A section contains one or a few diploid sexual taxa and a number of morphologically distinct apomictic polyploids (for which more than 3000 names have been published).” (Ľuboš Majeský,1 František Krahulec2 & Radim J. Vašut, 2017. “How apomictic taxa are treated in current taxonomy: A review” Taxon, 5 Oct 2017

More perplexing, of course, was the problem with limits of variation. If plants vary, what separates one “kind” of plant from another? What are the boundaries of a species? But we can defer these questions in that scientists still debate answers. The important tack to follow at this moment actually winds from other information.

In the first half of the 20th century, many important studies would inform understanding of variation within populations and the existence of regional adaptations we came to call ecotypes. Studies of plant-animal co-evolution led to a clearer understanding of isolation barriers and speciation.

Chemotaxonomy emerged as botanists began to extract and analyze various “secondary” compounds, such as flavonoids, anthocyanins, and betalains. Protein analysis and isozyme studies developed also. Under the “one gene, one protein” banner, researchers analyzed variation in proteins with the assumption we were getting closer to direct evidence of genetic variation.

These many techniques (cytotaxomy, phenotypic & population studies, and chemotaxomy) aggregated under the banner of biosystematics, suggesting the entire biology of a plant (not simply its external morphology) should be characterized in the effort to determine what distinguishes the life of one plant species as compared to another.

Though all of the kinds of studies we considered biosystematics remain important and continue to occupy graduate students around the world, the term “biosystematics” seemed to evaporate, almost overnight. The contemporary systematist now studied Phylogenetics, and if we must identify a year in which the the transition occurred, 2004 seems about right.

Phylogenetics, or is it Cladistics….

Willi Hennig gets credit and blame for launching the most recent era in systematics. The obvious impact of this methodology and philosophy has been at higher levels of organization. Biosystematics focused on the daily life of plant species, how they manage their business and interact. Phylogenetics, in reality, focuses on different issues – the course of evolution and the best way to determine the relationships among major plant groups.

Phylogenetics attempts to take the artifice out of constructing genera and families. All known, usable characteristics are scored and compared using different kinds of computational models. This weighting, most importantly, includes the growing addition of genetic information. The best result of analysis would be reconstruction and representation of the evolutionary lineage for all known plants. But most importantly, that representation would, somehow, imply equivalence. Equivalence suggests that family recognition is granted based on similar levels of divergence, or similar estimates of antiquity. Hennig worked with insects, and came to his own fresh system for determining relationships.

Where has this pursuit brought systematics? What has changed about the way we view plants? If you were grounded in systematics in 1985, you would have recognized names such as Robert Thorne, Armen Takhtajan, and Arthur Cronquist, These were the last authors of artful systems, outlines that attempted to organize major plant groups to reflect a presumption of evolutionary history. What you would remember from these authors is no longer very relevant.

Studies and statements increasingly reflect phylogenetic methods, particularly guided by the Angiosperm Phylogeny Group. Regarding flowering plants, perhaps the most fundamental change has been reconsideration of the simple monocots vs. dicots breakdown. Simpson (2010) explains that having embryos with two cotyledons is “an ancestral feature for the taxa of the flowering plants and not an apomorphy for any group within. Thus, “dicots” as traditionally delimited (all angiosperm other than monocots), are paraphyletic and must be abandoned as a formal taxonomic unit” This means the evolutionary groups of flowering plants are now mapped out differently, with several more ancient groups (Amborellas, Waterlilies, Chloranthoids, Magnolids) having emerged before the Monocots and other groups. That leaves a more tightly defined group of plants with similar pollen, which is termed the Eudicots. For botanists trained earlier in the 20th Century, another major departure has been resolution of plants previously in the Lily Alliance. Major groups, such as the Agaves, the Alliums & Amaryllids, the Irids, the daylilies, and the Orchids are now placed in Asparagales with Xanthorrhea and Hypoxis. The re-interpretation segregates the lily growth-habit from garden plants with equitant leaves and fleshy stems, representing a significant change for those of us who have always thought of daylilies and tiger lilies as reasonably closely-related.

Post Script: John Wilkins’ Species: A History of the Idea provides a very useful dialogue concerning the practical and philosophical origins of species concepts. To me, as a field biologist, much of this discussion is moot. In the field, in the garden, and in every day discussion, plant study requires a simple system of names, in order to attach information to groupings of plants you expect to closely share characteristics, to have the same biologies and chemistries, and to respond similarly to climate and stimuli.

At that hands-on level, the historical underpinnings of a plant name can be arcane. To horticulturists and field botanists, different plant groups speciate through differing processes, such that the parameters as to how you’d discriminate one “species” from another vary from orchids to cacti. Philosophically, scholars argue from a top-down perspective; a general kind has specific examples, each with its own essence. From a contemporary phylogenetic viewpoint, each plant is derivative as a branch in the continuum of evolutionary change that began hundreds of millions of years ago.

Phylogenetics traces the lineage of plants, establishes putative evolutionary relationships, and comes to some determination as to where breaks should be drawn. Reconciling that emerging, complex diagram with traditional taxonomy means a bit of a compromise, drawing lines that respect historical hierarchies, such as family, genus, and species but accepting the idea that one “species” will never be equivalent to another.

Detailed TimeLine


1500 A papyrus was created, documenting ancient information concerning plants of Egypt. The scroll describes hundreds of cures and treatments, including many ointments and extracts from plants. Purchased at Luxor (Thebes) by Georg Ebers around 1874, it is held by the library of the University of Leipzig, Germany (and thus is known as the Papyrus Ebers).

c300 Theophrastus (ca. 372-287 B.C.), the Father of Greek Botany, taught about plants from his own working knowledge of them, experience reflected in the “Inquiry” (Historia Plantarum) and “Causes” (De Causis Plantarum). Text covers 550 kinds of plants, including strawberry tree (Arbutus unedo), date palm, figs, and water lilies. His avoidance of more mystical notions about plants made a seemingly auspicious beginning for botanical study. During the middle ages, however, the Theophrastan works were generally unavailable, and second-hand versions were corrupted with misinformation – thus the level of botanical knowledge available in writing actually declined. The rediscovery and printing of his works beginning in 1483 replaced muddled interpretations of plants and helped rekindle an interest in botany. (HNT)


c50 Dioscorides (the Father of Medical Botany) authored his Materia Medica (HNT), a compilation of descriptions and medicinal uses for plants, including about 650 different species.

As the most widely known western botanical text during the middle ages, Dioscorides’ work became the basis for the earliest herbals. With an expanding awareness of the natural world in the 16th-century, herbalists began to make their own descriptions of plants, and at last Dioscorides’s influence waned.

c70 Pliny (Caius Plinius Secundus, A.D. 23-79), in his compilation called a Natural History (HNT), discussed about 1000 different plants. Well known throughout the middle ages, Pliny’s book constituted a major source of information on plants. Primarily an historian and storyteller, Pliny related accounts uncritically, even fancifully. Once original, rarer source documents were discovered and printed, errors in Pliny’s account became more obvious. Still the work remains valuable; it is through Pliny that we know the exact costs of many products, and that farmers alternated crops, such as beans and spelt.

Recognized in his comments was the growing trend of farm land consolidation into slave-maintained plantations. (Gras, 1946) On teaching: “Yes indeed, those who have gained a little knowledge keep it in a grudging spirit secret to themselves, and to teach nobody else increase the prestige of their learning.” (transl. Eamon, 1994)

1025 Approximate date for compilation of The Canon of Medicine, by Persian polymath Avicenna (Ibn Sina). The Canon describes about 760 different kinds of plants as the sources of useful medicines.

1256 Albertus Magnus produced De Vegetabilibus, based on ancient treatises and herbals, with added observations and descriptions. Gilla Wöllmer explains: “Albert the Great’s botanical work De vegetabilibus libri VII is based on the little treatise De plantis, which was ascribed to Aristotle in the Middle Ages, but is in fact a work of Nicholas of Damascus… Albert the Great’s treatise comprises seven books. Books 1–4 belong to the general theoretical area of botany; Books 1 and 4 have the treatise De plantis as their principal point of reference… In Books 6 and 7 he presents a special and applied botany. Beginning with the individual presentation of trees and herbs in Book 6, the account he gives of non-indigenous plants is indebted to the Canon medicinae of Avicenna, the Circa instans, and texts by other authors. Book 7 is devoted to agricultural botany…, here, he draws several times on the Opus agriculturae of Palladius. His treatise De vegetabilibus, which is the most extensive Latin commentary on De plantis, is much longer than the latter, both in terms of its sheer size and in its elaboration of the subject matter, ‘but it remains indebted to this work in its title and its theme: without the De plantis and its position in the Corpus Aristotelicum, the De vegetabilibus could not have existed in this form.’ ” (G. Wöllmer, 2012. “Albert the Great and His Botany,” in A Companion to Albert the Great, Brill OnLine.) As philosophers on which he relied, Albertus Magnus grouped plants as trees, bushes, shrubs, herbs, and fungi, expressing a worry that the categories overlap, but considering trees the highest and noblest of plants. Common belief held plants as the simplest of three orders of life (vegetative, sentient, & rational). Plant form and function were interpreted as compared to human(or animal) structure. Thus the sap of a plant was central to its life. Roots, which bring in water and nutrients, were regarded as homologous to a mouth, eating the earth for nourishment. Flowers were present to announce the coming of fruit. Etc.

1406 Zhu Xiao (朱橚) published Jiuhuang Bencao , an illustrated description of famine foods., describing and illustrating (woodblock) 414 kinds of plants.

1484, The Latin Herbarius, the first German herbal, was published in Mainz. An earlier natural history manuscript, Puch der Nature (the Book of Nature, ascribed to Conrad von Megenberg), was in circulation well before 1400, and then printed in 1475, but it was not a predecessor to the Latin Herbarius. (Locy, 1921)

1491 Jacob Meydenbach published Ortus Sanitatis in Mainz, which was basically the most extensive of a suite of related publications beginning with the German Herbarius zu Teutsch (Gart der Gesuntheit,) issued in 1485. The 1491 Ortus (in Latin) included 1066 botanical plates, that are said to be inferior to the 386 plant images provided in the earlier Herbarius (which is sometimes called the “smaller Ortus.”) (Locy, 1921)

1493 Carolus Clusius, having relocated to Leiden, established the Hortus Academicus, said to be the first botanical garden dedicated to ornamental plants. The valuable collection of tulips he cultivated there provided much of the material for the growing Dutch tulip industry – apparently through theft as much as sale or gift. (Grimshaw, 1998)

1526 The accepted year for production of The Grete Herball, an illustrated volume in the English vernacular. Believed to have been printed by Peter Traveris, the herbal is a translation of the 1498 French volume Le Grant Herbier, which itself compiles and “borrows” from earlier writings. The illustrations, likewise, are taken from the French volume, which is indebted to earlier books, including the 1485 German herbal, Herbarius zu Teutsch (Gart der Gesuntheit.) (Wikipedia, 2018 & Locy, 1921)

1530 Brunfels published Herbarium Vivae Eicones, the first newly written and printed botanical book/herbal. The “German fathers” were working with the North European cultivated and natural flora, which presented regional challenges for traditional Dioscoridean texts that continued as the basis for both materia medica and floristic plant studies.

1542 Fuchs published De Historia Stirpium Commentarii. By 1543 he had issued the German version, New Kreüterbuch. Illustrations for his herbals were based on studies of living plants, rather than on the simplified images that had become common in various scribed editions of the Apuleius herbal. [See c. 350] When describing European plants, however, text was taken essentially from information ascribed to Dioscorides. (HNT) Much later, the plant genus Fuchsia was named in his honor. The 1543 Dutch translation of New Kreüterbuch may have been generated by Dodoens, forming the basis for his Cruydeboeck.

1552 Created in Mexico, Libellus de Medicinalibus Indorum Herbis (Little Book of Medicinal Herbs of the Indies), was reportedly composed by native herbalist Martín Cruz in Nauhuatl (a Nauhuatl version is not extant.) The manuscript we know today was transcribed to Latin by Juan Badiano (applying Nauhuatl names to plants), and became known as the Badianus Manuscript. The manuscript, which had been transported to Spain, and then acquired by Cardinal Francesco Barberini, became property of the Vatican by 1902 ,and transferred to Mexico’s National Institute of Anthropology and History by Pope John Paul II in 1990. (Wikipedia, 2018)

1554 Rembert Dodoens published his herbal, Cruydeboeck, complete with 714 plates. The book was very well-received throughout temperate Europe, and soon translated to other vernacular languages – into French by Charles de L’Ecluse in 1557 and English (from the French) by Henry Lyte in 1578, and Latin in 1583 (an expansion and translation called Stirpium historiae pemptades sex.) The cumulative growth of the Dodoens’ works appeared through his productive life under various titles (culminating with the 1583 Latin Stirpium), an evolution that explains the many simple references to Dodoens’ work – as Cruydeboedk, Pemptades, and Stirpium….. Claims are that Dodoens’ growing product was one of Europe’s most translated books, second only to the Bible. Various renditions remained in use for two centuries. (Wikipedia, 2015) Excerpting from A.G.M. van Asseldonk, 2001

(“Traditional and modern herbalism in the Netherlands” a Research Report presented as a short paper at the Ethnobotany Conference, Antigua, Guatemala, 14-18 sept. 2001): “The first herbal in the Dutch language was (1554) the “Cruydeboek” (herbarium) of Rembert Dodoens (1517- 1586) (fig. 5). Dodoens was a Flemish Physician and also professor in Leiden University. He chose to call himself Dodoneus. Contributions to this herbal were also made by Dutch botanists such as Clusius and Lobelius. The work added new plants and experiences to the work of Dioscorides, which after 1500 years was still the most important in those days. It contained 1060 plant descriptions (of which 109 were original) and 715 figures (200 again original, 515 were used from the German herbal of Fuchs) 7. The book has played an important role in the past and present. So many copies were printed (last edition was the 6th in 1644) that they are still easily obtainable in the antique market and some herbal healers today still use the original antique book as a work of reference.” A Dutch website, “Plantaardigheden” ( makes two of the Latin (postmortem) versions available. See also:

1554 Pietro Andrea Mattioli published a Latin translation/his version of Dioscorides, having composed a very successful Italian translation ten years before (based on Ruel’s Latin translation of a Greek text). Mattioli’s Latin edition “was a publishing success on all counts,” perhaps because he added names from other sources, which made the text more widely useful, or because the book was well-illustrated. Mattioli followed this with a 1558 edition with 133 new illustrations and expanded commentary (set in typeface so as to stand out as clear additions). By 1665, Mattioli had produced a fourth edition, which included Greek, Arabic, German and French synonyms for the Latin and Italian plant names and larger illustrations (that had been created for a translation). Over the two decades since the first Italian publication, Mattioli’s many editions had spawned numerous translations, French, German, even Bohemian, as well as new drawings and woodcuts. (Stannard, 1999; XIV 71)

1557 Carolus Clusius issued his French edition of Dodoens’ Cruydeboeck.

1559 Conrad Gessner recorded the earliest known instance of a tulip flowering in cultivation in Europe, in the garden of Johann Heinrich Herwart of Augsburg (Pavord, 1999.) Gessner is said to have received these bulbs himself from Ogier Ghiselin de Busbecq, ambassador from Holy Roman Emperor Ferdinand I to the Ottoman court of Suleiman the Magnificent. Busbecq reported to Gesner that the highly colored flowers were called tulipam by their Turkish admirers, though the native word for these plants is lalé. Confusion as to the name could have had something to do with the turban (dulban) shape of the bulbs and flowers, but the true origins of the word “tulip” are lost in time. (Grimshaw, 1998)

1568 The New Herball of William Turner was published in completed form (in Cologne), including all three parts. Part 1 had been published in 1551 (in Antwerp), part 2 in Cologne in 1561. (Sanecki, 1992) Though woodcuts were copied from Fuchs, Turner added observations from his own working knowledge of herbs. Turner’s book is often described as the first herbal in the English vernacular, though earlier titles, such as the Grete Herball, might claim this distinction [See 1526]. (Wikipedia, 2015)

1568 Dodoens Florum, et coronariarum odoratarumque nonnularum herbarum historia was issued. One of the earliest European treatments of ornamental plants, the Florum includes garden introductions from Asia.

1569 Joyful News… published by Monardes from Seville between 1569 and 1574, later published by John Frampton in English, 1577, as Joyfull News out of the Newfounde Worlde. Many new plants are discussed in this book, including tobacco and sunflower (the first mention). In 1596 John Gerard described the sunflower, which he had grown in his own garden. By 1665 John Ray commented that the flower’s popularity had subsided. Joyfull News… seems also to be the first mention in Europe of the American native tree sassafras [See 1586].

1570 Matthias de L’Obel published his Stirpium adversaria nova, a compendium of approximately 1200 kinds of plants, which he organized based on leaf characteristics. His system of organization influenced the layout of Gerard’s Herball (1597), as well as the much later work of Linnaeus. L’Obel moved from Flanders to England in 1584, where he was appointed Physician to James I. (Encyclopaedia Britannica, on-line)

1574 Nine years following his death, Swiss naturalist Conrad Gessner’s Historia Plantarum was published. The Historia included 1500 illustrations, most (or all) of which were drawn by Gessner himself.

1578 Henry Lyte, also Henry Lite (sometimes awkwardly listed as John Lite or John Lyte) published A niewe herball or Historie of Plantes…, as the English translation of Charles de L’Ecluse’s Histoire des Plantes (1557), which was the French translation of Cruydeboeck (1554) by Rembert Dodoens.

Though published nearly two decades prior to Gerard’s Herball, and even though both are essentially translations of Dodoens, Lyte’s version of Histoire never became as widespread or well-known. (Wikipedia, 2015 – note dates for both L’Cluse and Dodoens in the Wikipedia treatment of Lyte were incorrect.) In the Herball, Lyte introduced the word “arborist” to English, based on the Latin arborator. (Campana, 1999) Lyte’s preface (in the 1619 edition) reads: “TO THE FRIENDLY AND indifferent Reader… IF thou be ignorant (gentle Reader) and desirous to know, either how profitable this Historie of Plants is, or how worthy to be studied, A good thing, the more common it is, the better it is ”

1583 De Plantis libri by Andrea Cesalpino became the greatest botanical book of the 16th century and the first general plant science text to supersede ancient writings. In the preceding 2000 years, little had been added to our knowledge about plants. Like his predecessors, Cesalpino accepted anecdotal information, but he advanced plant study in some regards, particularly in his grouping of plants by their physical characteristics (such as habit, i.e. tree versus herb, and fruit type) rather than by their supposed medicinal properties. As an introduction to his topic, Cesalpino classified around 1500 plants. Laurence (1951) is not so kind, i.e.: “Cesalpino was an Aristotelian scientist in that his conclusions were based on reasoning rather then an analysis by observation. Teleology was accepted by him and treated as of major importance. He believed leaves to have been provided for the protection of buds, flowers, or fruit, denied the existence of sex in flowers, treated the pith of dicots as a homologue of the spinal column of vertebrated animals, and contended that plants had a nutritive soul.” Cesalpino was a student of Luca Ghini [See 1533; 1543.] The bean genus Caesalpinia was named for him.

1583 Publication of Stirpium historiae pemptades sex, the Latin culmination of Dodoens’ studies (see 1554).

1592 Prospero Alpino published De Plantis Aegypti liber (which was included in Bauhin’s 1623 Pinax), relating plant information he garnered from having lived in Egypt. The book included citations of many tropical plants not previously mentioned in European herbals. Among his better-cited comments were observations of requisite pollination of date palms: “the female date-trees or palms do not bear fruit unless the branches of the male and female plants are mixed together; or, as is generally done, unless the dust found in the male sheath or male flowers is sprinkled over the female flowers”.

1596 In a catalog of plants in his Holborn garden, Gerard included what may be the first mention of the garden Nasturtium (probably Trapaeolum majus). A much later publication, Aiton’s Hortus Kewensis, notes that Lumley Lloyd introduced this plant to horticulture, in 1686. (Halliwell, 1987)

1597 Gerard published the first edition of his Herball, followed eventually by a second edition in 1633, which was edited and expanded by Thomas Johnston. Titled The Herball or General Historie of Plants. (Sanecki, 1992). Jackson (see 1876 in this TimeLine) describes the Herball as an English translation of Dodoen’s Stirpium Pemptades (the Latin translation of Cruydeboeck) reorganized along the lines of L’Obel’s work.

1598 Caspar Bauhin edited the Opera quae extant Omnia, the consolidated life works of Pietro Andrea Mattioli. The importance of this publication is overlooked, but it marks the beginnings of botany as emerging separately from medicinal realms. Mattioli had, through his many studies, labors, and disagreements become the Renaissance spokesperson for ancient Dioscoridean text. His translations and commentaries (beginning in 1544) were the basis for the re-emergence of classical thought on medicinal plants throughout Europe. For the Opera, Bauhin authored a list of synonyms for the many plants Mattioli had covered, a list that became the basis for his 1623 Pinax. (Stannard, 1999,XIV, 73)

1603 Spigelius published instructions on making dried herbarium specimens (in his Isagoges in Rem Herbarium) – a technique that had only come into practice during the previous 50 years. The collecting, exchange, archiving, and study of pressed, dried plants that are mounted to sheets of paper engendered a quiet revolution in taxonomy, floristics, and systematics. (Morton, 1981)

1605 Carolus Clusius published Exoticorum libri decem, an important step in the documentation of his work. (On line: The Exotic World of Carolus Clusius (1526-1609), Catalogue of an exhibition on the quatercentenary of Clusius’ death, 4 April 2009, Ed. Kasper van Ommen, Leiden University Library: For the first time in European history a work was dedicated to exotic nature as such, and not to its medicinal effects. Published when he had reached the ripe old age of 79, it includes an Appendix to Clusius’ collected works of four years earlier (the Rariorum) which lists newer discoveries without even bothering to include page numbers. The sense of the urgency of discovery that emerges from these pages symbolizes Clusius’ involvement with the exotic and his unwavering fascination with rare naturalia.)

1608 Jean Robin and Pierre Valet published the first European florilegium, Jardin du Roy tres Chrestien Henri IV. It was followed closely by Florilegium Novum (1611-1614), Florilegium Renovatum (1641) by Jean Theodore de Bry, Besler’s Hortus Eystettensis (1613), Emanuel Sweert’s Florilegium (1612), and Hortus Floridus by Crispin de Passe (1614). These books covered extensive numbers of horticultural floral forms.

1613 Basilus Besler published Hortus Eystettensis, a highly illustrated codex documenting plants in the garden (hortus) of Johann Konrad von Gemmingen, the Bisho of Eichstätt. Organized based on the four seasons, Besler’s team of artists and writers worked over 16 years to produce and publish the hundreds of engravings. Besler’s work included 660 species and more than 400 variants (doubles, variegates, etc); 400 of his plants had medicinal value, 180 were used in cooking, and 250 were grown principally for ornament. The publication included numerous forms of lilies, campanulas, delphiniums, hollyhocks, scabiosas, iris, tulips, narcissus, roses, hyacinths, and anemones.

1618 The London Pharmacopoeia was first issued, attempting to standardize and improve medicinal preparations available in commerce. Subsequent editions issued in 1621, 1632, 1639, and 1677, were modest improvements. The 1721 edition, published through oversight of Hans Sloane, overhauled cures (eliminating many unsubstantiated formulae), included better plant names as sources, and standardized strengths of solutions. (Wikipedia, 2018)

1623 Gaspard Bauhin produced the Pinax, a monumental compilation that pulled together uncoordinated plant names and descriptions on 6,000 species that had appeared in Theophrastus and Dioscorides, as well as in later herbals and other plant records. Pinax is among the first publications to distinguish between general categories (genera) and more specific examples (today’s species) of plants. Gaspard’s brother, Jean Bauhin created a separate comprehensive synonymy that expanded the listing with descriptions that constitute some of the earliest examples of appropriate plant diagnoses. His compilation, Historia plantarum universalis was published in 1650, nearly two decades following his death. The Bauhins’ publications remained core authorities for a century, and provide the basis for Linnaeus’s floristic works. By accepting their compilations, Linnaeus avoided many complications of the ancient literature. Linnaeus recognized the brothers with the genus Bauhinia, a legume that produces bilobed leaves.

1629 John Parkinson produced Paradisi in Sole Paradisus Terrestris, a gardening book/florilegium that resulted in his gaining the title Botanicus Regius Primarius – Royal Botanist.

1637 Tradescant f. (the son, filius, of elder Tradescant) made his first trip to Virginia, returning to England with living material of bald cypress and American sycamore. Tradescant f. made his second trip to Virginia in 1642. John Tradescant introduced Mimosa pudica, the South American sensitive plant, to cultivation in England. (Grimshaw, 1998)

1640 John Parkinson published his Theatrum Botanicum in which plants are classified according to 17 classes or tribes; i.e. 1. Sweet smelling Plants; 2. Purging Plants; 3. Venemous Sleepy and Hurtfull plants and their Counter Poysons; 4. Saxifrages; 5. Vulnerary or Wound Herbs; 6. Cooling and Succory Herbs; 7. Hot and Sharpe Biting Plants; 8. Umbelliferous Plants; 9. Thistles and Thorny Plants; 10. Fearnes and Capillary Herbes; 11. Pulses; 12. Cornes; 13. Grasses; 14. Marsh Water and Sea Plants and Mosses and Mushroomes; 15. The Unordered Tribe; 16. Trees and Shrubbes; 17. Strange and Outlandish Plants. (Sanecki, 1992) This is said to have been the largest English herbal produced, covering nearly 4,000 plants with more than 2,700 woodcuts.

1649 Nicholas Culpeper published his herball, The English Physician or an Astrologophysical Discourse of the Vulgar Herbs of this Nation Being a Compleat Method of Physic Whereby a man may preserve his body in health or cure himself being sick for thee pence charge with such things onely as grow in England, they being most fit for English Bodies. The English Physician dealt considerably with astrology and the signatures of plants. (Sanecki, 1992)

1672 Robert Morison published the first scientific study of a single plant group (the carrot family) [the first monograph.] (HNT)

1686 John Ray, in his Historia plantarum (published in volumes through 1704) arrived at an early natural grouping of plants through looking at their many different characteristics. His study dealt with plants worldwide, establishing standards and giving currency to much of our modern botanical terminology and summarizing the current state of botanical knowledge. Ray, unaware of the work by Rudolf J. Camerer, concluded in his discussion on fertility in date palm, willow, and other plants that: “in our opinion the pollen is equivalent to the sperm of animals.” His definition of species was quite modern: “each produces only its own kind; one must distinguish between essential, accidental, and environmental characters.” Ray’s summary of plant physiology was so thorough that he could be considered the founder of that field. (HNT) (Isely, 1994; Morton, 1981)

1694 Rudolf Jacob Camerer (in Latin, Joachim Camerarius) wrote a scientific letter (later published by Valentini in his Polychresta exotica, 1700, HNT) that made the first clear case (with solid experimental evidence) for the nature of sex in plants and the actual role of pollen and ovule in this process. The publication documented years of work with plants such as the dioecious Morus (mulberry), Mercurialis, and Spinacia (spinach), as well as Ricinus (castor bean) and Zea (corn), which are both monoecious. In all cases, removal of staminate plants or flowers either greatly reduced or completely eliminated fertility. In his experiments with Cannabis (hemp), removal of staminate plants from a field did not completely deter production of fertile seed, a result “at which I must admit I was quite upset” Camerer reported. (Morton, 1981) [See 1718] Of pollen: “In the vegetable kingdom there is accomplished no reproduction by seeds, that most perfect gift of nature, and the usual means of perpetuating the species, unless the previously appearing apices of the flower have already prepared the plant therefor. It appears reasonable to attribute to these anthers a nobler name and the office of male sexual organs.” (reproduced from Duncan Starr Johnson, 1915. History of the Discovery of Sexuality in Plants, Smithsonian Report for 1914, available free from G**gle Play)

1694, 1700 Joseph Pitton de Tournefort regarded flowering plants as either trees or herbs, with each divided based on floral construction and shape. He has been called “the father of the modern genus concept.” (Lawrence, 1951)

1697 Father Francisco Cupani published the first scientific description of Lathyrus odoratus, a plant from Sicily and the parent stock of today’s sweet pea. Seed that he sent in 1699 to Robert Uvedale, headmaster of Enfield Grammar School near London, resulted in cultivated forms, and by 1731, a famous selection called ‘Painted Lady’ – the exact origins of which are not known. (Grimshaw, 1998)

1699 Dr. Uvedale of Enfield received a shipment of Lathyrus odoratus (Sweet Pea) form Father Cupani in Sicily. Due to their form, color, and fragrance these plants became popular and their cultivation spread. A century later many variants were recognized, including the ‘Painted Lady’ which remains in cultivation today. (Fletcher, 1969)

1703 John Ray’s Methodus plantarum covered nearly 18,000 plant species, which he separated as either woody or herbaceous. Ray recognized monocots as separate from dicots, basin his classes on fruit (or cone) type, with finer segregation based on leaf and flower characters. Linnaeus substantially based his Critica botanica and his Philosophia botanica on Ray’s classification. (Lawrence, 1951)

1703 Charles Plumier published Nova Plantarum Americanarum Genera over a two year period. Plumier was an accomplished plant collector, and a prodigious author – leaving (at his death) 31 well-developed but unpublished manuscript volumes and 4,000 plant illustrations. Plumier described the genus Fuchsia from specimens collected in Hispaniola, and was himself recognized through designation of the genus Plumeria. (Wikipedia, 2108)

1712 Engelbert Kaempfer published Amoenitates Exoticae, the first western description of the Japanese flora (as well as other information). Kaempfer was a physician with the Dutch East India Company at Deshima from 1690 to 1692. Other Kaempfer notes, published by Hans Sloane as History of Japan, include the first western description of ginkgo.

1712 Mark Catesby made his first trip to America, traveling first to Virginia. He returned to England in 1719, but his time in the New World included travels to Jamaica in 1714. (Meyers & Pritchard, 1998) [See 1729]

1718 Sébastian Vaillant was one of the earliest supporters of Camerarius’s ideas concerning the sexual nature of plants. He contributed to the development of terminology necessary to discuss flower structure and function (some of which shocked his contemporaries, such as his comparing stamens to animal testicles and penis). Originally Vaillant delivered his information in a talk at the Jardin du Roi in Paris. By 1718 he had published the remarks as Discours sur la structure des fleurs… (HNT) (Morton, 1981) [See 1694](see also: Paul Bernasconi and Lincoln Taiz, transl. 2002. “Sebastian Vaillant’s 1717 lecture on the structure and function of flowers “Huntia 11(2) 2002.)

1722 Mark Catesby journeyed to America, leaving England in February, arriving in South Carolina on 23 May. Prior to his return to England in 1726, Catesby traveled to the Bahamas. (Meyers & Pritchard, 1998) [See 1729]

1722 Philip Miller began management of the Chelsea Physic Garden.

1722 Joseph Miller published Botanicum Officinale, a substantial compilation of plants, their origins and characteristics, and their uses in medicine. Miller’s treatment adopted names that had been listed in the 1721 London Pharmacopoeia (see 1618). Miller’s text was heavily borrowed (with permission) by Elizabeth Blackwell for A Curious Herbal (1737-1739), suggesting that Blackwell’s serialized prints constitute the illustrated version of Botanicum Officinale.

1729 As a student at Uppsala University, at the age of 20, Carl Linnaeus published Praeludia Sponsaliorum Plantarum, a discourse reflecting his understanding of the significance of the sexual nature of plants. Having been impressed by Vaillant’s letter, published just a decade earlier (though he seems to have denied that influence), Linnaeus applied that important revelation to his appreciation of plant diversity. Indeed, Linnaeus realized that floral characteristics provide incredibly useful characteristics for grouping and identifying plants, an observation bolstered by the knowledge that stamens and pistils are are a flower’s sexual parts.

1733 John Bartram of Philadelphia began correspondence with Collinson, Miller, and others. Their exchange is the likely source of pawpaw, sourwood, and other American plants introduced to cultivation in Europe. (Spongberg, 1990)

1735 Linnaeus arrived in Holland (for a 3-year stay), visiting the Amsterdam Hortus botanicus on his first day. In Holland Linnaeus gained the respect and support of three important botanists: Herman Boerhaave, Jan Frederik Gronovius, and Johannes Burman. Through Burman he gained the acquaintance and support of George Clifford, wealthy banker and owner of de Hartecamp. Since purchasing that estate in 1709, Clifford had transformed it into a botanist’s paradise of exotic plants. During his three years in Holland, Linnaeus published 14 books, laying the groundwork for his entire career. (Stafleu, 1971)

1735 – 1758 Soon after arriving in the Netherlands, Linnaeus met Jan Frederik Gronovius, who was sufficiently impressed by the young naturalist so as to finance publication of the first edition of Systema Naturae, the manuscript for which Linnaeus carried on the journey form Sweden. Over the next 23 years, Linnaeus published 10 editions of Systema Naturae (with two more editions surfacing the next decade). This publication constitutes his attempt at a comprehensive accounting of Earth’s plants and animals. The 10th edition (1759) included 7,700 plant species. It is documented that Linnaeus believed the world’s flora would total less than 10,000 species, which means he believed Systema Naturae approached a comprehensive accounting of the world’s plants.

1737 Linnaeus authored Hortus Cliffortianus, with illustrations by Ehret. This record of plants cultivated by George Clifford in his garden at Hartekamp (Holland) is the forerunner of Species Plantarum. The illustrations demonstrate Linnaeus’ belief that botanical drawings should be of superb detail and must result from close collaboration between botanist and artist. In his introduction, Linnaeus waxed “I gazed at Your garden in the very center of Holland bright with flowers, between Haarlem and Leiden, a charming spot between two thoroughfares, where boats, where carts pass by; my eyes were captivated by so many masterpieces of nature…” (Stafleu, 1971) (HNT)

1737 The magnificent Southern magnolia, Magnolia grandiflora (introduced from Southeastern North America to Europe by 1730) flowered in August at the London home of Charles Wagner, First Lord of the Admiralty. Georg Ehret immortalized this event with a sumptuous and justifiably famous illustration. Ehret, an apprentice gardener, had learned his artistic skills from his father during his youth in Heidelberg, Germany. (Grimshaw, 1998)

1737 Johannes Burman published Thesaurus zeylanicus, using plant specimens from Ceylon that were collected by Paulus Hermann and Jan Hertog. In the following two years Burman published Rariorum africanum plantarum decades I-X based on drawings made at the Cape of Good Hope by Hendrik Claudius. (Stafleu, 1971)

1737 Elizabeth Blackwell began publication of 500 illustrations of plants associated with Chelsea Physick Garden, which she drew (pinx), engraved (delin sculp.), and printed. The illustrations, issued in 125 weekly serials, constituted the two-volume set named A Curious Herbal. Accompanying text was taken from other sources, principally Miller’s Botanicum Officinale (see 1722). Blackwell’s herbal came at the close of the Herbal era.

1742 “Tokugawa Yoshimune decided that Dodoens’ Cruijdtboeck should be examined for useful products. He ordered his personal physician Noro Genjō to study Dutch and to interview the Dutch surgeons during the court journey to Edo.

These interviews took place between 1742 and 1750. Their purpose was to collect data on the medicinal properties of plants and to identify corresponding indigenous plants. The selected entries were compiled under the title Oranda Honzo-Wage. Each entry usually starts with the Dutch, Latin, Chinese and Japanese names of the plant, followed by its thera- peutic properties.” Quotation from: The Exotic World of Carolus Clusius (1526-1609), Catalogue of an exhibition on the quatercentenary of Clusius’ death, 4 April 2009, Ed. Kasper van Ommen, Leiden University Library

1747 Bernard de Jussieu received seed of Sophora japonica from d’Incarville in Beijing, via Moscow. This shipment probably also included Koelreuteria paniculata.

1748 Michel Adanson, a student of Bernard de Jussieu, arrived in Africa to collect until 1754.

1749 A near century old female specimen of the Mediterranean fan palm, Chamaerops humilis, had flowered for years in Berlin without fruiting. By 1751 Gleditsch reported that in 1749 he had applied pollen from a male plant grown in Leipzig to the flowers that remained fresh on one branch of the female plant. The seed produced proved viable, thus further confirming the male role of pollen. (Morton, 1981)

1752 Joseph G. Kölreuter (a medical student in Tübingen) published his survey of studies of sex in plants, one of the first that had been reported since Camerarius first suggested plant sexuality in his Epistola. Kölreuter had almost certainly been a student of S. G. Gmelin (a professor at Tübingen), who had republished Camerarius’s work and appended his own lectures calling for increased dedication to experimental work on this subject. (Morton, 1981) [See 1760; 1761]

1753 Linnaeus’ Species Plantarum established a standard for plant classification as well as nomenclature. Linnaeus included 5,940 kinds of plants in about 1,000 genera, organized in his 24 Classes. This treatise eventually became recognized as the beginning point for today’s binomial nomenclature. (HNT)

1759 Bernard de Jussieu organized plantings at La Trianon, Versailles, to reflect his developing concepts as to how plants groups are naturally (evolutionarily/phylogenetically) related. He segregated plants “on the basis of monocot vs. dicot, ovary position, presence or absence of petals, and fusion or distinctness of petals.” (Lawrence, 1951)

1759 Publication of the 10th edition of Linnaeus’s Systema Naturae (see 1735).

1761 Kölreuter reported his work on the role of insects in pollination. His detailed descriptions of insect activity and floral structure instructed botanists on the mechanisms and significance of insect pollination, and led directly to the work of Sprengel. (Morton, 1981) (HNT)[See 1716 & 1877]

1763 Michel Adanson’s Familles des plantes represented the first general attempt to group plants based on their relatedness, a “natural system.” The entry for each of his natural families presents a variety of characters common to the group Much of Adanson’s work predicted the system outlined in Genera plantarum, published in 1789 by Antoine Laurent de Jussieu. Adanson’s work was not widely recognized in his lifetime, perhaps because he did not use Linnaeus’s binomial nomenclature, which had already established its great utility. (Morton, 1981) (HNT)

1765 The Bartrams discovered the Franklin tree. Not until another trip, in 1773, would the younger Bartram collect seed in the only known population, near Fort Barrington, GA. In 1774, the supporter of this trip, John Fothergill, presented seedlings to Kew. Publication of William’s travel accounts was completed by 1781, but awaited identification of plants from specimens he had sent to Fothergill. At Fothergill’s death in 1780, his herbarium was purchased by Joseph Banks. (Spongberg, 1990)

1766 Joseph Banks explored Newfoundland and Labrador, charting waters and making collections.

1766 A colonial garden was established on St. Vincent, receiving mango trees as well as East Indian spice trees. (Sauer, 1993)

1766 Peonies and iris are said to have been first planted in Missouri by the Chouteau family, who brought the plants from Illinois. French settlers were the first to establish permanent settlements in Missouri (in St. Genevieve in 1755). (Edith Sinclair in Slosson, 1951)

1767 Pierre Poivre again was sent by France to Mauritius, as general intendant. The following year Poivre brought along his nephew, Pierre Sonnerat, who became a notable botanical explorer. Early in his career Sonnerat collected and transplanted the famous double coconut to the Mauritius garden from the Seychelles. Over the following two decades, Sonnerat contributed to our understanding of many tropical plants (such as dragon’s blood, breadfruit, banana, and cavalam), but his greatest energies were dedicated to the study of palms. (Duval, 1982) [See 1745]

1770 Australia was “discovered” by the British (though the Dutch had already named the area New Holland and had experienced at least 15 landings since 1606.) James Cook set out in the Endeavor on a scientific mission in 1768, with the young naturalists Joseph Banks and Daniel Charles Solander (a pupil of Linnaeus), as well as artists. On 29 April 1770, the ship stood into Botany Bay (an oceanic embayment 13 km south of Sydney), which Cook originally called Sting Ray Harbor – until the great collection of new plants by Banks and Solander provoked him to change the name.

1772 Joseph Banks was appointed scientific advisor for the royal gardens by George III.

1774 Antoine Laurent de Jussieu (nephew and collaborator of Bernard de Jussieu) proposed, in his Exposition d’un nouvel ordere de plantes, a classification of all plants as acotyledonae monocotyledoneae or dicotyledoneae. He divided the dicots into five groups, the apetalae, petalae, monopetalae, polypetalae, and diclinae. This system matured in Genera plantarum (1789). (Lawrence, 1951)

1774 Thomas Jefferson planted olive cuttings at Monticello – unsuccessfully. In 1791, he sent several hundred cuttings from France to South Carolina, only to be disappointed by the lack of commercialization. He was unaware that the Padres who established missions in California had planted olives there by 1769.

1775 Carl Pieter Thunberg arrived at Nagasaki harbor to work at Deshima with the Dutch East India Company. Thunberg received medical training in Sweden, and had been a student of Linnaeus. He was surprised to learn he had considerable freedom to collect dried specimens of plants on the Japanese mainland around Nagasaki. There he collected Hovenia dulcis and Rosa rugosa. Thunberg returned to Europe in 1776, having essentially smuggled his specimens out of Japan. He published Flora Japonica in 1784. (Spongberg, 1990)

1776 Antoine Laurent de Jussieu began using Linnaean binomials to identify plants at Paris’ Jardin du Roi. (Stevens, 1994)

1777 Carl Peter Thunberg, who eventually would occupy Linnaeus’s chair at Uppsala, was appointed botanical demonstrator at the botanical gardens. This followed seven years of travel and collecting in Europe, South Africa, Ceylon, Japan, and the East Indies. His work in Japan, because it was with Dutch merchants who held sole access to that country, required a stay in Java to learn the Dutch language. (Stafleu, 1971)

1778 Joseph Banks began his 42-year stint as president of the Royal Society.

1778 Lamarck (Jean-Baptiste-Pierre-Antoine de Monet de Lamarck) published his 3-volume Flore françoise, a contribution that continued his relationship with Bernard de Jussieu and established his credentials with many other scientists, most notably Georges-Louis Leclerc, Comte de Buffon.

1780 John Fraser traveled from England to Canada to collect plants; he entered US territory in 1785, receiving financial support from William Forsyth (Curator of the Chelsea Physic Garden), William Aiton (Head Gardener at Kew) and James Smith (President of the Linnean Society). He returned to America in 1788 and again in 1796. Fraser and his son returned yet later as collectors for the Russian Czar and Czarina. Their work was commemorated through plant names, Fraser fir and Fraser magnolia.

1783 José Celestino Mutis (a Spanish citizen who had moved to Bogota, Colombia in 1761 to serve as a physician) was successful in gaining support for a new “expedition” – an enterprise dedicated to documenting the region’s flora. Over the next three decades, Mutis employed and trained local peoples to paint several thousand spectacular and botanically-realistic paintings of local plants. The collection is held by the Real Jardín Botaníco de Madrid. (Bleichmar, 2017; Wikipedia)

1784 William Hamilton introduced ginkgo (Ginkgo biloba), Acer platanoides, and tree of heaven (Ailanthus altissima) to his garden near Philadelphia (the tree of heaven had first been planted in Europe by Miller at the Chelsea Physic Garden in 1751). Tree of heaven is now a major weed tree for eastern North America, and is “The Tree” that grew in Brooklyn. (Spongberg, 1990) [See 1770]

1784 David Landreth, along with his brother Cuthbert, established North America’s first substantial seed house in Philadelphia. D. Landreth & Co. was the country’s most important seed merchant for many years. (Hedrick, 1950)

1784 Thunberg published Flora Japonica. [see 1775]

1784 James Edward Smith, a botanist and associate of Joseph Banks, purchased Carl Linnaeus’ herbarium (about 14,000 sheets), correspondence, library, and other natural history collections from the Linnaeus family. By Smith’s death in 1828, his herbarium, totaling 27,185 specimens, became property of the Linnaean Society of London (which Smith and Samuel Goodenough had founded in 1788). The oldest collected specimen dates to 1708. (Linnaean Society website, 2018; Staffan Müller-Wille, 2006. “Linnaeus’ he Antoine rbarium cabinet: a piece of furniture and its function”. Endeavour Vol. 30 No. 2 June 2006; Jean-Baptiste Saint-Lager 1886. “Histoire des Herbiers” , Société Linnéenne de Lyon Année 1886 13 pp. 1-120)

1787 Publication began for Botanical Magazine by William Curtis, the world’s longest-running journal, dedicated to introducing exotic plants to an avid audience. Curtis resigned his position as Demonstrator in Botany for the Chelsea Physic Garden to produce this series.

1789 Aiton’s Hortus Kewensis recorded 15 exotic species of orchid at Kew. They were: Bletia verecunda, Epidendrum fragrans, Epidendrum cochleatum, Phaius grandifolius (syn P. tankervilliae), Cypripedium spectabilis, Cypripedium acaule, Liparis liliifolia, Calopogon pulchellus, Habenaria fimbriata, Arethusa bulbosa, Satyrium carneum, Satyrium coriifolium, Bartholina pectinata, Serapias lingua, and Nigritella angustifoliaEpidendrum cochleatum was the first epiphytic orchid known to have bloomed at Kew, in 1789. (Reinikka, 1972)

1789 Laurent de Jussieu achieved a workable system of naming and grouping plants in his Genera plantarum secundum odines naturales disposita, by combining Linnaeus’s nomenclature with a basic expectation that botanists would (eventually) discover plants that document a continuity in the forms seen in nature. This was not based on the concept that species further along the chain of life had evolved from earlier forms, rather the idea that nature would be shown to have provided examples of many different possibilities within each plant group. His treatment, by recognizing those sequences, also outlined the the manner in which those natural chains (not lineages) could be subdivided, thus establishing relatively natural groupings of major plant groups. Jussieu also had a fixation with numbers; groups could be neither too small nor too large. Genera plantarum grouped plants into 100 families, none of which included more than 100 genera. This caused some artificial adjustments, leading to his breaking the composites into three families. The book was published in Paris – during the same year as the beginning of the French Revolution. (Stevens, 1994)

1792 Carl Ludwig Willdenow, Grundriss der Kräuterkunde, accepted the idea that plant relationships do not form a single chain of being, that we must understand the continuity of plant life as reticulate (a network).

1798 A Franciscan botanist native to Brasil, José Mariano de Conceição Vellozo, began publishing his 11-volume encyclopedia (O fazendiero do Brazil) on the natural and agricultural circumstances in Brasil. His major taxonomic work, Flora Fluminensi, was published posthumously beginning in 1825. (JSTOR: The Text of Vellozo’s Flora Fluminensis and Its Effective Date of Publication, J. P. P. Carauta Taxon Vol. 22, No. 2/ 3 (May, 1973), pp. 281-284 Published by: International Association for Plant Taxonomy (IAPT) DOI: 10.2307/1218138 Stable URL: Page Count: 4) According to Frodin (see his Guide to the Standard Floras of the World. ), Martius criticized the work as duplicative and lacking citations.

1798 A chance encounter between Alexander von Humboldt and Aimé Bonpland (while both men were staying at Paris’s Hôtel Boston) resulted in an amazing journey of exploration [see 1799] that led to significant developments in our understanding of natural history, biodiversity, biogeography, and ecology. (Aniśko, 2013)

1799 John Lyon began collecting North American plants, at first for William Hamilton, and later for collectors in Europe. He followed the trails of Catesby, the Bartrams, Michaux, and the Frasiers. It has been suggested that Lyon may have contributed to the extinction of the Franklin tree by his aggressive and successful collecting, but that would also argue for the reality that the plant existed in very small numbers in a hyperlocal population. Lyon sent quantities of oakleaf hydrangea to England, a plant introduced by Hamilton in 1803.

1799 Alexander von Humboldt and Aimé Bonpland began their 5-year exploration of Central and South America. Among the six thousand plant specimens Bonpland had collected, scientists named 4500 new species. (Aniśko, 2013)

1801 In describing the new genus Lodoicea, J. J. H. Labillardiére commemorated an analogy made by P. Commerson (who served with Louis Bougainville on his historic voyages) between the form of the famous coco-de-mer fruit (the double coconut) and his image of the pelvis of Laodice (lovely daughter of King Priam of Troy). Previously, the plant had been included in the coconut genus, Cocos. (Emboden, 1974)

1802 Bernard M’Mahon established his nursery in Philadelphia and began his own limited publication series (1806) similar to Curtis’ Botanical Magazine . His seed lists are among the first published in the US. M’Mahon was selected to receive and germinate seed collected by the Lewis and Clark Expedition.

1802 Robert Brown arrived at Sydney (Australia) on the Investigator, along with botanical artist Ferdinand Bauer. George Caley, who had already been sent to collect plants in New South Wales by Banks, was furious that a second botanist was dispatched. (In 1803 Banks received seed of 170 species from Caley.)

1802 John Champneys of Charleston, South Carolina, created ‘Champneys’ Pink Cluster’ rose (eventual parent to the Noisette hybrids) through crossing ‘Parson’s Pink China’ with Rosa moschata, a white-flowered climbing rose from Asia. His new rose, a climber producing bunches of double, pink flowers, was quickly established in American gardens. Champney had acquired his China rose from the Noisette nursery in Charleston. Philippe Noisette produced seedlings from ‘Champneys’ Pink Cluster’ from which he selected the first Noisette, which was introduced in Europe through his brother in Paris. (Grimshaw, 1998)

1802 Seed were sold in packages in America, marketed by a Shaker community at Enfield, Connecticut. Keeping their own seed lines healthy and free of corruption was important to the community. (Connor, 1994)

1804 The Japanese devil lily (oniyuri) was brought into cultivation at Kew. Due to the ease of propagation from bulbils that form in the leaf axils, Kew gardeners were able to propagate and distribute over 10,000 plants within a decade. Scientific name Lilium lancifolium aside, the plant is known today most readily by its English common name, tiger lily. (Grimshaw, 1998)

1805 Alexander von Humboldt’s personal observations of many different plant habitats resulted in his important generalizations about the relationships of plants to their native climates. He is probably best known for making ecological correlations between the different plant habitats observed with rising elevation and the changing habitats seen when traveling from the tropics to arctic regions. Publication of his Essai sur la géographie des plantes… may be considered the beginning of the science of ecology. (HNT)

1810 Robert Brown’s Prodromus Florae Novae Hollandiae marked the beginning of his publications on the flora of Australia. Brown (the “Jupiter botanicus”) made important comparisons of plants from Australia with other floras, yielding a fresh approach to this type of study. With Brown’s work, botanists began to understand that significant information can result from studying the distributions and associations of plants. We also began to realize the distinctive nature of the Australian biota.

1813 While a professor of botany at the University of Montpellier, Agustin Pyramus de Candolle introduced the term taxonomy in his Théorie élémentaire de la botanique (Elementary Theory of Botany), as part of his new classification scheme.

1814 Frederick Pursh published his Flora Americae Septentrionalis. He had been engaged originally by Barton in 1805 to study the plant material collected by Lewis and Clark, and later he worked for Hosack at Elgin. In 1809 he returned to London with his own collections of plant material to study.

1815 Johann Friedrich Elsholz, a German, died while participating in a Russian expedition to California. Later, the German botanist, Adelbert von Chamisso, honored Elsholz through describing the new genus Eschscholzia for the California poppy, albeit misspelling the name of the honoree. Not a huge surprise; hardly anyone has been able to spell the scientific name of this plant, the state flower of California, ever since. (Grimshaw, 1998 – information on Chamisso in Grimshaw is incorrect – correction supplied by Ann Gardiner)

1816 John Reeves introduced Wisteria sinensis to European gardening from nurseries in Canton, China. The first two plants to be exported, each sent aboard a different ship, arrived in the same month of May. One of the ship Captains was Richard Rawes, famous for his involvement with introduction of the first camellias. (Grimshaw, 1998)

1818 Thomas Nuttall published the second volume of The Genera of North American Plants (and a Catalogue of the Species to the year 1817). On page 115 he described the genus Wisteria, “in memory of Caspar Wistar, M.D. late professor of Anatomy in the University of Pennsylvania and for many years president of the American Philosophical Society; a philanthropist of simple manner, and modest pretensions, but an active promoter of science.” Nuttall, with seeming purpose, named the genus Wisteria, rather than Wistaria; his spelling has been officially conserved by botanists. Horticulturists in England, however, continue to spell the genus Wistaria…..

Caspar Wistar’s contributions to medical science were significant, and in 1892 his great-nephew Issac Jones Wistar funded an endowment to establish The Wistar Institute of Anatomy and Biology, housing Caspar Wistar’s collections and honoring his contributions. The Wistar Institute stands as America’s earliest independent medical research organization. (Note: It would be easy enough to confuse Caspar Wistar with the later horticulturist John Caspar Wister, also of the Philadelphia area. J. C. Wister had a founding role in establishment of Swarthmore‘s Scott Arboretum during his 50-year tenure at the College.)

1821 Fifteen thousand specimens collected by Thaddaeus Haenke as part of the Malaspina Expedition [see 1879 entry] were secured by Prague’s National Museum. Haenke distinguished himself as the first Western botanist to explore the South American interior. As part of his continued explorations, Haenke had (in 1801) been the first scientist to see populations of the Giant Waterlily, Victoria amazonica. (Aniśko, 2013)

1823 Philipp Franz Balthasar von Siebold arrived in Japan to live there until 1830 as surgeon major in the Dutch East Indies Army, anxious for a career as a scientific explorer. He restored order to the botanical garden at Deshima. Because he accepted the gift of a map of Japan on his trip to Edo (foreigners were not allowed access to this type of information,) Siebold was imprisoned for a year, but pardoned in 1829. Banished from Japan in 1830, he was forced to abandon his Japanese wife and their child. The deck of the vessel on which he sailed was filled with plants he used to establish a nursery in Leiden. Among his introductions were Wisteria floribundaHydrangea paniculataHydrangea anomala, Malus floribunda and Rhodotypos scandens. He returned to Japan in 1859 and by 1863 produced a sales catalog that offered 838 species native to that country. (Spongberg, 1990)

1823 David Douglas was sent by The Royal Horticultural Society to the Eastern US to procure any new varieties of fruit trees and vegetables that might have been developed there. He met Thomas Nuttall (a British native recently appointed professor of Botany at the Harvard Botanic Garden) and others who helped him. Douglas returned to England with a wide variety of fruit trees, as well as Oregon grape holly. (Spongberg, 1990) [See 1804]

1823 Robert Bruce, and later his brother Charles, negotiated the process of acquiring seed and plants of the Assam form of tea (Camellia sinensis var. assamica) from the Singpho tribe of Upper Assam. Eleven years later, the East India Company recognized the value of this discovery and began establishing tea plantations in Assam, with the first tea arriving in London in 1838. The growth in this enterprise led to conscription and near-enslavement of several hundred thousand recruits from over India. Within 60 years, 340,000 acres in Assam were dedicated to tea plantations. With other growing areas established, Chinese tea exports plummeted from 100%, to 10% of the world market. (Hohenegger, 2007) East India Company employees Charles Alexander and Robert Bruce discovered a kind of tea previously unknown to Europeans (Camellia sinensis var. assamica) growing in Assam, a province of northern India. The first shipments of Assam tea arrived in England in 1838. Though attempts were made to cultivate China teas in India, it became clear that the native Assam tea was the better crop for that region. Today, Assam tea is grown in Africa as well as Papua New Guinea. (Lewington, 1990)

1824 Agustus Pyramus de Condolle published the first volume of Prodromus Systematis Naturalis Regni Vegetabilis, Prodr. (DC.), which continued through 17 volumes (completed in 1873), the last ten published by his son, Alphonse de Candolle and other authors. Intended to cover all seed plants, the series covered dicotyledons only. Condolle is known by his variance from the core Jussieu idea of continuity, having noted that newly discovered plants did not necessarily fill in the gaps between known species.

1826 In his paper (“Character and description of Kingia”) describing the Australian genus Kingia, Robert Brown also wanders afield and describes reproductive systems of cycads and conifers, which is recognized as constituting the first presentation of the significant differences between gymnosperms and angiosperms, particularly as relates to his concern that cycads and conifers do not appear to enclose the ovum in an ovularium (ovary). I’m a serious advocate for Robert Brown, and when I read this paper, I’m impressed by the fabulous recitation of historical work. But Brown so steadily hedges his bets in circular discussion about the cycads and conifers as to make his point fairly obtuse. To me, it isn’t a clear statement that instantly changes the field in a way other writers suggest.

1830 John Lindley, in his Introduction to the Natural System of Botany, explains: “The principle upon which I understand the natural system of botany to be founded is, that the affinities of plants may be determined by a consideration of all the points of resemblance between their various parts, properties, and qualities; and that hence an arrangement may be deduced in which those species will be placed next each other which have the greatest degree of relationship. A genus, order, or class, is therefore called natural, not because it exists in Nature, but because it comprehends species naturally resembling each other more than they resemble anything else.” Remember, this still does not impose any idea of evolutionary origin. Lindley is simply accepting the fact that there is a natural similarity (or affinity) among certain plants, which merits recognition and study. (Stevens, 1994)

1835 Having returned to London from nearly a decade of living, working, and collecting plants in Mexico and California, Thomas Coulter sorted through his notes and specimens. Early in this year, he loaned cones he had collected to botanist David Don. In June, Don read a paper detailing five new pine species from Coulter’s collections. Among those new species was Pinus coulteri, of which Don notes: “The leaves are longer and broader than those of any other Pine, and the cones which grow singly are the largest of all, being more than a foot long, half a foot in diameter, and weighing about four pounds… At the suggestion of Mr. Lambert I have applied to this remarkable tree the name of its discoverer, who is no less distinguished for his scientific acquirements than from the excellent qualities of his mind.” (Nelson and Probert, 1994)

1835 Hugh Cuming commenced a 4-year trip to the Philippines. He was probably the first person to ship living orchids successfully from Manila to England. Plants he sent included Phalaenopsis amabilis, first grown at Chatsworth. Cuming distributed 130,000 herbarium specimens.

1842 John Frémont began a series of five expeditions, which led to his arrival in Alta-California in 1844. Over a period of thirteen years, Frémont would be responsible for significant plant collections leading to designation of nineteen new genera. Included were specimens of Flannelbush, collected on 27 May 1946 and used by botanist John Torrey as the basis for the genus Fremontodendron. (Beidleman, 2006)

1843 Robert Fortune made the first of four journeys to China (ending in 1860), initially for the Royal Horticultural Society, later for the East India Company (as a result he sent 23,892 young tea plants and 17,000 germinated seedlings to northern India), and finally for the US Government. Never before had so many Chinese plants gotten to England. His success was based greatly on the newly invented Wardian case. Plants he sent included balloon flower, bleeding heart, golden larch, Chinese fringe tree, cryptomeria, hardy orange, abelia, weigela, winter honeysuckle, etc. Tea plants Fortune sent to Washington did not succeed, partly because of the War Between the States. (Spongberg, 1990) [See 1846]

1846 Robert Fortune delivered plant material (from his first of three China collecting trips) to the Horticultural Society’s gardens at Chiswick. Included was Jasminum nudiflorum (Winter Jasmine). Though originally cultivated in the glasshouse, the plant proved to be hardy and became a popular garden shrub. (Halliwell, 1987)

1846 William Lobb collected seed of Tropaeolum speciosum, the Flame Creeper, in Nothofagus forest of the south Chilean island of Chiloe. The plant was first grown by the Veitch nursery in Exeter, which had sponsored his trip. Flame Creeper is a close relative of Canary Vine (Tropaeolum peregrinum) and the garden Nasturtium (Tropaeolum majus) [See 1596], all of which are native to the Neotropics. (Halliwell, 1987)

1849 On 9 November, Joseph Paxton, Gardener to William Spencer Cavendish, and his staff flowered Victoria amazonica at Chatsworth. The seed had been delivered to Paxton by William Hooker, Director of Kew, whose staff had successfully germinated about 50 seed delivered in March. The event was regarded as a horticultural triumph, the first time this South American native plant had been cultivated and flowered successfully. (Aniśko, 2013)

1852 James Drummond (along with his son) made his last significant shipment of Australian plants to England, the results of his six major collecting expeditions. Drummond’s botanical legacy is strong; over 100 new species were named for him (a third of which have turned out to be taxonomic synonyms.) But he was known for fragments rather than full specimens, due partially to difficult circumstances and shortage of paper. (Webb, 2003)

1853 Albert Kellogg (native of South Carolina who had studied at Kentucky’s Transylvania College, and later traveled to San Francisco where he had a pharmacy) and six colleagues established the California Academy of Sciences. At one of the meetings, Kellogg relayed specimens as well as stories he had heard from A. T. Dowd about a giant new conifer in the Sierran foothills, southeast of Sacramento. William Lobb, who was at the meeting, left for the area soon afterward, collecting seed, mature cones, vegetative shoots, and two seedlings. Lobb returned to San Francisco and quickly departed for England. The two saplings were planted at the Veitch nursery in Exeter. John Lindley described the new species that December in Gardener’s Chronicles as Wellingtonia gigantea. (Spongberg, 1990) The name eventually accepted for this tree was Sequoiadendron giganteum.

1853 The first flower show held in California opened in San Francisco on 7 October. Among the entries were specimens from James Warren, of Sacramento, one of the first professional nurserymen to set up business in California (1849). Warren published the state’s first nursery catalog and initiated California Farmer, the state’s first agricultural and horticultural publication. By 1855, several nurseries operated along Folsom Street in San Francisco, including William Walker’s Golden Gate Nursery, James and William O’Donnell’s United States Nursery, and the Commercial Nurseries, a subsidiary of Highland Nursery in Newburgh, NY. A nursery from Napa County was represented in the 1854 flower show. (Taylor & Butterfield, 2003)

1853 The herbarium at Kew was established, based on: “the amalgamation of several formerly private collections, such as Sir William Hooker’s, George Bentham’s and M J. Berkeley’s mycological herbarium. The collections include the personal herbaria of some of Britain’s most celebrated scientists and explorers of the past. Charles Darwin, Joseph Hooker, David Livingstone, John Hanning Speke, Richard Spruce, Ernest ‘Chinese’ Wilson and Miles Joseph Berkeley are just a few of the famous names whose collections can still be studied in the Herbarium.” (Kew website,, 2018)

1856 Calanthe ×dominii flowered. This is the world’s first planned orchid hybrid, raised by John Dominy for Veitch & Sons. Though horticulturists were enthusiastic, botanist John Lindley was quoted as remarking: “You will drive the botanists mad.” (Fletcher, 1969)

1859 Louis Agassiz “insisted that the ranks of the hierarchy used in the classification of life were ‘instituted by the Divine intelligence as the categories of his mode of thinking.’ (Stevens, 1994)

1859 Charles Darwin published On the origin of species by means of natural selection As explained by Darwin, evolution is a simple change in character of a population of either plants or animals. Circumstances governing the success of a population are not neutral, rather the environment favors certain characteristics, which creates a natural system of selection that can lead to changes in the makeup of a population.

Gradual change in a population can lead to differences that qualify the population as a distinctive enough to become a new species – thus the “origin” of species. By identifying a mechanism that could lead to the diversity of life on earth, Darwin rewrote the book on relationships of plants and interpretations of plant adaptations. My favorite quotation from Origin: “We have seen that man by selection can certainly produce great results, and can adapt organic beings to his own uses, through the accumulation of slight but useful variations, given to him by the hand of nature. But Natural Selection as we shall hereafter see, is a power incessantly ready for action, and is as immeasurably superior to man’s feeble efforts, as the works of Nature are to those of Art.” (HNT)

1860 Joseph Hooker’s Flora Tasmaniae, which was published between 1855 and 1860 is dedicated to Ronald Campbell Gunn and William Archer. Of Gunn, Hooker writes: “There are few Tasmanian plants that Mr. Gunn has not seen alive, noted their habits in a living state, and collected large suits of specimens with singular tact and judgement. These have all been transmitted to England in perfect preservation, and are accompanied with notes that display remarkable powers of observation, and a facility for seizing important characters in the physiognomy of plants, such as few experienced botanists possess.” (Webb, 2003) William Archer, a native to Tasmania, was an architect and amateur botanist and botanical artist.

1861 A new treaty with Japan in 1858 [sic] led to a race by American and European plantspeople to collect and introduce plants from these islands. Field collectors included Carl Maximowicz who sent plants to Russia, Max Ernst Wichura from Germany, and Richard Oldham from the Royal Botanic Gardens at Kew (re. Bambusa oldhamii). George Rogers Hall, an American resident of Yokohama, sent a huge shipment in 1861 to Francis L. Lee of Chestnut Hill, MA. Lee went to war and left Francis Parkman, explorer, neighbor, and friend, to curate the growing collection. (Parkman would become Professor of Horticulture at Harvard in 1871). Thomas Hogg (son of a Scottish emigrant and nurseryman, sent to Japan by Lincoln as a US Marshal) shipped plants to his brother, James, as well as to the Parson’s firm at Flushing, NY. His introductions included the Japanese stewartia, the fragrant snowbell, the sapphire berry, and the katsura tree. (Spongberg, 1990) [See 1854]

1862 George Rogers Hall returned from Japan and brought seed, plants, and Wardian [See 1842] cases of material to Flushing, NY, which he entrusted to the Parsons & Co. Nursery. Included were the kobus magnolia, the star magnolia, zelkova, Japanese maples, wisterias, raisin tree, etc. Also in this shipment was the future weed, Japanese honeysuckle, initially called Hall’s honeysuckle. Some of Hall’s plants in Yokohama had been obtained from Siebold. (Spongberg, 1990)[See 1823].

1862 Specimens obtained by Jean Pierre Armand David, a Basque in the Lazarist priesthood who moved to China in 1862, form the basis of Plantae Davidianae, in which Adrien Franchet of the Museum at the Jardin des Plantes described nearly 1500 new species.

1862 George Bentham and Joseph Dalton Hooker began presenting a new system for organizing plants with publication of Genera plantarum ad exemplaria imprimis in herbariis kewensibus servata definita, three volumes published between 1862 and 1883. The monumental set, simply called Bentham and Hooker by most botanists, covered seed plants and included over 97,000 species in 7,569 genera, organized in 202 families. Bentham alone dedicated 27 years to this publication, and it stands apart from other comprehensive treatments of seed plants in that descriptions of families and genera were written by the two authors, based on study of specimens. (Lawrence, 1951; Wikipedia, 2018)

1866 Ernst Haeckel published Generelle Morphologie der Organismen: allgemeine Grundzüge der organischen Formen- Wissenschaft, mechanisch begründet durch die von Charles Darwin reformirte Descendenz-Theorie, which is noted as the first time phylogenetic trees were used to explain phylogenetic concepts. (Note: Haeckel studied and published on embryology, and is credited with the aphorism: “Ontogeny recapitulates phylogeny.”

1887 Initiation of the first edition of the 23 volume Die Natürlichen Pflanzenfamilien by Adolf Engler and Karl Anton Prantl (completed in 1915, followed by a partial second edition). Known simply as Engler & Prantl by botanists around the world, one might say constitutes the first approach at a true world flora. Engler’s schema for organizing plant families changed through many renditions, which were reflected in an 1889 guide to Breslau Botanic Garden (where Engler worked.) Paralleling his other publications, the catalog surfaced as Syllabus der Pflanzenfamilien in 1892, and passed through numerous editions, the 13th published in 2009.

Engler’s system was the standard basis for organizing many herbaria for decades, and thus held great sway over the world of systematists. Lawrence (1951) reminds us that: “the system was not conceived to be phylogenetic in the modern sense,” but, Lawerence does explain that Engler’s groupings of flowering plants are based “on the premises that evolutionary lines progressed from apetaly to polypetaly and gamopetaly, apocarpy to synarpy, hypogyny to epigyny, and actinomorphy to zygomorphy.” Alfred B. Rendle, Richard von Wettstein, and August A. Pulle would subsequently publish their own independent updates to the Engler and Prantl system. (Lawrence, 1951;Wikipedia, 2018)

1915 Charles E. Bessey published “The phylogenetic taxonomy of flowering plants” in Annals of Missouri Botanical Garden. (See The Reader, Chapter 5, Section 2 for a discussion of Bessey’s Dicta.) Bessey created his own system, borrowing strongly from ideas generated by Bentham and Hooker, as well as Engler. He thought of seed plants as polyphyletic, deriving the flowering plants (Anthophyta) from the cycads (Cycadophyta).

1922 Edgar Anderson began work in St. Louis, MO as “Geneticist to the Garden” (heading Missouri Botanical Garden’s School of Gardening) and professor at Washington University. Anderson would quickly become one of the principle figures in development of biosystematics. “His first biosystematic project was a look at the species problem in Iris. His 1928 paper took up the species problem concretely by looking at populations of two closely related yet distinct species of iris and allowed Anderson to test the relative importance of hybridization and mutations as sources of the variation on which natural selection works “ (Kim Kleinman, 2009. “Biosystematics and the Origin of Species: Edgar Anderson, W. H. Camp, and the Evolutionary Synthesis”, Transactions of the American Philosophical Society, New Series, Vol. 99(1) 73-91, in Descended from Darwin: Insights into the History of Evolutionary Studies, 1900-1970

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1926-1934 John Hutchinson published Families of Flowering Plants, a 2-volume classic that presented his phylogenetic system. Hutchinson separated predominantly woody groups of plants from those that are predominantly herbaceous, basing much of his thought on work of Charles Bessey.

1929 Göte Turreson explained several different and important concepts of ecological approaches to plant species, describing the Ecospecies, Agamospecies (a term he introduced in this paper), and Coenospecies. (Wilkins 2011)

1931 Walter Zimmerman published “Arbeitsweise der botanischen Phylogenetik und anderer Cruppierungswissenschaften“, which discussed the history of plant classification schemes and promoted what would (today) be considered a cladistic approach to establishing evolutionary history as the basis for systematics.

1937 In his publication Die Methoden der Phylogenetik, Walter Zimmermann explained that classificadtion must reflect phylogenetic relationships. Zimmerman’s ideas were important to Willi Hennig, as he developed his own statement on phylogenetics. (see Michael J. Donoghue and Joachim W. Kadereit, 1992. “Walter Zimmermann and the Growth of Phylogenetic Theory” Systematic Biology, Vol. 41(1): 74-85

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1939-1948 Jens Clausen, David D. Keck, and William M. Hiesey published a series of studies, using reciprocal plantings to document ecotypical variation across plant populations.

1939: “The concepts of species based on experimentation.” American Journal of Botany 26: 103–106; 1940: Experimental studies on the nature of species I. Effects of varied environments on western North American plants . Washington, DC: Carnegie Institution of Washington; “Heredity of geographically and ecologically isolated races”. American Naturalist 81: 114–133; 1948: Experimental studies on the nature of species. III: Environmental responses of climatic races of Achillea. Publication 581; Washington, D.C.: Carnegie Institution of Washington.

1942 Julian Huxley published Evolution: The Modern Synthesis, an influential book that called for overall integration of knowledge and techniques in establishing the origins and relationships among organisms.

1942 Ernst Mayr published notes from a lecture series in his book Systematics and the Origin of Species from the Viewpoint of a Zoologist. Considered a complement to Huxley’s Evolution: The Modern Synthesis, Mayr lays out his biological species concept, which informed working assumptions of biologists in all fields.

1943 With a goal to address the “rising tide of discontent with which the geneticist and cytologist view the systematist’s concept of species,” Wendell Holmes Camp and Charles Louis Gilly published “The Structure and Origin of Species” in Brittonia (vol 4: 323-385), in which they coined the term biosystematy, a term that morphed into the word biosystematics.

From their text: “The systematist has long recognized that many populations are diverse – that some species are morphologically homogeneous, while others are quite variable. Biosystematy seeks to explain the causes of these differences in the structure of species and so permit them to be arranged in a functional nomenclatural system,” and, later in the article: “biosystematy seeks (1) to delimit the natural biotic units and (2) to apply to these units a system of nomenclature adequate to the task of conveying precise information regarding their defined limits, relationships, variability, and dynamic structure.“ ( p. 327) [For more information on both the contemporary criticdl assessments and historical significance of the Camp and Gilly publication, see also: Kim Kleinman, 2009. Biosystematics and the Origin of Species: Edgar Anderson, W. H. Camp, and the Evolutionary Synthesis” Transactions of the American Philosophical Society, New Series, Vol. 99(1), in Descended from Darwin: Insights into the History of Evolutionary Studies, 1900-1970, pp. 73-91.

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1944 Chinese botanists reported discovery of living specimens of dawn redwood (Metasequoia glyptostroboides.) The tree hitherto had been known only from fossil material that was at least 20 million years old. (Rupp, 1990)

1948 John Hutchinson published British Flowering Plants, which updated and his Families of flowering plants, which was based on 22 principles. Lawrence (1951) simplified them as:

  1. Evolution is both upwards and downwards, the former tending toward preservation… and the latter to their reduction and suppression of characters.
  2. Evolution does not necessarily involve all organs at the same time…
  3. Broadly speaking, trees and shrubs are more primitive than herbs in any one family or genus…
  4. Trees and shrubs are older than climbers in any one family or genus.
  5. Perennials are older than biennials and annuals…
  6. Aquatic flowering plants are derived from terrestrial ancestors, and epiphytes, saprophytes, and parasites are more recent than plants of normal habit…
  7. Dicotyledons are more primitive than monocots.
  8. Spiral arrangement is more primitive than cyclic.
  9. Simple leaves are usually more primitive than compound leaves.
  10. Unisexual flowers are more advanced than bisexual; dioecious plants are more recent than monoecious.
  11. The solitary flower is more primitive than the inflorescence…
  12. Aestivation types are evolved from contorted to imbricate to valvate.
  13. Apetalous flowers are derived from petaliferous flowers.
  14. Polypetaly is more primitive than gamopetaly
  15. Actonomorphy is ore primitive than zygomorphy.
  16. Hypogyny is usually more primitive than perigyny, and epigyny is the most advanced.
  17. Apocarpy is more primitive than syncarpy.
  18. A gynoecium of many pistils preceded one of few pistils.
  19. Seeds with endosperm and small embryo are older than seeds without endosperm and a large embryo;…
  20. Numerous stamens, in general, indicate greater primitiveness than does an androecium of a few stamens (except Malvaceae).
  21. Separate anthers, in general, indicate greater primitiveness than does an androecium of either fused anthers or filaments.
  22. Aggregate fruits are more highly evolved than single fruits; as a rule the capsule precedes the berry or drupe.

1950 Willi Hennig, an entomologist, published his book Grundzüge einer Theorie der phylogenetischen Systematik, which is considered foundational in phylogenetic systematics, i.e. cladistics. The total impact of Hennig’s book was delayed until 1966, when the text was published in English as Phylogenetic Systematics. From Hoch and Raven (1995): ‘The most basic principles of Hennig’s theory of cladistics include: 1) relationship is clearly defined in terms of recency of common ancestry; 2) relationship is detected by means of shared derived characters (synapomorphies) in which are homologies that characterize monophyletic groups; 3) relationships and character distributions can be expressed in dichotomous branching diagram (cladograms); and 4) only natural groups, defined by synapomorphies to be monophyletic (i.e. descended from a common ancestor), are to be admitted into Linnaean classification schemes.”

1973 Over a decade, Gottlieb and collaborators refined interpretation of isozyme information detected from enzyme electrophoroesis to establish ideas as to evolutionary timelines. “The paper by Gottlieb (“Genetic differentiation, sympatric speciation and the origin of a diploid species of Stephanomeria.” Amer. J. Bot. 60: 545-553) should be required reading for graduate students in plant systematics, because it is an excellent example of how to apply isozymic data to studies of speciation.” Quotation and Information from: Daniel J. Crawford, 2000. “Plant Macromolecular Systematics in the Past 50 Years: One View” Taxon, 49(3): 479-501 Stable URL:

1981 So what is considered a plant? Zoologist Thomas Cavalier-Smith introduced the sub-Kingdom Viridaeplantae, which might be equivalent to Plantae, or Chlorobionta. In this organizational sense, the term plant includes green algae, most especially the Charophytes Quotation from, 20 December 2010: “Prof. Cavalier-Smith of Oxford University has produced a large body of work which is well regarded. Still, he is controversial in a way that is a bit difficult to describe. The issue may be one of writing style. Cavalier-Smith has a tendency to make pronouncements where others would use declarative sentences, to use declarative sentences where others would express an opinion, and to express opinions where angels would fear to tread. In addition, he can sound arrogant, reactionary, and even perverse. On the other [hand], he has a long history of being right when everyone else was wrong. To our way of thinking, all of this is over-shadowed by one incomparable virtue: the fact that he will grapple with the details. This makes for very long, very complex papers and causes all manner of dark murmuring, tearing of hair, and gnashing of teeth among those tasked with trying to explain his views of early life… Nevertheless, he deals with all of the relevant facts.”

1985 Rolf Dahlgren and K. Bremer cooperated to generate “the first computer-based analysis (using PAUP) of the angiosperms,” which was published in the first volume of the new journal, Cladistics. The tree structure generated did not long stand, as new algorithms were quickly developed and computer technology progressively improved.

1994 David Nobel (a national park staff member) discovered a stand of unusual trees in Wollemi National Park within 200 kilometers of Sydney, Australia. The trees were judged to represent an entirely new genus and species, Wollemia nobilis, in the Araucariaceae (the monkey-puzzle tree family).

1998 Publication of the first paper through the: Angiosperm Phylogeny Group “An ordinal classification for the families of flowering plants”, Annals of the Missouri Botanical Garden, 85 (4): 531–553, doi:10.2307/2992015, JSTOR 2992015

1998 One fallout of a meeting held at Harvard University was establishment of a committee dedicated to Phylogenetic Nomenclature, to devise a PhyloCode. Version 4c of the evolving PhyloCode, 2010, can be found on the internet, searching:

2004 In 1960, the International Organization of Biosystematics was organized at meeting of International Committee for Biosystematic Terminology, a Committee of the International Association for Plant Taxonomy (IAPT). The organization became fully independent in 1983, and following a symposium, 2004, in Valencia (Spain), reverted as an interest group of IAPT. In the same year, in Paris, the International Society for Phylogenetic Nomenclature was established at the First International Phylogenetic Nomenclature Meeting. (Taxon, November, 2004) Thus, 2004 has been selected as the year in which Biosystematics perished as a field of study.

2009 Publication of Pollination Biology of Basal Angiosperms (ANITA GRADE), by Leonard B. Thien, Peter Bernhardt, Margaret S. Devall, Zhi-duan Chen, Yi-bo Luo, Jian-Hua Fan, Liang-Chen Yuan, and Joseph H. Williams (American Journal of Botany 96(1): 166–182.) The paper defines characteristics of basal Angiosperms (their first three phylogenetic branches) which share similarities in pollination and chemistry: “Floral odors, floral thermogenesis (a resource), and colored tepals attract insects in deceit-based pollination syndromes throughout the first three branches of the phylogenetic tree.”

2009 John Wilkins’ Species: A History of the Idea

Some articles to explore:

Locy, William A. 1921. “The Earliest Printed Illustrations of Natural History,” The Scientific Monthly, 13 (3):238-258 Available on JSTOR