This and That: Travels of Sophora toromiro

2 Sophora

Toromiro, Sophora toromiro (Phil.) Skottsb, collected 28 June 1800, H. Herrenhus. [possibly Hannover Herrenhausen Royal Gardens], Germany. CC BY-NC-ND 4.0. Te Papa (SP107845)

As with most of the posts in this series of miscellanea (see last post), this story begins with a Tweet, one linked to a blog post and a research article connecting four countries over 250 years.  I’ll try to keep it as simple as possible, by starting in the middle.  In 1877, James Hector, director of the Colonial Museum in New Zealand (now the Museum of New Zealand: Te Papa), asked the British Museum (BM) for a collection of European plant specimens to compare with European plants colonists had brought into the country and were now flourishing, sometimes to the point of being nuisances.  Hector received 28,000 specimens collected by three British amateur botanists: a husband and wife, Silvanus and Bridget Thompson, and Thompson’s student, James Baker.  Most specimens were from cultivated plants gathered in German botanic gardens and the Cels nursery in France between 1764-1864.  Hector never got around to sorting through this gift from the BM; it remained in its original packaging until the 1950s; even today, the only vascular plants to be processed are the orchids.

Recently six specimens of Sophora were found in the collection.  Sophora is a small genus of 17 species in the Fabaceae family and native to the South Pacific.  With eight species, New Zealand is its center of diversity, hence the interest in these sheets that were dated from 1796 to 1822 and were presumably from cultivated plants.  This was surprisingly early for Sophora to be growing in Europe.  Until now, it was thought that the Sophora in Europe were all descended from seeds collected from the 1920-1950s.  There was little plant collecting in the South Pacific until the early 1800s, though Joseph Banks and Daniel Solander had gathered seeds of two Sophora species on Captain James Cook’s first round-the world voyage.  These were planted at Kew by 1772, and there were a few other early cultivations.

The six specimens of interest are in the herbarium of the descendent of the Colonial Museum, the Museum of New Zealand, with the Maori name, Te PapaCarlos Lehnebach, botany curator, and Lara Shepherd, research scientist specializing in DNA sequencing, decided to learn more about the genetics of these six specimens from the 19th-century BM gift.  When Shepherd got the results of her analysis, she was shocked:  one of the specimens, collected in 1800, had genes of Sophora toromiro, a species endemic to Easter Island, Rapa Nui.  It became extinct in the wild in the 20th century, though it is cultivated at several botanic gardens.  At first Shepherd couldn’t believe the results, but when she and Lehnebach looked at the specimen, they found that it did in fact have characteristics of the Rapa Nui plant.  But how did it end up growing in Germany in 1800?

The researchers speculate that seeds may have been collected during Captain Cook’s second round-the-world voyage (1772-1775), when the expedition stopped at Rapa Nui.  The botanists on that trip were Johann Reinhold Forster and his son Georg, with the Linnaean pupil Anders Sparrman as their assistant.   They were the first Europeans to collect specimens on the island, and Sparrman was known to have collected seeds.  He may very well have collected them from this plant, since it grew in thickets and was the only native shrub on the island.  If S. toromiro seeds were planted in the late 1770s, then the shrub would have been established enough to yield cuttings in 1800.  In looking for other Sophora specimens, Lehnebach and Shepherd have found one at the herbarium of the Botanic Garden and Botanical Museum Berlin-Dahlem that could be S. toromiro.  It has no collection date, but it is part of Carl Ludwig Willdenow’s (1765-1812) collection, and a large number of Forster specimens were included in it.  Willdenow had one of those bad habits that frustrates later curators:  he removed the old labels and replaced them with his own, often neglecting to transcribe what’s now considered essential information.

Admittedly, there are suppositions holding this story together, but further work, including analysis of chromosomal DNA from the Willdenow specimen, may make the picture clearer.  In any event, this case study presents a good argument for curating specimens that have been moldering in boxes for decades if not centuries.  This situation is not the result of bad management but of overworked curators without time to deal with the substantial work involved in mounting specimens and providing them with up-to-date identifications.  However, this example suggests the exhilaration that can result from the effort.  Though not every find is a jewel, that’s true of cleaning out any attic.  However, one never knows when a first edition book or a valuable art work might come to light.  My favorite statistic at the moment is that when the herbarium at the National Museum of Natural History in Paris was cleared out prior to renovations about 10 years ago, 830,000 unmounted specimens were found.  Most of them have since been mounted by an outside contractor called in for the massive job (Le Bras, 2017).  But the specimens still need to be curated and filed, a job that amounts to organizing a good-sized herbarium.

Reference

Le Bras, G., Pignal, M., Jeanson, M. L., Muller, S., Aupic, C., Carré, B., Flament, G., Gaudeul, M., Gonçalves, C., Invernón, V. R., Jabbour, F., Lerat, E., Lowry, P. P., Offroy, B., Pimparé, E. P., Poncy, O., Rouhan, G., & Haevermans, T. (2017). The French Muséum national d’histoire naturelle vascular plant herbarium collection dataset. Scientific Data, 4(1), 1–16. https://doi.org/10.1038/sdata.2017.16

Getting the Most Out of Herbaria: Systematics and Chemistry

2 Florid

Murder Most Florid by Mark Spencer, London: Quadrille, 2019

As mentioned in the last post, herbaria, both real and virtual, are most frequently visited by taxonomists, who are usually studying particular plant taxa or preparing flora of areas ranging in size from city parks to entire countries.  These are the traditional uses of plant collections and are still crucial.  However, several things have changed.  Now the “visit” is often to digital portals rather than onsite, making it much easier for researchers to look at specimens from far-flung institutions, IF the material has been digitized, and particularly if it is available through aggregators such as iDigBio, GBIF or JSTOR Global Plants with their links to massive numbers of specimens.  Still, coverage is uneven, with some collections more fully digitized than others.  Also changed is the way taxonomic information, once generated, is distributed.  Many flora are now published virtually, with or without an accompanying paper format.  The 2012 International Code of Nomenclature for Algae, Fungi, and Plants made it acceptable to publish descriptions of new species digitally as long as they were responsibly published and properly archived.

Plant taxonomy is also changing because of its increasing links with genetics.  Most treatments of species and genera now include DNA sequencing data.  While this has been going on for decades, the last ten years or so have seen greater use of DNA data derived from samples taken from herbarium specimens, with NGS, next-generation sequencing (NSG) making this possible. NGS techniques utilize small pieces of degraded DNA found in dried plant material easier to sequence and to determine how such sequences fit together to provide meaningful results.  That this work has revolutionized taxonomy is hardly news.  Still, it is interesting to look at how the information has solved various puzzles, such as the origin of European potatoes or the origin of the pathogenic Phytophthora strain responsible for the Irish potato famine of the 1840s.  In a study of the genetics of grapes, researchers used over 200-year-old specimens from the herbarium at the Royal Botanical Garden in Madrid.  These plants were collected by Simón de Rojas Clemente y Rubio, considered one of the founders of the botanical study of grape vines, especially varieties used in wine-making.

DNA is not the only chemical being extracted from specimens to glean useful information about plants and also about their ecological relationships.  For example, researchers in Copenhagen tested specimens of four species of Salvia used for medicinal purposes for levels of terpenoids, known to have medicinal applications.  These plants were collected over the past 150 years.  While the terpenoid levels did decrease with the specimen’s age, the “chemical composition of four Salvia species are predominantly defined by species, and there was a substantially smaller effect of year of sampling.  Given these results, herbarium collections may well represent a considerably underused resource for chemical analyses.”  Also being investigated are secondary metabolites that plants produce to control herbivore damage.  In one study researchers were able to extract pyrrolizidine alkaloids from plants in the Apocynaceae family that includes milkweed.  The specimens were as much as 150 years old, and even in those treated with alcohol or mercuric chloride, alkaloids were detectable.

There has also been work on the presence of heavy metal pollutants in collections as a way of tracking contamination.  A study at Brown University in Providence, Rhode Island analyzed samples from specimens collected around the city from 1846 to 1916, compared with newly collected ones.  Levels of copper and zinc remained relatively consistent, but lead levels were much lower in plants growing in Providence today.  It was impossible to test accurately for another toxic heavy metal, mercury, because mercuric chloride was so often used to prevent insect damage to specimens.  While toxic metals in plants might make them seem less palatable as food sources, there is an emerging field of agromining:  growing plants that are hyper-accumulators of metals like lead and mercury to eventually reduce soil contamination.  Herbarium specimens can be used to discover how long areas have been contaminated and also to identify species that are particularly good at extracting metals.  There are even some who think that growing plants in nickel-rich soil could be a way to extract this metal for sale.

Such studies suggest that the possible uses of specimens are only limited by the ingenuity of researchers in coming up with them.  It is fun to see what they can ferret out.   The British botanist Mark Spencer recently published a book on his work as a forensic botanist.  It has a great title:  Murder Most Florid (2019).  He was at the herbarium at the Natural History Museum, London curating the British and European collections when he was first asked by the police to aid in a murder investigation.  Human remains have been found in a forested area and had apparently been there for several years.  Would he be able to determine the time more precisely by studying plants at the site?  I don’t want to spoil this story or the other great ones in the book, but I will say that Spencer explains why a herbarium is essential for the work he does, now that he has become much more involved in forensics.

Reference

Spencer, M. (2019). Murder Most Florid: Inside the Mind of a Forensic Botanist. London, UK: Quadrille.

Oakes Ames at Harvard

Specimen of Malaxis dentata with watercolor by Blanche Ames in the Oakes Ames Orchid Herbarium at Harvard Univeristy

Now that I’ve introduced Oakes and Blanche Ames in the previous posts (1,2), I want to discuss some of Oakes’s contributions to botany at Harvard University.  As a student he was already obsessed by orchids and visited the great herbaria, including Kew.  He also kept adding to his own living collection.  After receiving his bachelor’s and master’s degrees at Harvard, the latter in 1899, he devoted himself full-time to orchid systematics.  Within a few years he became known as an expert on orchids, having hired assistants and set up a laboratory, herbarium, and library in his home.  He published work as coming from the “Ames Botanical Laboratory,” contributing the orchid section to the seventh edition of Gray’s New Manual of Botany (1908) and to the Cyclopedia of American Horticulture (Bailey, 1909).  An indication of his seriousness about systematics is that in 1906 he donated his living collection of orchids to New York Botanical Garden and concentrated on building his herbarium, which eventually grew to over 60,000 specimens.

Ames held a variety of appointments at Harvard.  First, he was assistant director and then director of Harvard’s Botanic Garden.  He gave up the position in 1922 because he was discouraged by the lack of support it received.  He thought that was the end of his connection with the university, but the next year he was hired as curator of the Harvard Botanical Museum and served in various capacities there, including director and later associate director until his death 1950.  He was responsible for reorganizing the famous Ware Collection of Blaschka glass models of plants from an alphabetical to a phylogenetic arrangement.  He also wrote a booklet about them that became a best seller at the museum, with 200,000 copies printed (Ames, 1947).

Ames’s other roles at the university are almost too numerous to mention.  Over the years he rose from instructor to professor of botany.  He became head of the Arnold Arboretum in 1927 after the sudden death of the long-time director, Charles Sprague Sargent.  This involved a diplomatic problem in that the administration feared that the British-born Ernest Wilson, a famous plant collector and Sargent’s assistant, wouldn’t be happy with being passed over for the position.  Ames met with him and proposed to make him Keeper of the arboretum.  British-born Wilson was thrilled with this very British title.  It is no wonder that Ames was given to several administrative jobs including ten years as Chairman of the Division of Biology (Plimpton, 1979).

For many years, Ames taught a graduate course in economic botany into which he poured much time and effort.  In the last post, I mentioned that his artist-wife Blanche produced posters for the class.  These remained hanging in the classroom for many years and are now preserved in the Botanical Libraries at Harvard.  A number of illustrious botanists took the course including Edgar Anderson, a noted plant geneticist who wrote Plants, Man and Life (1952), a book that is still worth reading.  It includes a chapter called “Uneconomic Botany,” about Ames’s course and his rather unique take on the subject that he also described in his book, Economic Annuals and Human Cultures (1939).  Anderson explains that Ames was leery of the anthropological evidence that agriculture had arisen a few thousand years ago.  Ames thought its origins were much older, because it would have taken a great deal more time for plants to evolve from their wild to cultivated forms.

Ames also contended that once humans discovered a useful plant, they usually found it was good for more than one thing:  a food might also have medicinal properties such as seed oil employed as a salve.  Anderson admits that at the time he took the course, while he loved it, he thought it was useless.  However, when he worked in a botanical garden and had to interact with the public, he found Ames’s interesting information very helpful.  Many of the plants Ames discussed produced psychoactive substances:  tobacco, tea, cannabis.  These intrigued another of his students, Richard Schultes, who became a leading expert on hallucinogenic plant products, discovering many of these plants during field trips to South American rainforests where he lived with indigenous peoples and learned from them.

Besides enlivening the intellectual life of Harvard students, Ames wrote over 300 research papers and seven volumes on the Orchidaceae.  Many of these books were published by the Harvard Botanical Press, which Ames set up at his own expense in the Botanical Museum’s basement.  At the end of his career, it published Orchids in Retrospect (1948), a collection of Ames’s essays that Schultes and his colleagues at Harvard edited.  In a forward to the second edition, Schultes wrote proudly that they were able to put the volume together without Ames finding out about it and managed to go through it so thoroughly that there were no printing errors.  A much later volume, Orchids at Christmas (2007), is a tribute to both Oakes and Blanche and includes the orchid etchings that Blanche created and that they sent as Christmas cards between 1937 and 1949.  It is a beautiful little book, with reminiscences by family members as well as photos, including one of the memorial gravestone that Blanche sculpted with some of Oakes’s favorite orchids (see image above).

References

Ames, O. (1939). Economic Annuals and Human Cultures. Cambridge, MA: Botanical Museum of Harvard University.

Ames, O. (1947). The Ware Collection of Blaschka Glass Models of Plants in the Botanical Museum of Harvard University,. Cambridge, Mass.: Botanical Museum of Harvard University.

Ames, O. (1948). Orchids in Retrospect: A Collection of Essays on the Orchidaceae. Cambridge, MA: Botanical Museum of Harvard University.

Ames, O., & Ames, B. (2007). Orchids at Christmas (Reprint edition). Cambridge, MA: Botanical Museum of Harvard University.

Anderson, E. (1952). Plants, Man and Life. Berkeley, CA: University of California Press.

Plimpton, O. (Ed.). (1979). Oakes Ames: Jottings of a Harvard Botanist. Cambridge, MA: Botanical Museum of Harvard University.

Humboldt and Bonpland

2 Bonpland Cinchona

Nature print of Cinchona made by Humboldt and Bonpland, in the Institut de France, Paris.

Since Alexander von Humboldt’s training was in geology and Aimé Bonpland’s in botany, it’s not surprising that Bonpland took the lead in plant collecting on their Latin American expedition (see last post).  However, because they were essentially on their own, picking up assistants along the way, their work in processing specimens, in taking meteorological and astronomical readings, etc., was usually a team effort.  They were overwhelmed by the exciting new plants they saw and within the first few months had already amassed 4000 specimens.  They had to order more paper, since they were using it up so quickly.  It is impossible to say how many plants they collected in total, but the number is over 60,000 including 6,000 species, over half of them new (Lack, 2009).  None of these numbers are precise because many of the plants passed through several different hands, but the record is clearer than for many collections of the era because the two kept careful records that became a model for later expeditions.  They numbered each specimen and recorded it in a journal along with a tentative ID, a description, and locality information.  As time went on, the entries became more detailed.  While they sent back a number of shipments divided among several ships to guard against loss, they kept a small herbarium with them as a memory aid for what they had seen.

Needless to say, none of this work was easy.  Humboldt and Bonpland were traveling through rough, often mountainous terrain in hot and humid equatorial regions where they were driven mad by insects.  These conditions damaged or destroyed many of their specimens, and at one point they were so discouraged by the losses that they made to nature prints to document the plants.  Over 200 of these are preserved at the Institut de France in Paris (Lack, 2001).  However, they persisted in collecting because they just couldn’t ignore all the new species they encountered.  Through much of their trip they were in areas that the eyes of trained botanists had never seen so they were inundated with novelty.  Along with all the environmental data they had amassed, this treasure trove made them anxious to return to Europe and begin writing up their findings.  After leaving South America, they spent a year in Mexico, then returned to Cuba to pack up their specimens for shipment to Paris.  Humboldt decided to live there rather than to return to his native Prussia, because Paris was an intellectually alive city at the time with the National Museum of Natural History (MNHN) as the center of the country’s botanical research.  There was a great herbarium there, as well as a botanical library and experts to assist them.

Humboldt and Bonpland organized and divided up the collection so they each had a set of specimens.  Humboldt arranged for Bonpland to receive a pension from the French government to support him.  Bonpland became botanist to the Empress Joséphine at Malmaison estate, where he oversaw the gardens, provided her with new exotics, and saw to the lavish publications on her plant collection.  It soon became obvious to Humboldt that even with frequent nudging, Bonpland wasn’t getting anywhere with describing their plants.  So in 1813, eight years after they returned, Humboldt invited Carl Kunth, a young German botanist, to come to Paris and work on the collection.  Kunth remained for over six years and eventually published seven volumes with descriptions of over 4,500 plant species, among which 3,600 were new to science (Lack, 2009).  However, this summary makes the process seem more clear cut than it was.

In 1814, Empress Joséphine died, and Bonpland decided to return to South America; he felt more comfortable exploring for new plants.  He took his herbarium with him, and perhaps more importantly, he packed the botanical journals where the specimens were catalogued.  Humboldt and Kunth were aghast, and Kunth hurried to the port of Le Havre to intercept Bonpland before his ship sailed.  Bonpland didn’t give up his specimens, after all Humboldt had a collection too, but he did return the notebooks to Kunth, making it possible for the latter to continue his taxonomic work (Lack, 2004).  Eventually, Bonpland returned his sheets to the herbarium at the MNHN in Paris, where they were filed in the general collection rather than kept separately as the Humboldt collection is.

Another wrinkle was that, while still in South America, Humboldt had sent specimens and seeds to his mentor Carl Willdenow, who wrote descriptions of a number of species.  Some of these were published by others after Willdenow’s death, and his herbarium was sold by his heirs to the Berlin herbarium.  Because of the hostility between France and German, Berlin botanists refused to share specimens with Kunth, who then named some of the same plants, causing years of nomenclatural difficulties.  Kunth returned to Germany after completing most of his work for Humboldt, and when he died his herbarium was also sold to the Berlin-Dahlem Botanical Garden.  During World War II, the Willdenow specimens were considered valuable enough to be stored in a vault offsite and survived the bombing that destroyed most of the herbarium’s collection, including the Kunth specimens.  Lest you assume that by now all of the Humboldt-Bonpland plants had been identified, that may not be the case.  In 2007, a new species, Solanum humboldtianum, was described from a relatively recent collection, but researchers discovered that Humboldt and Bonpland had collected it, and it had lain unidentified for two centuries (Granados-Tochoy et al., 2007).  This is all fodder to feed my love of herbaria.

References

Granados-Tochoy, J. C., Knapp, S., & Orozco, C. I. (2007). Solanum humboldtianum (Solanaceae): An endangered new species from Colombia rediscovered 200 years after its first collection. Systematic Botany, 32(1), 200–207.

Lack, H. W. (2001). The plant self impressions prepared by Humboldt and Bonpland in tropical America. Curtis’s Botanical Magazine, 18(4), 218–229.

Lack, H. W. (2004). The botanical field notes prepared by Humboldt and Bonpland in tropical America. Taxon, 53(2), 501–510.

Lack, H. W. (2009). Alexander von Humboldt and the Botanical Exploration of the Americas. New York, NY: Prestel.

Darwin’s Botanists: Joseph Dalton Hooker

Illustration of Rhododendron glaucum from Joseph Hooker’s The Rhododendrons of Sikkim-Himalaya, Biodiversity Heritage Library.

Joseph Dalton Hooker (1817-1911) was born into the botanical world.  His father was William Jackson Hooker (1785-1865), a botany professor at the University of Glasgow who then became director of the Royal Botanic Gardens, Kew.  Joseph eventually succeeded his father in that post, but his career had more bumps than this succession might suggest.  The Hookers did not have the wealth of the Darwins, so they needed salaried appointments in order to pursue their interest in science, something Charles Darwin never had to consider.  William could provide for his family, but as an adult, Joseph had to find his own means of support after graduating with a medical degree from the University of Glasgow.  Eight years younger than Darwin, Joseph Hooker took a similar route to gain experience in natural history by participating in the British Navy’s Ross Expedition to Antarctica, serving as assistant surgeon; both he and the surgeon were also charged with collecting natural history materials.  Setting out in 1839, they visited South Africa and several groups of islands on their way to and from Antarctica, as well as Australia, New Zealand, and Tierra del Fuego. 

By the time Hooker arrived back in Britain four years later, he had not only amassed a large herbarium but also made many drawings.  Like his father he was an accomplished botanical artist and created many of his own illustrations, especially for his early publications.  On his return to Britain, he began work on studying his collection and publishing descriptions of new species.  Hooker also analyzed some of Darwin’s specimens from the Beagle expedition.  Eventually Hooker described many of them and in the process became quite friendly with Darwin who was thrilled to have his plant collection studied after the long delay in John Henslow’s hands (see last post).  Their friendship flourished and continued until Darwin’s death.

In 1841 William Hooker became director of Royal Botanic Gardens, Kew, but there was no paid position available to his son.  Joseph applied to be professor of botany at Edinburgh, but didn’t get the job, so he worked for the British Geological Survey and learned paleobotany.  In 1847, he went on another expedition, this time to the Himalayas as a plant collector financed by Kew.  He sketched many of the plants, especially the rhododendrons, and it is amazing how beautiful these sketches are considering the rough conditions under which he worked (see above).  When Joseph returned to Britain he went to work on his collections, and finally obtained a paid position at Kew as his father’s assistant in 1855.  Ten years later, when his father died, he became director and was paid to cede the elder Hooker’s herbarium to Kew where he could still have access to it. 

In the meantime, Darwin had been developing his ideas on evolution, having written up a 230 page “summary” in 1844.  He had copies made and gave them, in sealed envelopes, to his wife Emma and to Joseph Hooker so that in case of his death it could be published, though he wasn’t ready to do the deed himself.  At one point after this, Hooker bluntly suggested that while Darwin’s interests were definitely broad, extending from variation in domesticated animals to fossils to plant breeding, he really hadn’t delved deeply into any one group of organisms.  He needed to study some segment of the living world so closely that he would get a sense of the issues involved in distinguishing one species from another.  This was the start of Darwin’s eight-year odyssey studying barnacles that resulted in a two-volume publication on them.  Janet Browne (1995) sees this as Hooker’s most significant contribution to Darwin’s thinking. 

Hooker was also very involved in dealing with the crisis that overwhelmed Darwin when he received a letter from Alfred Russel Wallace in 1858 outlining a theory of natural selection very similar to Darwin’s own.  Hooker and the geologist Charles Lyell calmed Darwin down and devised a plan in which they presented Wallace’s paper, along with a short summary of Darwin’s work, at a meeting of the Linnean Society of London (Browne, 2002).  This was another major event in Hooker’s relationship with Darwin and led to Darwin’s writing On the Origin of Species (1859) within a year.  Though they didn’t always agree on all the finer points of the theory, Hooker remained an important support to Darwin especially because in the years after publication of The Origin, Darwin wrote a number of major works on plants.  Hooker supplied not only advice and taxonomic assistance, but also sent Darwin orchids and other plants from Kew. 

Hooker had an illustrious career in his own right as described in Ray Desmond’s biography (1999).  In Imperial Botany, John Endersby (2008) takes a different tack toward Hooker’s profession and analyzes how he used his position at Kew to command an army of collectors around the world to add to the already outstanding herbarium his father had amassed.  Endersby argues that Hooker was intent on remaining in control of plant taxonomy, particularly of naming new species.  He sternly directed collectors to send the material to Kew rather than attempt to describe species themselves.  Such tight reins were difficult to maintain as collectors became more knowledgeable about the plants where they lived and collected, for example, in Australia and India.  Hooker’s argument was that they lacked the broad collection he had available and so tended to see something as a new species, when it was only a variant (Boulter, 2009).  In other words, colonial botanists were splitters and imperial botanists like Hooker were lumpers.  During his career, Hooker published an impressive array of books including Genera Plantarumwith George Bentham, The Rhododendrons of Sikkim-Himalaya, and Flora of British India, as well as works on plants he collected in Tasmania, New Zealand, and Antarctica. 

References

Boulter, M. (2009). Darwin’s Garden: Down House and the Origin of Species. Berkeley, CA: Counterpoint.

Browne, J. (1995). Charles Darwin: Voyaging. Princeton, NJ: Princeton University Press.

Browne, J. (2002). Charles Darwin: The power of place. Princeton, NJ: Princeton University Press.

Desmond, R. (1999). Sir Joseph Dalton Hooker: Traveller and Plant Collector. Kew, UK: Royal Botanic Gardens, Kew.

Endersby, J. (2008). Imperial Nature: Joseph Hooker and the Practices of Victorian Science. Chicago: University of Chicago Press.

The Linnaean Apostles: Daniel Solander

3 Solandra tridentata LINN 332a.2

Solandra tridentata (LINN 332.2) from the herbarium of the Linnean Society, London

In the last post I discussed one of Carl Linnaeus’s students, Peter Forsskål, who never returned from his expedition to the Near East.  Daniel Solander (1733-1782) traveled farther and also lived to study the fruits of his exploration.  He was born in Lapland and, not surprisingly, studied in Uppsala where he was considered Linnaeus’s favorite pupil.  When British botanists asked Linnaeus to send someone to England to boost the use of Linnaean taxonomy, Solander was chosen.  He left in 1760 and never again lived in Sweden.  At first, he spent time reorganizing the herbaria of wealthy patrons such as the Duchess of Portland and Peter Collinson, who was one of those who had encouraged Linnaeus to send an “apostle” to England.  He was influential in British botanical circles as a member of the Royal Society, a trustee of the newly formed British Museum, and a patron of the American nurseryman John Bartram.  Solander sent specimens from Collinson and others on to Linnaeus.  When the British Museum was looking for someone to care for the herbarium of Hans Sloane, the donor of the museum’s founding collection, Collinson asked Lord Bute, then Prime Minister, to suggest Solander to the King as the ideal choice (Rose, 2018).  This is a fascinating, though tiny, piece of history because all of the individuals involved were interested enough in plants that the care of a plant collection would be discussed at the highest government levels.

Solander began working at the museum in 1763 and set about giving the plants in the herbarium Linnaean names without disrupting the physical order of its 265 bound volumes.  This was a compromise that would allow those not familiar with the new system to still find plants in the collection using Sloane’s system, which was essentially an annotated copy of John Ray’s Historia Plantarum in which Sloane or his botanical curator had written the volume and folio numbers for each species, noting new species when necessary.  Solander began with the first volumes of the herbarium, those containing the collections Sloane had made while he served as physician to the Duke of Albemarle on the island of Jamaica in 1687-1688.  Among the 800 species were hundreds of new ones that Sloane described in his two volume Natural History of Jamaica (1702-1727).  Solander wrote the names on labels that he added to Sloane’s sheets, but retained the older labels, an approach that not been taken by many earlier botanists though later became the norm.

While at the museum, Solander began to receive visits from a young botanical enthusiast, Joseph Banks, heir to a large fortune who had attended but not graduated from the University of Oxford.  He supplied Solander with an unmarked copy of the Sloane Jamaica volumes to annotate.  This was a good way to keep track of the Jamaican species.  When Solander moved on to the rest of the collection, he used slips of paper to record the new names and crossreferenced them with the volume and folio numbers.  He also annotated a copy of Linnaeus’s Species Plantarum.  In this way, the collection was “modernized” without being rearranged.  This went on until 1768, when Solander took on a very different kind of challenge.

Banks had convinced the British Admiralty to make him part of the round-the-world expedition to be led by James Cook on the Endeavor.  It’s major aim, and the publicized one, was to observe the transit of Venus across the sun, which would take place on June 3, 1769.  However, Cook was also instructed to visit Australia, acquiring as much navigational and geographic information for future use in possible colonization (O’Brian, 1993).  Banks, at his own expense, put together a team to study natural history.  It included Solander, Herman Spöring of Finland as secretary, two artists, and two servants.  Banks paid to outfit the ship for his group as well as for scientific instruments and other supplies for preserving specimens and even for growing plants.  There is obviously a lot to this story, but for now I will stick to Solander and plants.

Apparently Banks and Solander made a good team.  They developed a system for dealing with the huge amount of material they collected, in all, about 30,000 plant specimens.  They would return to base each day, give the artist Sydney Parkinson the fresh material to sketch and make color notes, while they, with Spöring’s help, wrote up their notes.  The plants were pressed, though at times, as when they reached Australia, their system couldn’t keep up with collecting.  Plants weren’t drying fast enough, so the pair laid them out on deck on sails during the day.  Needless to say, this massive collection proved daunting to tackle for publication.  Banks and Solander worked on it for years, with engravings made of about 800 species from the Parkinson drawings.  The artist hadn’t survived the voyage, but he did produce 900 complete watercolors and as well as hundreds of sketches.  Unfortunately, Solander died in 1782 before the project was completed, and Banks seems to have given up first-hand work on botany after his death.  Instead, Banks became more involved in projects to promote botanical exploration as well as agriculture.  The Banks’ Florilegium wasn’t published until the 1980s in 34 massive volumes.  Solander did not publish much but he was obviously essential to the Endeavour mission, and perhaps even more importantly, to making the Sloane Herbarium a continuingly useful botanical resource.

References

O’Brian, P. (1993). Joseph Banks: A Life. Boston, MA: Godine.

Rose, E. D. (2018). Specimens, slips and systems: Daniel Solander and the classification of nature at the world’s first public museum, 1753–1768. The British Journal for the History of Science, 5 (2), 1–33.

Linnaeus Beyond the Netherlands

4 Chelsea Garden

Chelsea Botanical Garden, London

This is the last of a series of posts about Carl Linnaeus’s three-year stay in the Netherlands and how it shaped his future career.  While there he had two opportunities to travel to other parts of Europe and meet leading botanists of the day.  It was while living on the estate of George Clifford at Hartekamp and working on cataloging his collection (see last post), that Linnaeus took time off for a month in England to look into what he had heard to be a vibrant botanical community there.  Clifford agreed to this hiatus and even financed it, with the stipulation that Linnaeus return with new plants for his estate.

Not surprisingly, Linnaeus first visited Hans Sloane, then an aged icon among collectors, who opened his herbarium to the Swede.  Jan Frederik Gronovius had already sent Sloane a copy of Linnaeus’s Systema Naturae, and Herman Boerhaave wrote a letter of introduction in which he put Linnaeus on a par with Sloane, describing them as “a pair of men whose equal is hardly to be found in all the world” (quoted in Blunt, 1971, p. 110).  Sloane didn’t quite see things that way and didn’t pay that much attention to Linnaeus who later described Sloane’s herbarium as disorganized.  His first meeting with Philip Miller, the head of the Chelsea Physic Garden, was also less than a success, but eventually Miller gave Linnaeus a good selection of plants to take back to Clifford, as well as herbarium specimens that William Houston had collected in Central America.

In London, Linnaeus met another key member of the botanical confederacy, Peter Collinson, who had already begun a long-term correspondence with John Bartram, the Philadelphia naturalist and nurseryman.  Over a 30-year period, Bartram sent a large array of specimens, seeds, and cuttings to Collinson, who in turn distributed them to a number of the leading gardeners of the day who were anxious to have the latest finds from North America.  Collinson got along well with Linnaeus, and they continued to correspond over the years, with Linnaeus examining some Bartram specimens that thus became types for Linnaean species.  Linnaeus must have met up with Georg Ehret in London, since the artist wrote that he had given him plates to finish Clifford’s catalogue.  In addition, John Martyn, a professor of botany at Cambridge and a London physician, was impressed enough with Linnaeus that their meeting led to a regular correspondence.

Linnaeus also managed time for a trip to Oxford where Johann Jacob Dillenius was professor of botany.  As with several other Linnaean first meetings, this one did not go well because Dillenius had read some of Linnaeus’s early publications, and he felt they threw botany into confusion.  After a few frosty meetings, they finally reconciled when Linnaeus showed Dillenius that he was wrong about his description of the genus BlitumThen Dillenius finally appreciated the depth of Linnaeus’s knowledge, and they had a lively conversation and continued to correspond afterwards.  Obviously Linnaeus’s time in England was very fruitful and provided him with several important contacts who would continue writing to him with information for years to come.

When Linnaeus left Hartekamp in fall of 1937 after finishing the catalogue that would become Hortus Cliffortianus, he went back to Leiden and spent the winter there, working with Adriaan van Royen in the botanic garden, classifying plants according to his sexual system (Rutgers, 2008).  In the spring, he started out for his return to Sweden by going in the opposite direction, to Paris, to visit the famous Jardin des Rois where he met the de Jussieu brothers.  Antoine was older, a professor of botany at the Jardin and a physician; he was a busy man.  He had one meeting with Linnaeus and introduced him to Bernard who then served as his guide.  Bernard de Jussieu showed him the herbarium, and they went through Joseph de Pitton Tournefort’s specimens, a broad collection that included plants from his voyages to the Middle East as well as to the Caribbean area.

Linnaeus also worked in the Jardin’s botanical library, where there were many books of which he had been unaware.  He prepared a ‘wish list’ and later procured a number of these titles.  At the Jardin, he met two of the most accomplished botanical artists of the day, the elderly Claude Aubriet, who had worked with Tournefort, and his pupil Françoise Madeleine Basseporte.  Aubriet showed Linnaeus the large collection of paintings of plants in the Jardin done over the years, so again, as with the time he spent with Georg Ehret, Linnaeus developed a taste of what the best botanical art looked like.  Paris allowed him to deepen still further his knowledge of botany in terms of specimens, living plants, books, and art.  All these were to figure in his future work, and he left for Sweden having made the best possible use of his three years away from home.  Those who read the first post in this series might remember that Linnaeus’s journey had in part been urged upon him by his future father-in-law who agreed to his daughter’s engagement only with the proviso that there be a three-year hiatus in their relationship.  Having fulfilled the agreement, Linnaeus was still an ardent suitor, and when he got back to Sweden, plans for the wedding proceeded.

References

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.

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.

References

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.

References

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.

References

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.