Linnaeus in the Netherlands: George Clifford

3 Clifford Hypericum androsaemum

Hypericum androsaemum from the Clifford Herbarium, courtesy of the Natural History Museum, London

As I mentioned in the first post in this series, Carl Linnaeus had just begun work with Johannes Burman at the Leiden Botanic Garden when George Clifford (1685-1760) asked Linnaeus to write a catalogue of the plants in his garden at Hartekamp, near Haarlem in the Netherlands.  It took some convincing for Burman to release him, but it ended up well for Linnaeus.  He spent over two years at Hartekamp, where he had available to him a large collection of tropical plants from around the world.  Linnaeus had already sketched out his Systema Naturae (1735) before he left Sweden, but his knowledge of plant diversity was limited to northern Europe.  Then he met Jan Frederik Gronovius, who had studied plants that John Clayton had sent him from Virginia and Burman, who had Paul Herman’s specimen collection from Ceylon (now Sri Lanka).  His horizons were broadening (see last post).

Clifford was a wealthy Dutch financier and a director of the Dutch East India Company (VOC) that oversaw a worldwide shipping organization making the Netherlands a mercantile power.  From the VOC’s creation in 1602, its captains and ship surgeons were given directions on how to make collections and transport specimens, seeds, bulbs, and cuttings back home.  The more exotics that reached home, the more the Dutch became avid gardeners hungry for still more plant novelties.  Because of his position, Clifford had first dibs on the plants that arrived in Holland, and he had the interest and knowledge to appreciate them.  To give a sense of the scope of his collection, he had four greenhouses, one each for plants from Europe, Asia, Africa, and the Americas.  At this time, gardening and sophisticated plant collecting were status symbols for the elite; Clifford’s Hartekamp was obviously a premier example.  Even his herbarium specimens reflected his status.  The sheets had elaborately printed labels, and the cut end of each plant was covered with a printed urn (Thijsee, 2018).  This became a fad at the time among the rich and botanically sophisticated (see figure below).

Among the living plants in Clifford’s unique collection was a banana tree, which was growing well but had never blossomed or produced fruit.  Linnaeus gave it special attention and took credit for inducing it into flower in four months with a regimen of restricting watering, and then watering generously.  This was one of the first times this feat had been achieved in Europe and was so noteworthy that Linnaeus wrote a short book on the plant, and Clifford had it published (Rutgers, 2008).  This added luster to both their names; it also indicated Linnaeus’s skills with living plants as well as with identifying specimens.

Another important event during this time was the arrival of the German artist Georg Ehret at Hartekamp in 1736.  Ehret had already produced a large portfolio of botanical watercolors for several patrons, none of whom paid very well.  He had come to the Netherlands after doing some work in England and called on Clifford in the hope of finding further employment.  Clifford was indeed interested in Ehret’s work and even paid his asking price for a number of paintings.  Ehret remained at Hartekamp for a month, working on illustrations for Clifford’s catalogue.  Linnaeus explained to Ehret his plant classification system based on the reproductive structures in flowers.  He had worked out 24 classes simply by counting the number and arrangement of the stamens or pollen-producing male organs, with the 24th class reserved for those without visible stamens.  Within each class were subclasses depending up on the number of female organs.  The beauty of the system was its relative simplicity, grounded in traits that were usually visible and countable.

Ehret illustrated the system with a chart that has become famous, a simple visual representation of the 24 classes (see figure below).  He published it shortly after leaving Hartekamp and Linnaeus also published it much later, but not crediting Ehret.  Working in close proximity together, even for a month, must have been important to them both during this early formative period in their careers.  Ehret, who had already developed the practice of dissecting flowers and illustrating their parts, often with magnification, learned from Linnaeus the pivotal importance of these structures in identifying species.  On the other hand, Linnaeus was able to see the artistic and intellectual work that went into creating first-rate botanical art.  In their book Objectivity, Lorraine Daston and Peter Galison (2007) write of four-eyed sight, which results from an artist and a scientist working and looking together, resulting in an image that satisfies both.  Linnaeus and Ehret could very well have collaborated in this way.  After he left Hartekamp, Ehret had a long career in England producing illustrations for many major botanical works including those of Philip Miller and Christoph Jacob Trew, who had been an early patron of Ehret’s in Germany.

3 Ehret
Georg Ehret’s diagram of Carl Linnaeus’s classification system, courtesy of the Biodiversity Heritage Library

Most of the illustrations in the Clifford catalogue were done by Ehret and the remainder by Jan Wanderlaar, who also engraved the plates.  It took Linnaeus nine months to write the text (Blunt, 1971).  The species descriptions were organized according the classification system Linnaeus had laid it out in his Genera Plantarum, which was also published during this time (1737).  While he was in Hartekamp, he published early versions of other works as well.  Clifford also afforded him the time and the resources to become better educated in botany.  Besides his herbarium and garden, Clifford also had a substantial library, with all the leading botanical references of the day.  Hartekamp must have been a difficult place to leave.  However, after spending almost three years in the Netherlands, Linnaeus’s thoughts were of Sweden.  Yet he didn’t go directly home.  His further wanderings will be examined in the next post.


Blunt, W. (1971). The Compleat Naturalist: A Life of Linnaeus. New York, NY: Viking.

Daston, L., & Galison, P. (2007). Objectivity. New York: Zone.

Rutgers, J. (2008). Linnaeus in the Netherlands. TijdSchrift Voor Skandinavistiek, 29, 103–116.

Thijsse, G. (2018). A contribution to the history of the herbaria of George Clifford III (1685–1760). Archives of Natural History, 45(1), 134–148.


Linnaeus in the Netherlands: Mentors

2 Claytonia virginica

Specimen of Claytonia viriginia collected by John Clayton, courtesy of the Natural History Museum, London.

In the last post, I outlined the early days of Linnaeus’s three years of travel (1735-1738) and mentioned his early meetings with Herman Boerhaave, a physician and retired director of the Leiden botanic garden, and Jan Frederik Gronovius, a botanist with a large herbarium.  Linnaeus was much younger than them, and he learned a great deal from both, especially because they allowed him to study their specimen collections.  So they deserve more attention in this series of posts on Linnaeus’s travel experiences (Blunt, 1971).

For many years, Herman Boerhaave taught medicine at the University of Leiden and elevated the institution’s stature.  He then headed the university’s botanical garden and worked to increase its holdings of exotic plants.  He was aided in this by his contacts with the Dutch East India Company ( VOC), one of the leaders at the time in trading with Asia.  Following company instructions, surgeons and captains on VOC ships brought back cuttings, seeds, and specimens of plants they encountered on their travels.  Boerhaave was able to add many of these to his garden and herbarium, four volumes of which are now in the Sloane Herbarium at the Natural History Museum, London.  In addition, he published descriptions of new species and built on the work of botanists such as John Ray and Joseph de Pitton Tournefort in attempting to develop a natural classification system (Rutgers, 2008).  It is no wonder that with this background Boerhaave appreciated what Linnaeus was attempting to do with his Systema Naturae, which he had already sketched out by the time he went to Leiden.

Jan Gronovius was a student of Boerhaave’s.  He was an avid specimen collector and kept up a wide correspondence with naturalists in Europe and beyond.  It was through this network that he obtained John Clayton’s specimens from Virginia (see figure above).  Clayton became interested in botany and plant collecting after meeting Mark Catesby on his second trip to the American Southeast collecting for what became the impressive The Natural History of Carolina, Florida, and the Bahama IslandsAfter Catesby returned to England, Clayton shipped him specimens, which Catesby then passed on to Gronovius.  Eventually Clayton sent specimens and letters directly to Gronovius.

At this time, “sending a letter” across the Atlantic could mean waiting months to a year or more for a response, if indeed a response ever came.  Also at that time there was great interest in North American plants and not only because of their novelty.  Since the climate there was temperate as was that of Europe, species were more likely to acclimatize well and could be introduced into gardens.  Wealthy landowners were clamoring for the latest novelties, and botanists wanted to be the first to describe new species.  This helps to explain why Gronovius published a book, Flora Virginica, based on Clayton’s manuscript and specimens without letting him know about it ahead of time and gaining his permission.  This sounds rather dubious, but he did credit Clayton with finding the plants and sending him information on them along with the specimens.  Also, later observers have noted that because Gronovius was so well connected, his publication likely made Clayton’s work more broadly known than if Clayton himself had written on them.  As a case in point, Gronovius allowed Linnaeus to study the Clayton specimens, and so they became type specimens for a number of the North American plants Linnaeus described in Species Plantarum.  Linnaeus spent the winter of 1737-1738 with Gronovius right before returning to Sweden.  They worked on Clayton’s 1737 shipment of plants, to which they gave Linnaean names, a very early use of his system.

Gronovius was also in touch with another American botanist, John Bartram in Philadelphia.  They were originally connected by Bartram’s patron in England, Peter Collinson, another adept networker.  Bartram sent material to Gronovius, who again allowed Linnaeus to examine it.  This was later than with the Clayton material; Linnaeus by then had his long-term academic position in Uppsala and the two sent packages of specimens back and forth between them.  Eventually Gronovius and Bartram corresponded directly, as did Gronovius and Cadwallader Colden, a New York naturalist whose daughter Jane Colden was also involved in botany and produced an illustrated manuscript on New World plants (Colden, 1963).

One last name that should be mentioned as a Linnaean mentor is someone of his own age whom he had worked with while studying at the university in Uppsala.  There they planned to develop a system to organize all living things.  They divided up different groups between them.  For example, Linnaeus opted for most of the plants, and Peter Artedi selected fish and the Brassicaceae as among his favorites.  Finishing their studies, they went their separate ways, then met by chance in Amsterdam and took up where they left off.  Unfortunately, Artedi soon drowned in one of the city’s canals.  Linnaeus saw to the publication of Artedi’s manuscript on fish, and the approaches they developed to classification greatly influenced Linnaeus’s future work.  This is one of those cases where it’s interesting to speculate on what they would have achieved if they had been able to work together for years.

While the three individuals discussed here were important to Linnaeus’s career, it could be argued that the most important individual of his Netherlands sojourn was George Clifford with whom Linnaeus lived and worked for over two years.  Clifford will be the subject of the next post.


Blunt, W. (1971). The Compleat Naturalist: A Life of Linnaeus. New York, NY: Viking.

Colden, J., Rickett, H. W., & Hall, E. C. (1963). Botanic Manuscript of Jane Colden, 1724-1766. New York: Garden Club of Orange and Dutchess Counties.

Rutgers, J. (2008). Linnaeus in the Netherlands. TijdSchrift Voor Skandinavistiek, 29, 103–116.

Linnaeus in the Netherlands


1 Systema Naturae

Title page of Carl Linnaeus’s Systema Naturae (1735), courtesy of the Biodiversity Heritage Library.

There is a great deal of talk about the European Union these days, and the advantages of open travel among nations.  Freedom of movement is a wonderful concept in any age, and it’s one experienced by Carl Linnaeus (1707-1778) when he was in his late 20s.  Having completed his education in Uppsala, Sweden and having become engaged to a woman whom he very much desired, he set out for three years of study and travel.  This wasn’t entirely his own idea.  His future father-in-law was not thrilled with his daughter’s beloved, a physician with few financial resources, so he would only bless the match by having Carl agree to a three-year hiatus.  Linnaeus might not have been a man of means, but he was a man who had already learned a great deal about botany and had developed original ideas about how plant diversity should be organized.  He also had some experience of travel having spent a few months exploring Lapland, the northern reaches of Scandinavia.  So in 1735 he took his manuscripts, packed his bags, and headed to the Netherlands, traveling through Germany on the way.  His experiences in Holland and elsewhere in Europe did a great deal to form his ideas and shape his career.  This series of posts will look at some of those influences (Blunt, 1971).

It seems that Linnaeus did not make a good first impression on many people.  There are a number of stories about men who were put off by his self-possessed manner, and then, as they realized what a good mind lurked behind the bravado, became good friends with him.  This was the case with Johannes Burman, a professor of botany and director of the Amsterdam botanic garden.  Burman, who was the same age as Linnaeus, had been at the garden for several years working on the Flora of Ceylon, using primarily the herbarium of Paul Hermann, who had collected there in the 1670s.  After this brief meeting where Burman was unimpressed by Linnaeus, it would probably have been difficult for either of them to predict that they would be lifelong friends.  At this time Linnaeus also visited Albertus Seba who had amassed a large cabinet of curiosities including materials he collected on trips to the East and West Indies.  During these years the Netherlands was an important naval power with far-flung mercantile interests, so along with trade goods—like spices and silks, exotic plants, animals, and artifacts also poured into Dutch ports.  Even though Seba had sold his original massive collection, he was able to build another and showed some of it to Linnaeus on two visits to his home.  He later asked Linnaeus to assist him in preparing a book he was writing on his holdings, but by then the Swede had made other connections (Rutgers, 2008).

Linnaeus next spent two weeks in Harderwijk, the site of a university where for a week’s residency he qualified as a doctor, submitting a thesis he had written in Sweden.  Then he went to Leiden and showed his manuscript of Systema Naturae to the Dutch botanist Jan Frederik Gronovius, who was so impressed with the work that he arranged for its immediate publication as a thin volume of 14 pages that set out the rudiments of Linnaeus’s taxonomic system.  Gronovius also gave him a letter of introduction to Herman Boerhaave, who had retired as head of the Leiden Botanic Garden.  As with Burman, their relation did not begin smoothly, but eventually Boerhaave appreciated Linnaeus’s intelligence and energy.  However, none of these meetings landed him a position where he could earn enough money to allow him to remain in the Netherlands.  He told them that he would have to return home.  That’s when Boerhaave offered to fund a trip to Cape Town, South Africa which was then under Dutch control and was proving to be a botanically rich area.  Linnaeus, however, after his Lapland expedition, did not much relish a long journey with many probable hardships; Sweden was a safer and easier option.

There are more twists to this story.  On his way home, Linnaeus stopped in Amsterdam and again visited Burman, this time with a letter of introduction from Boerhaave.  Burman paid more attention to his visitor, especially after Linnaeus was able to identify a rare plant Burman showed him.  The latter offered to pay Linnaeus for helping to prepare the Flora of Ceylon, and also convinced him that he should definitely call on George Clifford, a wealthy merchant and horticulturalist who lived near Haarlem.  Clifford and Linnaeus got on well because Linnaeus identified many of his hosts’ Indian plants and was sorely tempted by Clifford’s offer to live and work on his estate, with access to his garden and herbarium.  But Linnaeus was committed to Burman.  In the end, Burman and Linnaeus visited Clifford, and Burman agreed to free Linnaeus if Clifford would give him a very desirable book displayed in his library: the second volume of Hans Sloane’s Natural History of Jamaica.  Clifford and Linnaeus were both very fortunate, with the gardener/financier getting an expert to bring order to his collection of specimens, properly name his plants—those in the herbarium and those in the garden—and help in producing a catalogue to make public his botanical treasures.  Linnaeus, on the other hand, was freed of economic worries, had a very comfortable place to live, and great resources to work with, including a first-class library.  What happens then will be the subject of a later post.


Blunt, W. (1971). The Compleat Naturalist: A Life of Linnaeus. New York, NY: Viking.

Rutgers, J. (2008). Linnaeus in the Netherlands. TijdSchrift Voor Skandinavistiek, 29, 103–116.

Natural History in 17th-Century Britain: Nehemiah Grew

4 Grew plum

Transverse section through plum branch, from The Anatomy of Plants, Biodiversity Heritage Library

When I think of Nehemiah Grew (1641-1712), an image of a cross section through a stem appears in my mind’s eye (see above).  I remember Grew as the creator of magnified images of plant tissue that have a rather inorganic feel to them in their rigid rows of cells.  He was doing microscopic studies at about the same time as Robert Hooke, whose illustrations of plant cells are less regular, and somehow have a more organic feel.  They were both attempting to communicate the new world they were exploring and trying to make sense of it.  In an article to commemorate the tercentenary of Grew’s birth, the plant morphologist Agnes Arber (1941) noted that Grew held a mechanistic view of the universe and saw the microscope as a way to clear up mysteries of life by observing its constituents.  Because of this viewpoint he also developed mathematical descriptions and was concerned with how to communicate the scale of objects seen under the microscope.  This is a reminder that at the time there were no adequate standards of measurement for the microscopic world.  So Grew used comparison to give his reader some idea of the size of what he was seeing, for example, that something was one-fifth the size of the cheese mite or the width of a marsh mallow seed.

Another major contributor to the beginnings of plant anatomy was the Italian Marcello Malpighi who like Grew was a physician, though Grew practiced medicine while Malpighi taught in a medical school and also did a great deal of research on animal anatomy.  In fact, he began studying plants because he found animal tissue so complex and wanted to see if the “simpler” structures of plants could give him clues.  Grew and Malpighi are usually mentioned together because in some ways, their work is similar.  They labored independently without any knowledge of the other’s research until Grew produced a paper for the Royal Society of London (RS) shortly before Malpighi sent a manuscript read at an RS meeting.  After that they followed and cited each other work.

The consensus is that they achieved similar results.  Alan Morton  (1981) claims that Malpighi saw more clearly than Grew in some details, but Grew’s culminating The Anatomy of Plants is the fuller and clearer work, with a more integrated view of plant structure.  Agnes Arber (1942) also wrote a comparison of their contributions and contends that Grew may be better known because he wrote in English, while Malpighi published in Latin.  Arber notes:  “His Latin, though lively, is not very correct, and its interpretation is often by no means easy” (p. 15).  But she considers Malpighi’s illustrations, made from his drawings, as excellent.  Some of Grew’s illustrations are noteworthy because, while Malpighi made attempts at depicting microscopic structures in three dimensions, Grew did it more successfully.

Since this set of posts is on British botanists, I’ll end by noting some of Grew’s most important findings, though in many cases, Malpighi also produced similar results.  Grew described the main anatomical differences between roots and stem.  This required a great deal of work examining a variety of different species.  The same was true of deciphering the vascular network in these structures.  Grew admits that he got the idea for the spiral form of vessels from Malpighi, but he came up with the name “parenchyma” for the material packed around the vessels.  While he depicted a great deal of order in plant tissue as orderly, he did not really conceive of cells as the basic unit of plant structure, though Robert Hooke had already coined the term for the structures he saw in cork cambium.  Grew was able to differentiate between the scattered vascular bundles in monocots and the more ordered structures in dicots; he identified the medullary rays in dicot stems as well.  Grew compared the cell walls to woven fibers and more generally compared a plant’s inner structure to a textile fabric.  Arber (1913) quotes a long passage where he describes plant tissue as fine lace with an intricate and delicate texture.  At one point he writes:  “One who walks with the meanest Stick, holds a Piece of Nature’s Handicraft, which far surpasses the most elaborate Needle-Work in the World” (p. 54).

Grew also went into detail on the structure of flowers.  Though he accepted the idea of sexual reproduction in plants, he wasn’t able to work out the process.  He presented much information on seed structure in a variety of species and carefully observed seed development, coining the term radicle for the embryonic root.  He also did simple experiments on the movement of sap, but his major work was anatomical.  For a century and a half after Grew and Malpighi there was little further development in the field.  It may be that it had to wait for the creation of better microscopes, or for further work in plant physiology.  Or perhaps some of the lack of interest in the field may have been due to those images that so intrigue me.  They presented such a well-developed, finished view of plant structure, that others might have considered the job of working out plant architecture to be complete.  After all, plants were simpler than animals, how much more was there to know?


Arber, A. (1913). Nehemiah Grew (1641-1712). In F. W. Oliver (Ed.), Makers of British Botany (pp. 42–64). Cambridge, UK: Cambridge University Press.

Arber, A. (1941). Tercentenary of Nehemiah Grew (1641-1712). Nature, 147(3734), 630–632.

Arber, A. (1942). Nehemiah Grew (1641-1712) and Marcello Malpighi (1628-1694): An essay in comparison. Isis, 34, 7–16.

Morton, A. G. (1981). History of Botanical Science. New York, NY: Academic Press.

Natural History in 17th-Century Britain: John Evelyn

3 Evelyn tree

Plate from Sylva of collecting birch tree sap, Biodiversity Heritage Library

While John Ray, the subject of the first post in this series, is an important figure in the history of botany, this post’s subject would be considered more a horticulturalist than a botanist and is best remembered for Sylva, his book on trees and how to grow them.  However, John Evelyn (1620-1706) published a number of other books, including several translations.  After studying at Oxford, he trained in the law, but the disruption of the English Civil War led him to spend several years on the Continent, visiting botanical gardens in Paris, Leiden and Padua, where he purchased a herbarium.  Twenty years later, he showed his own to his friend, the diarist Samuel Pepys, who had never seen one before and was taken with the way a plant’s characteristics were so clearly preserved.

While in Europe Evelyn also toured private gardens to broaden his understanding of horticultural design.  This need for information led him to translate two French works, The French Gard’ner by Nicolas de Bonnefons (1658) and later, Jean-Baptiste de La Quintinie’s The Compleat Gardner (1699)  Translation was a way to not only disseminate ideas, but to understand them better.  As was common, Evelyn added commentary and in the case of the latter work, also other writing by Quintinie not in his original book.  Publishing at the time was looser than today in that authors used the opportunity to pack as much into a book as possible and were less concerned about cohesiveness.  Evelyn also translated books from the French on other subjects including painting and architecture.

Evelyn returned to England with the restoration of the monarchy and the convening of a new parliament in 1661.  He was also relieved that the Church of England, of which he was a devote member, was again legitimized.  Evelyn became involved in several government projects at the behest of the king.  In addition, the spirit of renewal led to plans for the creation of the Royal Society of London (RS) for the advancement of science based on the writings of Francis Bacon.  They aimed to promote empirical studies, the collection of information on a subject thorough enough to allow for analysis and firmly based conclusions, in other words, inductive reasoning.  Evelyn was engaged in the organization of the society and delivered a paper on forest trees at an October 1662 meeting.  This became the first formal publication of the society, Sylva, or A Discourse of Forest-trees.  He presented descriptions of a number of species, but since the society was interested in practical outcomes from science, he also argued for restoration of forests in Britain where they had been drastically retracting over the centuries.  If the country was to remain economically viable and become a world power then its navy and its industry required timber.

As part of his involvement in the RS, Evelyn was a member of the “Georgical” Committee, named after Virgil’s horticultural text, the Georgics.  This group worked for the improvement of English agriculture, horticulture, and landscape.  At the same time, Evelyn was improving his own garden on property he leased from his father-in-law.  This involved extensive reworking of the garden’s organization with the planting of allées of trees.  He was also interested in the kitchen garden and was particularly taken with salads, even writing a book on the subject.  He also wrote a book on fruit trees (1706) and another on what seems a 21st-century topic:  the use of plants to deal with air pollution.  Called Fumifugium, it dealt with among other topics fragrant plants whose scents would compete with the stench of the city.

With all this endeavors, Evelyn never completed his largest project, an encyclopedia of British gardening, Elysium Britannicum.  By the late 1650s, he already had an outline for the work and sent it to several friends for their comments.  All urged him to continue with it, but it was a huge undertaking.  He wanted to cover every aspect of the subject from garden design, to how to manage its development and maintenance.  Evelyn was a member of the upper class so he focused on large-scale gardens, not those surrounding a cottage.  In the 17th century, the interest in plants that had emerged in the previous century developed into an industry, with professional gardeners and nurserymen providing services to wealthy landowners.  Evelyn took this into account, but he was still a hands-on gardener interested in the growth habits of individual species as well as larger issues.  He even planned a chapter on why and how to create a herbarium as a reference for what was growing in the garden.

In John Evelyn: A Life of Domesticity, John Dixon Hunt (2017) explores several possible reasons for why Evelyn never finished the project.  All that remains are manuscripts of the original outline as well as parts of the first of three projected sections.  Not surprisingly, Hunt sees the size of the project as so massive it discouraged Evelyn who was occupied with family issues as well as his work with the government and the RS.  Though he was a man of means, he would have needed financial as well as technical support in completing the manuscript and producing the illustrations.  Hunt also conjectures that Evelyn felt a sense of guilt about his preoccupation with gardens which seemed such a worldly pursuit for a man whose religious beliefs led him to focus on the otherworldly.  Despite this failure, Evelyn remains a symbol of love of gardening, and especially love of trees.


Bonnefons, N. de & Evelyn, J. (1658). The French Gardiner. London, UK: John Crooke.

Evelyn, J. (1706). Pomona. London, UK: Scot, Chiswell, Sawbridge and Tooke.

Hunt, J. D. (2017). John Evelyn: A Life of Domesticity. London: Reaktion.

La Quintinie, J. de, & Evelyn, J. (1699). The Compleat Gard’ner. London: M. Gillyflower.

Natural History in 17th-Century Britain: Francis Willughby and Martin Lister

2 Lister shells

Plate from Historiae Conchyliorum, Biodiversity Heritage Library

In this series of posts, I’m looking at the lives of a number of British naturalists active during the early years of the Royal Society of London (RS), of which they were all members.  This week’s post deals with two men who are not known for their work in botany, so why would I even give them a second thought?  Like most naturalists of the time, they did not restrict themselves to a single group of organisms.  Particularly in their early years, they delved into the plant world and spent time with John Ray, the subject of the last post.  Also, Francis Willughby (1635-1672) and Martin Lister (1639-1712) are both subjects of new books on their areas of expertise:  Willughby in ornithology (Birkhead, 2018) and Lister in conchology (Roos, 2018).

Ray and Willughby worked closely together for a number of years.  They met at Cambridge University where Ray tutored Willughby and became good friends who often went on collecting trips together, several through Britain that lasted for weeks or months.  Eventually, they toured Europe for a year and half, visiting the Low Countries, Italy, and France, often accompanied by one or two other naturalists.  While I focused on Ray’s interest in plants in my post, he shared some of Willughby’s passion for insects, birds, and fish.

Willughby had the money to collect art depicting interesting species, including entire albums, that are still housed at his family’s estate.  Also there is one of the two specimen cabinets he ordered while in Europe, and it contains some of his collection of bird eggs as well as drawers of seeds.  In the most personal passage in his book, Tim Birkhead describes his thrill in sifting through this treasure chest and finding notations in Willughby’s handwriting.  Ray and Willughby themselves worked their way through Ulisse Aldrovandi’s massive natural history collection in Bologna, attempting to identify some of the new species they had encountered on their travels.  Eventually the two went on to Montpellier and collected plants in the area.  Some of these specimens are in Ray’s herbarium, now part of the Sloane Herbarium at the Natural History Museum, London.

The two continued to collaborate after they returned to England in 1666.  Willughby was supporting Ray after Ray lost his position at Cambridge because he refused to take an oath of loyalty to the king.  This arrangement lasted until Willughby’s death in 1672.  Before then, they worked on Willughby’s interests in insects, birds, and fishes with an eye toward publications on all three, though none had been completed before he died.  What made it difficult for Ray to complete them is that the family would not give him access to Willughby’s specimens or his notes.  Ray had to rely on what he had written about the collections.  The fish book was farthest along and was published first, while the insects needed the most research and came out last.  But the most original work, with the most first-hand information, was in ornithology.

In his biography of Willughby, Birkhead makes the argument that the importance of the Willughby/Ray publication on birds was due much more to Willughby’s contribution.  He takes a stand against that of Charles Raven (1950) in his Ray biography where he considers Willughby the junior partner, and Ray the genius of the project.  To this day, Willughby’s descendants fume about what they see as the injustice of this viewpoint, and Birkhead makes a good case for their opinion, despite the fact that little remains of Willughby’s notebooks and manuscripts.

While Ray was in Montpellier, he also collected with Martin Lister, who was spending three years there as a student.  It’s not surprising that fellow countrymen, particularly with shared interests, would gravitate toward each other.  However, they had to leave France in 1666 by order of the French king, in anticipation of a war between France and Britain.  Once back in England, Ray spent three months botanizing with Lister and the apothecary Peter Dent.  Lister was particularly interested in grasses, insects, and spiders, but eventually became enthralled with mollusks and focused on conchology, the study of their shells.  He wrote a three-part natural history of spiders, terrestrial and river mollusks that was illustrated by William Lodge.  However, when he set out to create a more comprehensive work on mollusks, Lodge was not particularly interested, so Lister decided to teach his two oldest daughters, Susanna then eleven years old and Anna aged nine to draw.  Eventually they even learned to engrave.

One of the reasons I’ve decided to include Lister here is because some of his experiences in studying mollusks are similar to those of botanists.  A number of the latter also used the talents of family members, usually females, in creating illustrations.  From the notes and drawings left by the Listers, it’s clear that in their case, the sisters made very careful observations, sometimes using a microscope, to work out mollusk anatomy; their drawings weren’t limited to shells.  In the last edition of Historiae Conchyliorum of 1692, there were 1067 plates.  In addition, Susanna created illustrations for papers by other authors in the RS’s Philosophical Transactions.  Roos’s book does a good job of presenting the work of her and her sister as integral to the success and value of her father’s opus.


Birkhead, T. (2018). The Wonderful Mr. Willughby. London, UK: Bloomsbury.

Raven, C. E. (1950). John Ray Naturalist: His Life and Work. Cambridge, UK: Cambridge University Press.

Roos, A. M. (2019). Martin Lister and His Remarkable Daughters: THE Art of Science in the Seventeenth Century. Oxford, UK: Bodleian Library.

Natural History in 17th-Century Britain: John Ray

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Title page of John Ray’s Historia Plantarum, from the Biodiversity Heritage Library

I’ve written a number of posts about 16th-century naturalists who opened the field of early modern botany, especially about Luca Ghini, an early advocate for the herbarium, and those who worked with him (1,2,3,4).  In this series, I want move on to the 17th century and discuss a circle of naturalists working in Britain.  They knew each other, as least causally, in part because they were all members of the Royal Society of London (RS), which was founded in 1660.  This was in the latter days of a turbulent time in Britain when kings had been killed, Oliver Cromwell took over the government, and finally the monarchy was restored by bringing Charles II to the throne.  Part of this political turmoil was religious as well, with Catholics and different Protestant factions at odds with each other—and with the government.

But what does this have to do with plants?  Well, quite a bit, because many of those interested in natural history were products of Cambridge and Oxford Universities, which were then religious institutions devoted primarily to training clergy.  They were affected by the political upheavals, especially when Cromwell.  When King Charles II reached the throne, those working at the universities was asked to sign a loyalty oath to the Church.  John Ray (1627-1705), a naturalist who had been teaching at Oxford for 13 years, refused and lost his job.  He responded by not only leaving the university but the country and spent the next three years traveling in Europe, most of the time with Francis Willughby  and Philip Skippon, both of whom he had tutored.

Ray and Willughby had already made several collecting trips to areas in Britain, and Ray had published a Flora of Cambridgeshire (1660), the first of its kind for the British Isles and one that served as a model of such books.  This came to my attention recently when I read Tim Dee’s (2015) Four Fields, one of which is in Cambridge where he lives.  Dee writes of Ray’s work:  “I can think of nothing more thrilling, nothing that our species has done better, than this benign capture and permanent vivifying of a season, a pathway and a field edge, and its simpling, or its lovable mapping of what might be in front of us” (p. 236).  In other words, Ray makes the nature of Cambridge come alive.  Early his book, Dee himself writes that as a child he was enchanted by the world of books and found that books about the living world made that world more vivid and real.

Ray’s interests, like those of most naturalists of his time, extended well beyond plants.  With Willughby, he investigated birds, insects, and fish, publishing the results of their work after Willughby’s early death.  There will be more on Willughby in the next post.  For now I want to stick with Ray’s major work, his massive three-volume Historia Plantarum (1686-1704).  Agnes Arber (1943) suggests that its size, as well as its Latin text, led to its lack of popularity, but it’s nonetheless an important resource.  Ray is credited with one of the best pre-Linnaean classification schemes.  He built on the much earlier work of Andrea Cesalpino and still divided plants into trees, shrubs and herbs, but he also differentiated between monocots and dicots, and between angiosperms and gymnosperms.  These were not totally new discoveries, and much of the terminology came from the writings of Joachim Jung.  But Ray’s genius was in gathering all this information together and presenting it in a clear, organized way.  His work is considered one of the forerunners of Antoine Laurent de Jussieu’s natural system of classification.

Alexander Wragge-Morley (2010) argues that Ray’s descriptions were a form of picturing, they “enjoyed the same epistemic status as graphic representations, because they provoked images—and knowledge—of the same sort.  An image revealed immediately, a verbal description, more slowly” (p. 174).  Ray himself wrote that images can enhance the intelligibility of text, but can’t replace it because there is information about a species that can’t be conveyed in an engraving.  This was an opinion Ray shared with others like Robert Hooke and Nehemiah Grew, both of whom did use illustrations in their publications.  One reason Ray didn’t was their cost, but he also seemed to gravitate toward words.  He didn’t think much of herbaria, though there is one in the Sloane Herbarium at the Natural History Museum, London containing plants he collected on his European tour.  He doesn’t seem to have kept specimens relating to his Historia, though he did examine plants in a number of herbaria including those of Hans Sloane, Leonard Plukenet, and James Petiver.

Ray wrote a number of other works, including some I’ll mention in the next post on Francis Willughby.  He also produced a collection of translations of travel writings by authors who had toured parts of the Middle East.  The major portion was a translation by a German writer into English of Leonhard Rauwolf’s travelogue, noting the many plants he encountered along the way (see earlier post).  Ray also wrote a theological tract.  Throughout his life he remained religiously fervent, like many of his day, and saw the study of nature as a way to learn more about the creator.


Arber, A. R. (1943). A seventeenth-century naturalist: John Ray. Isis, 34, 319–324

Dee, T. (2015). Four Fields. Berkeley, CA: Counterpoint.

Ray, J. (1660, 1975). Ray’s Flora of Cambridgeshire. Hitchin, UK: Wheldon and Wesley.

Wragge‐Morley, A. (2010). The work of verbal picturing for John Ray and some of his contemporaries. Intellectual History Review, 20(1), 165–179.

Vicki Funk: Thinking Big about Collections

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This is a last in a series of posts [1,2,3] on the plant systematist Vicky Funk and her recent review article on collections-based research.  Since Funk is a research scientist and curator in the National Museum of Natural History’s (NMNH) Botany Department, it isn’t surprising that she begins a section on the future use of collections with stats on herbaria.  The NMNH, part of the Smithsonian Institution, is home to the U.S. National Herbarium, with a collection of over five million specimens.  The goal there and at many herbaria is to digitize the data for all specimens and in some cases to also image them.  If this could be done at every herbarium, the data would serve as a potent research tool not only for taxonomists but for ecologists, conservationists, and researchers in other fields who never before considered using the information about plants available in herbaria.

One burgeoning field based on the availability of digital specimen images is computer vision and machine learning techniques that make automated plant identification possible.  It is sort of face recognition for plants and is developing to the point that herbarium specimens can be sorted rather well, though the processes are hardly at the point where identification is as good as that done by taxonomists.  However, machine sorting could be employed as a way to narrow down the number of specimens a researcher would have to look at in hunting for new species.  One recent report the computer was able to distinguish between moss groups better than the human eye could.

Funk cites several successful digitization projects, noting that the Atlas of Living Australia is a particularly comprehensive one that has resulted in online access to all records of Australian plant specimens held in the country’s national herbaria.  Australia is also at the forefront in developing software tools to assist researchers in extracting as much information as possible and in the most effective ways.  However, Funk sees the future as going beyond national or even regional databases:  “A Central Portal so all resources are available to everyone is critical.  It is particularly important that these efforts are making the data and images available to researchers in the countries where the specimens were collected, thereby supporting research in those countries” (p. 185).  She is referring to the fact that the bulk of specimens collected in developing countries, particularly during their colonial pasts, are held in European and North American herbaria.  A first attempt to make these specimens broadly available was the Andrew W. Mellon Foundation funding of type specimen digitization, the results now accessible through JSTOR Global Plants along with a great deal of supporting botanical literature.

But what Funk visualizes is something more comprehensive, and as an example, she describes a project funded by the Powell Center of the US Geological Service.  It focuses on the approximately 2500 species of North American Compositae (Asteraceae) and the location data on hundreds of thousands of specimens aggregated from GBIF (includes information from institutions outside the US), BISON (from US government institutions) and iDigBio (US private institutions).  Funk notes that this data is not only aggregated but “cleaned” to make sure it is of high quality, an issue that critics of aggregation emphasize.  The data is then integrated with environmental and geophysical data on geochemistry, climate, topography, etc., as well as phylogenetics—including gene sequences from GenBank.  Think of the power of this:  linking specimens with sequence and environmental data.   This is truly a harbinger of a new age in collections-based research.  It is amazing that ten years ago, just digitizing data and imaging specimens was considered a feat, with the Paris Herbarium’s plan to digitize most of its specimens considered daring.  Now the assembly line method they used has become relatively common, and other large herbaria have substantial percentages of their collections digitized and imaged.

Linking natural history collections to genetic data banks means uniting the two great arms of bioinformatics.  It is a biologist’s dream come true, and this connection will become even more powerful when environmental data is brought into the mix—a much more complex process.  But Funk has seen the digital world burgeon and has been one of the forces behind making it applicable to systematics.  She has also helped make systematics valuable to other fields such as phylogenetics and the growing discipline of phylogenomic—being able to sequence and compare entire genomes.  This is the result of new sequencing techniques that utilize fragmented DNA, just the type available in herbarium specimens.  Drawing on an example from the Asteraceae, Funk cites a study in which the entire genomes of 93 of 95 Solidago, goldenrod, herbarium specimens were sequenced with the plants ranging in age from 5-45 years (Beck & Simple, 2015).

In closing Funk notes:  “One exciting trend is the developing field of Integrative Systematics where collections-based systematics is combined with extensive field studies, phylogenetics, phylogenomics, detailed morphological studies, biogeographic inferences and diversification analysis to present a more comprehensive global” (p. 187).  She also argues for the maintenance of collections in educational institutions to insure the instruction of future generations of systematists; the digitization of cleared leaf slides, anatomy slides, pollen images, chromosome count images, and illustrations to fill out the information available to researches; and finally a series of symposia on the Tree of Life where systematists can map out a research agenda for the rest of the 21st century.


Beck, J. B., & Semple, J. C. (2015). Next-Generation Sampling: Pairing Genomics with Herbarium Specimens Provides Species-Level Signal in Solidago (Asteraceae). Applications in Plant Sciences, 3(6), 1500014.

Vicki Funk: The Age of Tree Thinking

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In the last post, I began my discussion of Vicki Funk’s (2018) recent article on “Collections-Based Science in the 21st Century” that I’ll continue here.  In this review, she terms the first 15 years of the 21st century “An Age of Tree Thinking,” in other words a time of investigating evolutionary relationships and the use of phylogenies.  This is a major interest of Pam and Douglas Soltis of the University of Florida, two other leaders in the field of collections-based research (Allen et al., 2019).   Funk gives examples of what she means by this term, beginning with evolutionary medicine.  This field’s work includes tracing changes in viruses as they are transmitted through a population and even within one body over time.  Funk notes that museum specimens of woodrats have been found to harbor viruses similar to those causing Chagas disease.  She also touches on food safety, beginning with GenomeTrakr a pathogen database set up by the Food and Drug Administration.  It hosts whole genome sequences for pathogens, mostly those implicated in food poisoning.  When an outbreak occurs, the pathogen involved can now be quickly sequenced, and then compared to sequences in the database; this helps to identify the source of contamination and speed control of the outbreak.

Moving on to evolutionary ecology, Funk cites a number of examples of how phylogenetics can illuminate ecological questions.  For example, DNA was sequenced from ragweed (Ambrosia artemisiifolia) specimens collected through time, both before and after deforestation in particular areas.  Pollen core data suggest that ragweed, an aggressive weed, was uncommon before deforestation.  The DNA sequencing data indicates that there was a hybridization before deforestation that may have permitted the hybrid to grow more aggressively when trees were removed.  This is a good example of pairing historical data with molecular analysis.

Funk’s paper also explores the idea of DNA barcoding, a technique that her colleague John Kress at the Smithsonian has fostered.  For plants, it involves sequencing two regions of the chloroplast genome that serve as a fingerprint for species identification.  Kress and his colleagues (2009) barcoded all tree species growing in a plot on Barro Colorado Island in Panama, a long-term Smithsonian study site.  The resulting phylogenies are being employed to investigate the relationship between habitat and community structure.  Barcodes are also used to monitor illegal traffic in endangered species, for example, as a way to identify illegal shipments of rare woods.  Since her article’s title, “Collections-Based Science in the 21st Century,” doesn’t limit Funk to only plants, she slips in a reference to molecular phylogenetics in human evolution studies, noting how DNA extracts from fossils of Neandertals and of a hominin population called the Denisovans found in the Siberian Altai Mountains, as well as from present-day humans, were employed to work out the relationship among them, with Neanderthals and today’s humans more closely related to each other than to Denisovans.  In an example relevant to botany, medically important plants have been barcoded over the past ten years, and molecular phylogenetics can be used to test the purity of ingredients in herbal medicines.  This is a perennial problem due to varying levels of quality control for these materials, resulting in impure or ineffective products.

What these examples of tree thinking have in common is that they involve DNA sequencing and the storage of that information so it can be used in future studies.  In other words, there is a summative process going on here, and these databases, if properly maintained and utilized will only become more and more valuable and effective.  In the next section of her article, Funk deals with the future, and calls it “An Age of Thinking Big.”  This theme is also taken up by a group of European researchers (Besnard et al., 2018).  Funk discusses not only collections of DNA sequences, and the voucher specimens that back them up, but also the increasing availability of online data about natural history specimens as well as images of them.  Digitization has been going on for years, especially since the development of BISON, which is a database for specimens from US government facilities such as the Smithsonian, and iDigBio, for private research and educational collections.  While more and more information is coming online, there is still a great deal to do.  To date, less than half of all plant specimens are databased, and that percentage is even lower for animals—there are an awful lot of insects out there, which were relatively easy to collect, but not so easy to image, to say nothing of jellyfish, etc.

Funk considers some of the questions that could be tackled if all specimen data were available to researchers:  “What parts of the world need additional collecting expeditions?  How many species are rare?  How many species have not been collected in the last 50 years and may be extinct?  Are there certain areas that have a lot of rarely collected species and are these areas endangered ecosystems?  How fast have invasive species moved into new areas?  How has community composition changed through time?” (p. 182).  This list is reminiscent of Funk’s “100 Uses for an Herbarium.”  With her vast experience she is very good at thinking about why collections are valuable as research tools, and this analysis is especially useful today as many collections are facing uncertain futures.  In an earlier post I cited one example of Funk’s writing on this topic.  Here I’ll end with another citation, a review article she wrote with several of her colleagues on what collection based systematics should look like in 2050 (Wen et al., 2015).  Her answers to this question will be covered in the next and last post in these series.


Allen, J. M., Folk, R. A., Soltis, P. S., Soltis, D. E., & Guralnick, R. P. (2019). Biodiversity synthesis across the green branches of the tree of life. Nature Plants, 5(1), 11–13.

Besnard, G., Gaudeul, M., Lavergne, S., Muller, S., Rouhan, G., Sukhorukov, A. P., … Jabbour, F. (2018). Herbarium-based science in the twenty-first century. Botany Letters, 165(3–4), 323–327.

Kress, W. J., Erickson, D. L., Jones, F. A., Swenson, N. G., Perez, R., Sanjur, O., & Bermingham, E. (2009). Plant DNA barcodes and a community phylogeny of a tropical forest dynamics plot in Panama. Proceedings of the National Academy of Sciences, 106(44), 18621–18626.

Wen, J., Ickert‐Bond, S. M., Appelhans, M. S., Dorr, L. J., & Funk, V. A. (2015). Collections-based systematics: Opportunities and outlook for 2050. Journal of Systematics and Evolution, 53(6), 477–488.

Vicki Funk: The History of Collections-Based Science

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In the last post, I introduced Vicki Funk, a plant systematist who is a research scientist and curator at the U.S. National Herbarium, part of the Smithsonian Institution’s National Museum of Natural History.  There I mentioned that Funk had recently published a review article called “Collections-Based Science in the 21 Century,” published in the May 2018 issue of the Journal of Systematics and Evolution.  As with most review articles, it begins with a historical perspective.  The first sentence is a bold claim:  “Major revolutions in scientific thought have occurred because of collections-based research” (p. 175).  Funk is in a position to know both because she works in an institution with a premier natural history collection, and because she herself has contributed to today’s revolution in how collections are accessed and utilized.

Funk begins with the age of classification and Carl Linnaeus’s heavy reliance on natural history collections in creating his artificial system of classification and nomenclatural reform.  Michel Adanson and Antoine Laurent de Jussieu, working at the botanical garden in Paris with its notable herbarium, devised natural classification schemes that in various forms eventually replaced the Linnaean artificial system.  The 19th century, Funk notes, began with Alexander von Humboldt’s expedition to Latin America that gave him the perspective to develop the field of biological and physical geography, along with ecology and meteorology.  He and his traveling partner Aimée Bonpland collected 50,000 specimens, documenting many new genera and species as well as the relationship between geography and species distributions.  Later, Charles Darwin, Joseph Dalton Hooker, and Alfred Russel Wallace not only collected specimens but used them to build on Humboldt’s work and to document the concept of species change.  With examples like this Funk makes clear the connection between collection and theory building, as well as the importance of great natural history museum collections, many of which were built in the 19th century.

Funk terms the 20th century the “Age of Synthesis” in reference to the evolutionary synthesis that developed at mid-century and to “four collection-based ideas and methods that changed . . . the way we do science” (p. 178).  The first was the concept of continental drift and with it the idea that land bridges between continents had existed in the past.  Both Humboldt and J.D. Hooker argued for these from the similarities among organisms in areas that are now separated by great distances.  Second was the development of phylogenetic systematics or cladistics, a field to which Funk has contributed a good deal both theoretically (1991) and in terms of her research, especially on the Asteraceae.  Cladistics deals with using derived characters to objectively construct relationships, then grouping taxa so all are descended from a single common ancestor without omitting any of its descendants.  This is a complex field, and as a recent issue of the American Journal of Botany (August 2018) on fossil plants reveals, there are problems that arise when only living species are used in creating monophyletic groups, so fossil collections are crucial to the process.

Under the third 20th-century trend, Funk lists databasing collections, biodiversity science, and niche modeling.  This is a huge triumvirate, but with its parts closely tied together.  Databasing collection data—specimen identification as well as place and time of collection—makes it possible to more easily assess data on the biodiversity of a region as well as on how it may be changing over time.  It also allows rigorous niche modeling, a term for techniques employing occurrence data to model the possible spatial extent of a species based on geographical and climatic data.  Ecology has always been a field using sophisticated mathematical models but the availability of digital data and high-speed computing have caused an explosion in research.  And this is really only the beginning, as more collection data and analytic tools come online.

The final concept Funk cites as developing in the 20th century is molecular phylogenetics, the analysis of gene sequences as a way to discover phylogenetic relationships.  She writes:  “Collections are an excellent source of material for the extraction of DNA, but they are also important because they provide the vouchers of the DNA sequences, and their presence allows us to check the identification of samples and to gather the data needed to ask questions about character evolution and modes of speciation” (p. 180).  These vouchers usually contain at least some geographic information, bringing in the biogeography she mentioned earlier.  Molecular systematics helped to clear up some arguments about derived characters used in cladistics and resulted in a major reorganization of plant phylogenetics.  As will become apparent in the next two posts, sequencing techniques have changed rapidly during the latter part of the 20th and into the 21st century, increasing the efficacy of DNA analysis with herbarium specimens.  These tools now allow sequencing of species for which no fresh material is available because the species are rare, inaccessible, or even extinct.  If historical material is available, they also enable work on how the genetics of a species may have changed over the last few hundred years.