Botany Today: Herbaria

Global map of biodiversity from GBIF occurrce data, showing continued bias in records from the Northern Hemisphere

While the articles I discussed in the last post on important questions facing plant research hardly mentioned herbaria, they are front and center in Charles Davis’s (2023) article: “The Herbarium of the Future.”  It’s an opinion piece in Trends in Ecology and Evolution, so it perhaps takes a broader view of the uses of herbaria than might be found in a systematics journal.  It is also written with a vitality, a lively pace, as if Davis is trying to fit in as much as possible about the promising future of herbaria—and plants—before a reader’s interest might flag.  But this is unlikely since he does a good job of introducing, one after another, aspects of the world of plant collections and how they can be used now and in the future in researching many questions that appeared in the lists of critical issues in the field (see last post).

Davis employs terms that connote change and growth.  His first heading is “A Revolution in Herbarium Use” where he outlines changes in herbaria and in how they are used.  One is what he terms the development of the global metaherbarium:  the growing collection of herbarium specimen data and images available on the internet, in most cases without paywalls.  The Global Biodiversity Information Facility (GBIF) is the largest of these portals, but there are many others including DiSSCo for European herbaria and iDigBio for those in the United States.  The data in these repositories overlap, and yet there really is no “metaherbarium” which harvests information from all other sources.  And there may never be, or at least it will take a long time to get there.

Davis is presenting what the plant science question group calls “horizon scanning,” peering into the future of what might be (Armstrong et al., 2023).  However, there are enormous technical difficulties in linking even collections that are using similar hardware and software.  The plus side is that as these problems have come to light so has the realization that they must be dealt with on a global level (Manzano & Julier, 2021).  The Alliance for Biodiversity Knowledge and other organizations such as the long-standing Biodiversity Information Standards (TDWG) are important forces in moving these goals forward.  The reason for urgency in effectively mobilizing data for all natural history collections is the crying need to use them for research on biodiversity and its conservation, or as Davis puts it: “Innovating Traditional Applications of Herbaria to Speed Discovery.” 

Writing of innovation and speed are rhetorical devices Davis uses to emphasize how critical the situation is.  There are still new species to be discovered, many of these already sitting in herbarium cabinets.  Could AI help to recognize some of them?  Here again, we are still in the early stages, but there have been significant advances in training machine learning systems to identify specimens.  The same is even more true of improvements in “herbariomics,” that is, extracting and sequencing DNA from herbarium specimens, even in cases where they are hundreds of years old.  Davis writes that:  “The metaherbarium soon will become the central resource for such [phylogenomic] investigations spanning populations, communities, and whole continents (p. 4).”  This is definitely on the far horizon.  If collection databases are often difficult to link together, how much more challenging will it be to extract DNA from far-flung collections?  Still, such forward thinking is essential so that possibilities feed into the groundwork now being laid for this bright future.  It includes training individuals worldwide in the skills needed to bring such work to fruition. 

The final section before the conclusion is entitled:  “Breathing New Life into Herbaria: Expanding Users and Novel Applications.”  This doesn’t require as much stretching to see the horizon because much has already happened here.  Ecologists are becoming more aware of herbaria as sources of information on life cycle traits and how they may change over time (Heberling, 2022).  Fifteen years ago phenological studies of the effect of climate change on flowering times were novel; now they have increased to the point of indicating the complexity and variety of species responses, on both small and large geographical scales.  Insect herbivory, fungal relationships, and pollinator interactions can be investigated, often by using more than one kind of natural history collection. 

Herbaria are also important in conservation work, in comparing past plant distributions with those of the present, and in studying how the genetics of populations may have changed over time.  There are really just too many ways herbaria can be used to list them all here or in Davis’s article (Funk, 2003).  However he does give a rather extensive list of uses, including devoting a full-page spread of photos from an exhibit at his home institution, Harvard University.  The Harvard Museum of Natural History opened “In Search of Thoreau’s Flowers: An Exploration of Change and Loss” in June 2022.  I’ve written about it before (see earlier post), but I want to mention it again here in the context of Davis’s article.  All 600 Thoreau herbarium specimens held at Harvard have been digitized.  These images are in the exhibit, presented through the work of several artists.  Davis is highlighting a trend that has become much more common in the 21^st century:  the use of herbarium specimens as inspiration for artists.  The great thing about this exhibit is that it remained up for almost a year, was at a popular museum, and highlighted the work of a well-known figure.  It was a wonderful way to introduce herbaria to a wider audience, while also highlighting the changes in the environment in which Thoreau collected.

References

Armstrong, E. M., Larson, E. R., Harper, H., Webb, C. R., Dohleman, F., Araya, Y., Meade, C., Feng, X., Mukoye, B., Levin, M. J., Lacombe, B., Bakirbas, A., Cardoso, A. A., Fleury, D., Gessler, A., Jaiswal, D., Onkokesung, N., Pathare, V. S., Phartyal, S. S., … Grierson, C. S. (2023). One hundred important questions facing plant science: An international perspective. New Phytologist, 238(2), 470–481. https://doi.org/10.1111/nph.18771

Davis, C. C. (2023). The herbarium of the future. Trends in Ecology & Evolution, 38(5), 412–423. https://doi.org/10.1016/j.tree.2022.11.015

Funk, V. A. (2003). 100 uses for a herbarium. American Society of Plant Taxonomists Newsletter, 17(2), 17–19.

Heberling, J. M. (2022). Herbaria as big data sources of plant traits. International Journal of Plant Sciences, 183(2), 87–118. https://doi.org/10.1086/717623

Manzano, S., & Julier, A. C. M. (2021). How FAIR are plant sciences in the twenty-first century? The pressing need for reproducibility in plant ecology and evolution. Proceedings of the Royal Society B: Biological Sciences, 288(1944), 20202597. https://doi.org/10.1098/rspb.2020.2597

Specimens: Multiples

Jany Renz’s specimens of Serapias orchids, Basel Herbarium at the University of Basel, Switzerland

It is considered good practice in herbaria today to place just one collection on a sheet.  This might include more than one plant, if they are small, but these are the result of one collection event, in one location.  That wasn’t always the case in the past, and even today some curators, conscious of the high cost of herbarium sheets, hate to see a great deal of space go to waste.  If the specimen is small, it might be positioned on a sheet so that there is room for at least one other specimen of the same species collected sometime in the future.  But if the plant is very small the temptation may remain, and result in rather interesting sheets.  At one point on Twitter, there seemed to be a contest to see who could come up with the most crowded sheet, with one entry from the Natural History Museum, London (NHM) a sheet with six specimens, followed by the Royal Botanic Garden, Edinburgh with eight fern specimens.

Then NHM countered with a nine-specimen example, where the barcodes seemed more obvious than most of the plants.  They also upped the ante by noting that one of the specimens had been collected by the great British ecologist Charles Elton.  Later, the Royal Botanic Garden, Kew entered the fray with another nine-specimen sheet, this time of orchids, made more crowded with lengthy accompanying notes.  All these sheets were of flowering plant specimens.  When it comes to mosses, at least in the past, such examples multiply, with several packets attached to a sheet.  Now many curators favor envelopes or packets stored in boxes or drawers, and much work has been done, often in combination with digitization projects, to remove packets from sheets.

Sometimes it is not entire specimens that are pasted on sheets, but multiples of some plant part, often to show variation.  On a visit to the Basel Herbarium, the visual artist Bea Eggli saw pages of orchid flowers and Tweeted that “I always find a piece of my identity in herbaria.”  To me, this is rather cryptic, yet I can relate to it; the order and variation of form are intriguing (see image above).  The flowers from several species of Serapias were preserved by the botanist Jany Renz in the 17^th century to show variation within and among species.  They were displayed by the Basel Herbarium during a 2022 conference to commemorate the 400th anniversary of the publication of Caspar Bauhin’s Flora of Basel.

A somewhat similar approach was taken more than 400 years later by Norman Douglas Simpson (1890-1974) with leaves of Hedera ivy species and cultivars.  This reminds me of work done in the tradition of Johann Wolfgang von Goethe, with variations on a theme or type.  They display just how much variation there can be even within a single specimen, let alone in genera, and also different kinds of variation from size to wide variations in shape; these leaves vary from entire to having from three to five lobes in various forms. 

There is yet another play on the unity and variety theme.  That’s when collectors amass a number of specimens from the same area at the same time.  Most commonly, the aim is to provide duplicates, herbarium coinage, to be sent to other institutions as insurance against a future calamity, or in trade to build species or geographic diversity in a collection.  But in a post from his lab, Mason Heberling, curator of botany at the Carnegie Museum of Natural History, Pittsburg, discussed five specimens of large-flowered trillium (Trillium grandiflorum).  All of them remain in the herbarium because they are vouchers for a study the collectors, Frederick Utech and Mashashi Ohara, published along with Shoichi Kawano in 1984.  Each sheet shows a different stage in the species’ life cycle, from small seedlings to large plants with flowers.  Since this plant is a spring ephemeral, such a wide variety of stages is more likely to be found in one place.  Heberling notes that Utech and Ohara were ecologists, not taxonomists, so they were more interested in life history.  They found that many herbarium collections did not provide specimens that adequately documented different developmental stages.  So their collections were an attempt to collect differently, and their article an argument for why others should do the same so ecologists would find more valuable material in herbaria and thus become more likely to use what could be an excellent research resource. 

Heberling himself does ecological studies and has published a good review of why ecologists have tended to underutilize herbarium collections and what can be done to make them more useful to this community (Heberling, 2022).  Documenting life history is one strategy and obviously not a new one.  Also used in the past was something that botanist and corn expert Edgar Anderson and W.B. Turrell wrote about in 1935:  mass collections.  This involves supplementing specimens with large numbers of a particular plant part.  One example would be collecting regular specimens from two or three maple trees at a site, and then gathering one leaf from each of 30-50 trees.   In other cases, inflorescences or fruits might be saved.  These could allow studies of the frequency of variations, any discontinuities in these variations, and also correlations between variations.  In other words, multiples matter.

References

Anderson, E., & Turrill, W. B. (1935). Biometrical studies on herbarium material. Nature, 136 (3451), 986. https://doi.org/10.1038/136986a0

Heberling, J. M. (2022). Herbaria as big data sources of plant traits. International Journal of Plant Sciences, 183(2), 87–118. https://doi.org/10.1086/717623

Specimens: Sloane’s Collections of Herbaria

Specimen of Anemone thalictroides in the Sloane Herbarium, Natural History Museum, London

In the last post, I discussed Hans Sloane’s herbarium at the Natural History Museum, London (NHM) and work being done on exploring its contents.  In this post, I want to highlight some of the fascinating specimens found by Brad Scott in his doctoral research on the collection.  I’ll begin with medieval philosopher and theologian Thomas Aquinas, not the likeliest person to come to mind in relation to herbaria.  However, Scott found labels on James Cuninghame specimens cut from what Scott has identified as a 1567 edition of Aquinas.  Using scrap paper for labels or for pressing specimens was not uncommon since paper was often scarce particularly for a shipboard surgeon like Cuninghame who traveled twice to China.  The book may have been abandoned by him or someone else, and served as a ready source of scrap paper.  The fact that Scott hunted down the paper’s provenance suggests the thoroughness of his work.  He did something similar with scraps the physician and botanist Leonard Plukenet employed in making packets for seeds he attached to sheets along with the specimens.  In one case, the paper seems to be an advertisement for “Nendick’s P,” with the rest ripped off.  Scott couldn’t find the exact same version, but did discover another praising the benefits of Nendick’s Popular Pills for scurvy.  This speaks to aspects of material culture relating to specimens.  Beyond their scientific value, sheets often hold revelations about the culture of a period well beyond their scientific value.   

Another indication of careful research is a recent lecture Scott gave on George Handisyd, also a ship’s surgeon who was involved in plant collecting, particularly in South America and around the Straits of Magellan.  To flesh out the information on the specimens, Scott also examined ship logs and Handisyd’s correspondence to correlate items with specific dates and locations.  In another case, he examined the correspondence of Charles Preston to learn more about a package of 70 mosses that the Edinburgh botanist had sent to Sloane.  And going even farther afield, Scott went to the French Academy of Sciences in Paris to look at an album of specimens from Aleppo, Syria to compare it to a similar album in the Sloane Herbarium.  It turned out that the two were almost identical not only in binding, but in the specimens included and how they were arranged.  Apparently they were both created by an apothecary in Aleppo, Jean Bigot, and possibly brought to Paris by the traveler and botanist Jean Thévenot and sold to collector like Sloane who later donated one of his two copies to the Academy. 

William Courten, also known as Charleton, created some of the tiniest labels in the Sloane Herbarium.  Besides the normal labels with species information, in some cases each specimen—and there were often several on a page—had a tiny dot of paper with a cipher he used.  In other cases, ciphers were affixed to the sheets.  Scott has been able to decode many of them and found that they often cite from whom Courten received the specimens or the geographic locations where they were collected.  He was apparently secretive in part because he was hounded by creditors, but he may also have found that this shorthand saved space and time. 

Scott is not the only one who has been finding interesting items in the Sloane collection.  The NHM botanist Sandy Knapp posted about pages of blighted leaves she saw along with Mark Carine, the NHM curator responsible for the Sloane collection and a major investigator for the Sloane Lab project (see last post).  This might not seem like an exciting find, but think about it:  an opportunity to study what was obviously a serious infection from centuries ago.  And there are other examples of non-plants that made their way into the volumes.  Scott reported on finding hake egg cases and even a starfish.  I remember reading about a woodpecker scull secreted in one volume (Jarvis, 2014), and there are also a number of cases of insects not infesting sheets after the fact, but purposefully placed.  Mark Carine notes that the insects often seem to be used as decoration and are not labeled.  However, he did post on a page where the insects were labeled and most of the plants weren’t.  In the British Library, one of Sloane’s books on insects has specimens pasted into the relevant pages, suggesting it’s impossible to sort a collection like this into absolute categories. 

The Sloane collection is definitely full of surprises, and I am sure there are more to be unearthed because what one researcher might find uninteresting, might very well tickle the fancy of another.  A page fragment from Thomas Aquinas comes to mind as something that could definitely be valued differently by different scholars.  I’ll close with one specimen I found interesting almost in its nonexistence.  It is a typotype specimen for Anemone thalictroides named by Leonard Plukenet and recognized as a species by Linnaeus.  Brad Scott describes it as “barely existent type specimen” (see photo above). 

Note: Much of the information for this post came from Brad Scott’s Twitter feed @Trichocolea.  I realize that Twitter is now a platform with many issues, to say the least.  However, as a recent Nature article noted (Insall, 2023), scientists have found it a useful way of sharing information and ideas.  I’ve learned so much from Twitter posts like Scott’s and many others, that I am not ready to give up that link.  I remain @flannerm, though I’m also on Mastodon at @flannerm@mastondon.online.

References

Insall, R. (2023). Science Twitter—Navigating change in science communication. Nature Reviews Molecular Cell Biology, 1–2. https://doi.org/10.1038/s41580-023-00581-3

Jarvis, C. E., & Cooper, J. H. (2014). Maidstone’s woodpecker – an unexpected bird specimen in the herbarium of Sir Hans Sloane. Archives of Natural History, 41(2), 230–239. https://doi.org/10.3366/anh.2014.0244

Herbarium Story: Veronica

Veronica, collected in Dec. 1922 by H.L. Darton, [Cultivated] Lawrence, New Zealand. CC BY 4.0. Te Papa

As became clear in the last series of posts (1,2,3,4) on my herbarium “home” at the University of South Carolina, every plant collection is replete with stories.  Discovering them is an exhilarating experience that may play out over a period of time as the story’s elements are pieced together.  The digitization of collections is one way many stories are now being unearthed as was the case described in a blog post from the Museum of New Zealand Te Papa Tongarewa.  The herbarium staff held an informal “botany blitz” for two weeks during which they devoted themselves to tackling some of the unsorted material that’s a staple of most collections.  Among the finds was a folder labeled Veronica hartiana, but digging failed to come up with any information on this species, so it must never have been published.   

The New Zealand species of Veronica used to belong to a separate genus called Hebe, but these plants were found to be monophyletic with Veronica; hebe is still the common name and also the name of over 800 cultivars.  The six specimens in the folder in question were collected by Henry Darton in 1922-1923 and annotated by Donald Petrie.  Darton taught at the local high school in Lawrence, on New Zealand’s South Island.  He and his friend Henry Hart were plant collectors and breeders who had a nursery where they grew many native species.  Donald Petrie was a Scottish botanist who spent nearly 50 years in New Zealand, working a school inspector for the state of Otago that includes Lawrence.  He named a species of Veronica for Darton, and from the evidence in the folder planned to name one for Hart as well. 

Heidi Meudt, who wrote the blog post, is a curator at the herbarium and went on to investigate this story further.  Scientists and historians have much in common.  Both groups want to answer questions, and in a case like this both science and history are involved.  Petrie noted on the specimen that it had a prostrate growth habit and designated it Veronica hartiana sp. Nov.  He added that “It certainly came from the Chatham Islands and was first grown by a solicitor in Timaru to whom it was sent by Mr. Cox.”  Meudt found that Felix Cox, a sheep farmer, lived in the Chatham Islands, over 600 miles east of New Zealand, and sent many specimens to botanists.  Timaru is on the South Island, a few hours north of Lawrence, so it is likely that the solicitor, who probably was a horticultural enthusiast, had contact with Darton. 

Checking further, Meudt discovered a 1941 letter from Erica Baillie, secretary of the New Zealand Alpine Rock Garden Society.  It accompanied a hebe specimen identified as Veronica chathamica that was “absolutely prostrate.”  She asked that it be identified, noting that someone named Baker said that Captain Hooper of the Amokura brought it back from one of the outlying Chatham Islands.  Meudt points out that two decades after Petrie’s notes, the plant was being cultivated by Baillie, who lived in Wellington on the North Island, so it had gotten around.  The fact that it was prostrate suggests what was identified as Veronica chathamica might be the same or similar to what Petrie proposed as Veronica hartiana. 

More digging revealed that from 1907 to 1921, George Hooper was captain of the Amokura, a training vessel for young men who wanted to become sailors.  He was interested in natural history and there are several of his plant specimens in the herbarium.  At the end of her post, Meudt summarizes:  “We still don’t know for sure if Veronica ‘Hartii’ is the same as V. chathamica, but these specimens seem to fit well within the variation seen in the specimens in the V. chathamica box at Te Papa, and they match most of the characters in other botanist’s descriptions of V. chathamica.”  She thinks that perhaps more information about the plant will come out of the Darton Hart Project aimed at recreating some of the gardens at Lawrence. 

This is definitely a New Zealand story from start to finish and suggests how herbarium specimens can provide windows into the way plants move around and become part of human culture, of horticulture.  It also reveals how people in diverse walks of life:  a sheep farmer, a ship’s captain, a lawyer, and a school teacher all contributed to the movement and cultivation of this species.  And Meudt was able to document this with specimens.  It would be difficult to ferret out all the stories lurking in herbarium cabinets, but it’s nice to see ones like this come to light.  Meudt not only took the time to investigate but then cared enough to document her work in this fascinating post.  What I didn’t mention is that she also gives a good description of what cultivars are and how they are named. 

I have to admit that I also learned a lot from digging into this story.  My knowledge of New Zealand geography was almost nil.  Yes, I knew there was a North and a South Island but I didn’t know that the Chatham Islands are a NZ Territory.  I had heard of Otago, but didn’t know it was region of New Zealand or that the country is divided into regions, not states.  As always, specimens have ended up making me a slightly more educated person, not only in terms of botany, but in this case, history, geography, and horticulture.

Botany for Amateurs: Pressing Plants

Dr. Priestley’s specimen of Carex depauperata, Natural History Museum, London (BM000059255)

Flow is a German magazine dedicated to the paper arts.  It had an English edition until the pandemic, and a friend of my sent me a section on plants from Issue 17, the last English-language number.  It was in three parts.  The first included a brief history of herbaria, a description of a Dutch stationery store’s line of herbarium-themed paper products, and of course, instructions on how to press plants between sheets of paper.  Next was a small Pocket Herbarium, a booklet pasted right onto on the magazine’s pages, ready for use in saving specimens.  It was created by Saskia de Valk who has already marketed a larger version.  The third section included three sheets of much heavier paper with reproductions of Maria Merian prints, suitable for framing as they say.  This entire feature, really the entire magazine, was definitely aimed at amateurs and women.  It could easily be dismissed as DIY fluff, but in the first section Luca Ghini is mentioned as an early champion of plant collections, and the herbarium at the Royal Botanic Garden, Kew is highlighted as the world’s largest.  Presenting Merian’s work provides exposure to some of the best botanical illustration.  In other words, these elements might just encourage some to explore plants, and herbaria, more avidly.

I was seduced by herbaria when I saw a couple of seaweed scrapbooks from 19^th-century Rhode Island produced by local women, one the governor’s wife.  Anna Atkins was not a “professional” botanist, but she could be classified as a professional photographer, and her volumes of seaweed cyanotypes were the first published photography books.  Cyanotypes of plant material are still popular today, as is scrapbooking of all kinds.  I myself am not enamored of this medium, but as I discussed in the last post, there is a spectrum of approaches and levels of expertise in any endeavor.  It can be hard to tell at what point a herbarium morphs into a scrapbook or visa versa.  Leopold Grindon, who worked as a cashier for a Manchester textile company, donated 39,000 specimens to the Manchester Herbarium; this is one of its three foundational collections.   What makes it distinctive is that Grindon often attached illustrations, drawings, and entire articles to a specimen sheet, and in many cases, the accessory material was so extensive it needed a second or third sheet.  The texts included botanical journal articles as well as cuttings from magazines and newspapers.  It is an amazing archive, but there are many collectors who less vigorously augmented specimens.  The Harvard botanist Oakes Ames was one, often including drawings by his wife Blanche Ames (Flannery, 2012). 

Moving along the spectrum are those, mostly amateurs, who kept their specimens in books, and added either their own art or printed illustrations to the specimens.  There are many 19^th-century scrapbooks with poems and other musings either printed or by the maker, along with cuttings; the language of flowers was popular at this time and often leaked into collections that also included scientific nomenclature.  In other words, amateurs ignored the borders between science and art, or science and life.  Even when the use of plant material was quite whimsical, as in a scrapbook of literary clippings with small plant cuttings—and feathers—as decoration, the attention to detail belies a great deal of observation.  Another notebook, the Bible Album of the naturalist Eliza Brightwen has only a few cuttings, but many drawings and prints of plants, along with religious art and texts.  Plants were woven deeply into the lived experience of women who documented them in these books.

The Department of Rare Books and Manuscripts at the Yale Center for British Art has a few of these gems which were highlighted in a wonderful book Of Green, Bird, and Flower: Artist’s Books and the Natural World (Fairman, 2014).  One example is an herbarium created by a Miss Rowe apparently as an entry in an herbarium contest conducted by the Liverpool Naturalists’ Field Club in 1861.  Such competitions were relatively common in the 19^th century and were akin to horticultural competitions for the best rose or geranium or flower arrangement.  There is no record of who won this particular contest but this entry should have.  Each carefully labeled specimen was enclosed in a blue envelope with a watercolor of the plant painted on it.  These were arranged in a wooden stationary box.  Miss Rowe was definitely someone who took her botany seriously, and her art as well. 

But lest you think that only women were careful in their presentation of plants, I have to mention a single specimen that I saw on the Twitter feed for the Natural History Museum, London (@NHM_Botany).  It is a Carex depauperata specimen collected by William Overend Priestley.  In the upper left hand corner, outside a blue sheet framed in gold there is a note: “Prepared by Dr. Priestley, and presented by him 1889.”  I don’t know if this sheet is unique, or if Dr. Priestley, whose Wikipedia entry describes him as an obstetrician and makes no mention of botanical interests, made a habit of creating such extravaganzas.  All I know is that this one sheet has everything:  not only the specimen, but illustrations of the flower parts, along with dissected parts (see above).  There are also seeds and even nature prints of seeds at the bottom.  The illustrations are very delicate, done with a fine hand.  And I have to say the gold trim is a nice touch.  This specimen is light on information, though it does give the date and location of collection and the plant’s scientific name.  It’s hard to see this as a serious scientific artifact, but it is, and illustrates just how hard it is to fit botanical work into neat categories.

References

Fairman, E. R. (Ed.). (2014). Of Green Leaf, Bird, and Flower: Artist’s Books and the Natural World. New Haven: Yale University Press.

Flannery, M. C. (2012). Blanche and Oakes Ames: A relationship of art and science. Plant Science Bulletin, 58(2), 60–64.

Herbaria: Specimens Get Around

Cover of Capitulum, Volume 2, Number 1

Years ago, my husband would attend meetings of the Popular Culture Association because he was interested in visual aspects of the popularization of science in 19^th century magazines and books.  I of course tagged along, and soon discovered that the PCA was about much more than comic books and plastic toys.  There are dozens of sections and I found ones that dealt with material culture, the study of things—including two of my loves: natural history specimens and textiles.  Material culture deals with human-made artifacts and would seem to exclude specimens, but that’s hardly the case.  A specimen is an artifact.  A herbarium sheet is more than a plant.  It’s a piece of paper with at least one label, and the plant has been processed by humans to behave well in two dimensions.  The botanical artist Rachel Pedder-Smith wrote about herbaria and material culture in her dissertation that accompanied her spectacular painting of Kew specimens, Herbarium Specimen Painting.

Over time, a specimen accretes greater significance as more humans interact with it.  Deborah Harkness (2007) writes:  “Every time a dried plant specimen changed hands it became infused with new cultural and intellectual currency as its provenance became richer, its associations greater” (p. 31).  Often though not always, the transfer is noted physically on the sheet, making it easier to explain how a particular plant from, for example Germany, ended up in New Zealand, or why plants collected on a Captain James Cook expedition found a home in Philadelphia.  In the last three posts (1,2,3) I’ve discussed attempts to puzzle out the provenance of herbaria, now I want to take this down to the “microhistory” level and look at the travels of individual sheets. 

I got this idea from a lovely article I read in Capitulum, the recently revamped newsletter of The International Compositae Alliance (TICA).  Abigail Moore (2022) writes about a single sheet at Philadelphia’s Academy of Natural Sciences of Drexel University that has two gatherings of what is now Grindelia ciliata(Nutt.) Spreng.  The first was collected in 1819 by Thomas Nuttall, a British botanist who worked for many years in the United States and collected widely in the West.  The label just gives “Arkansas” as the locale, but in the species description, Nuttall (1821) added “on the alluvial banks of the Arkansas, and Great Salt River.”  The second gathering was from Samuel Woodhouse, who was like many collectors in the West, a U.S. Army surgeon/naturalist.  He was on the Sitgreaves Expedition 1849-1851 to map in the same region Nuttall had visited 30 years earlier.  The locale is “Cherokee Nation,” and Moore explains the technicalities of that term at the time, pointing to an area close to where Nuttall had found the first specimen.  She ends by going into the rather complex nomenclatural history of this species, explaining the various names on the labels.  It’s a lovely article and a reminder of how much can be learned from a single sheet.

In a very different example of the rich history that ANS specimens can reveal, Earle Spamer (1998) writes of 26 sheets there that were collected by Johann Reingold Forster and his son Johann Georg, naturalists on Captain James Cook’s second around-the-world voyage.  Spamer describes how much they collected and how broadly the specimens were distributed, in most cases to British or at least European herbaria.  The ANS specimens were given in 1834 by Thomas Nuttall, who apparently got them from Aylmer Bourke Lambert, an avid British botanist and collector (they are marked “Lambert Herbarium”).  Some Lewis and Clark specimens at the ANS were also once part of Lambert’s herbarium but arrived by a very different route—a story for another time (Spamer & McCourt, 2002). 

On the other side of the world, the herbarium of the Museum of New Zealand Te Papa Tongarewa has a briefer history than ANS but none-the-less an interesting collection.  Eighty-four of its specimens traveled from Japan to Russian to Britain before arriving back in the Pacific.  The Russian botanist Carl Maximowich collected in Japan from 1860-1864 and returned to Saint Petersburg with 72 chests of specimens.  The Natural History Museum, London received 1500 from this hoard, and they sent 84 on to New Zealand.  At the time, New Zealand was a young colony, trying to develop its scientific infrastructure and the museum sought assistance from London.

These specimens were part of a much larger European collection of 28,000 specimens bought from NHM by the museum’s director James Hector in 1865.  He thought Australian botanists needed a reference collection to aid in identification of the many non-native plants spreading through the colony, either inadvertently or purposefully brought in by settlers.  As often happens in understaffed herbaria, most of this material lay in storage until the 1950s, and some of it has only recently been examined in detail.  The collection was put together by three British collectors, but within it were materials collected by others, including at least one specimen of an Easter Island plant (see earlier post).  There were also specimens gathered by a German botanist Johannes Flügge (1775-1816) who established a botanical garden in Hamburg.  It was destroyed by Napoleon’s forces in 1813, yet here is a record of what was growing on the other side of the globe when New Zealand wasn’t even an official colony.  That’s the wonder of herbaria and why I study them.  I bore easily, but I am confident that there is an endless supply of examples like this to keep me intrigued.    

References

Harkness, D. E. (2007). The Jewel House: Elizabethan London and the Scientific Revolution. New Haven, CT: Yale University Press.

Moore, A. (2022). Grindelia ciliata (Astereae), Thomas Nuttall, and the exploration of the American West. Capitulum, 1(2). https://doi.org/10.53875/capitulum.01.2.07

Pedder-Smith, R. (2012). Herbarium Specimen Painting. Rachel Pedder-Smith. http://www.rachelpeddersmith.com/Herbarium/Herbarium.html

Spamer, E. (1998). Circumventing Captain Cook. Lewisia, 2, 2–5.

Spamer, E., & McCourt, R. (2002). The Lewis and Clark Herbarium of The Academy of Natural Sciences. Part 1. Notulae Naturae, 475, 1–46.

Herbaria: Sorting Things Out

Specimen of Zollernia glabra from Brazil, Herbarium Wied [140], photo by P.L.R. des Moraes from article (2009)

It’s hardly news that the preponderance of type specimens are in Northern Hemisphere collections (Park et al., 2021).  To increase accessibility for countries in the species-rich Southern Hemisphere, the Andrew W. Mellon Foundation funded the digitization of over two million botanical type specimens that are now available through JSTOR Global Plants.  Yes, for most of us there is a paywall to scale to fully use the site, but support is available for third-world institutions to gain access.  It is an impressive resource and botanists are using it for more than just finding type specimens of interest.  For two Latin-American botanists, Sandra Reinales and Carlos Parra-O. (2022), Global Plants was a major tool in “disentangling” the specimens of José Jerónimo Triana (1828-1890).  He was a Colombian botanist who collected plants from 1851-1857 for the Chorographic Commission set up by the newly organized government of Colombia.  After the survey was completed, he turned over to the commission a full set of the plants he collected along with a catalogue where the specimens were numbered and organized taxonomically.  This became the “Colombian Catalogue.” 

Triana then took his duplicates to Europe and worked at the Paris herbarium.  There he created a new list, renumbering the specimens.  It ended up in the Natural History Museum, London and so is the “London Catalogue.”  I think you can probably figure out where this story is going, but to add one more level of complexity.  In the listing of some species in the second catalogue is another set of numbers:  collection numbers for specimens gathered by Jean Jules Linden with whom Triana had a long collaboration.  These are designated “Linden numbers.”  The article includes photographs of pages from the catalogues; they are hand written neatly, with the information given in columns. 

The problem is how to relate these catalogues to the many collections containing Triana material.  Obviously the catalogue numbers and specimens sync for those that remain in Colombia.  However, there were multiple duplicates for many of his gatherings located in the NHM, Paris, Geneva, Vienna, and several other European and North American herbaria.  To attempt to figure out the location of type specimens, the authors searched for Triana specimens in JSTOR Global Plants and found over 5000 records.  They then searched in other databases for additional types and cleaned the data by reading the label information and removing those specimens that didn’t fit their criteria.  Obviously this was a lengthy and tedious process, and they were rewarded with some knotty problems to solve.  I can’t even scratch the surface of their detailed work, but I’ll give a brief summary of a couple of issues.  There were cases where the same Triana gathering was used to describe different species; the different numbers on the labels of duplicates was one of the issues.  There were also cases where Triana and other botanists collected in the same area at the same time. One of the specimens designated as a type for Meriania umbellata, a species collected and described by Karl Wilhelm Karsten, also has a Triana label and collection number on it.  To alleviate some issues, Reinales and Parro-O. present guidelines for lectotypification of some names of specimens that Triana described based on his specimens.

Now I soldier on to another herbarium, no less problematic (Moraes, 2009).  Again, it involves South American plants, this time collected by Prince Maximillian of Wied when he was in Brazil from 1815 to 1817.  He explored along the southeastern coast, a species-rich rainforest area.  In 1998  historians were searching family records in what had been his palace and rediscovered his private herbarium.  It had been missing for 20 years and was found when an intrepid researcher decided to investigate a difficult to get at cabinet.  In it were 22 parcels of plants collected over 26 years, so they obviously contained more than the Brazilian material.  In all there were 7000 plants including some from his trip to North America and his European collections, and there were 125 Brazilian plants.  Though this is modest compared to the 5000 specimens of 1000 species that he gathered in Brazil, it does contribute to knowledge of Wied’s work because there are still many of his specimens that haven’t been located.  As with a number of German collections, some might have been destroyed in the large-scale damage to the Berlin-Dalhem herbarium during WWII. 

To bring up the major issue with the Triana specimens of collection numbers, the situation is not as confusing here, though hardly ideal.  Wied didn’t used collection numbers, but he did number some specimens later as he studied them, and some were also numbered by others in the course of their work.  Of the 125 Brazilian specimens in his personal collection, there are 98 species represented, several of which are not found in other Wied material.  Unfortunately, he rarely gave location information on his labels.  Still, Moraes notes:  “Species kept in the private collection of Brazilian plants gathered by Wied represent a precious register of the flora of the Atlantic rainforest of the 19th century.  Its historical value is indisputable since Wied’s vouchers are among the first ones collected in Brazil that are still extant”(p. 46).  In other words, the contents of that cabinet were a pleasant botanical surprise.

References

Moraes, P. L. R. (2009). The Brazilian herbarium of Maximilian, Prince of Wied. Neodiversity, 4(2), 16–51. https://doi.org/10.13102/neod.42.1

Park, D. S., Feng, X., Akiyama, S., Ardiyani, M., Avendaño, N., Barina, Z., Bärtschi, B., Belgrano, M., Betancur, J., Bijmoer, R., Bogaerts, A., Cano, A., Danihelka, J., Garg, A., Giblin, D. E., Gogoi, R., Guggisberg, A., Hyvärinen, M., James, S. A., … Davis, C. C. (2021). The colonial legacy of herbaria. bioRxiv (p. 2021.10.27.466174). https://doi.org/10.1101/2021.10.27.466174

Reinales, S., & Parra-O., C. (2022). Disentangling the historical collection of José Jerónimo Triana from the República de la Nueva Granada between 1851 and 1857. Taxon, 71(2), 420–439. https://doi.org/10.1002/tax.12653

Herbaria: Two Views of the Zierikzee Herbarium

Specimen of common soapwort (Saponaria officinalis), Zierikzee Herbarium

As discussed in the last post, older herbaria are being given increased attention because they shed light on what plants were in growing in Europe and in European gardens at the time, and how botanists approached their work.  But as with any old documents, it is often difficult to unlock their secrets.  A case in point is a beautiful herbarium at the natural history museum in Zierikzee in the Netherlands with no information on who the collector might have been.  In 2021, two articles were published on its contents and provenance, each presenting different conclusions as to who created it and when.  The article by a team from the Naturalis Biodiversity Center in Leiden came out first, and I read it first (Offerhaus et al., 2021).  Like the herbaria discussed in the last post, also studied by a Naturalis team, efforts were made to identify all the species and provide up-to-date names.  The organization of the sheets, the information on the labels, the type of paper used, and many other possible clues were presented. 

In the case of the Zierikzee Herbarium, there was an analysis of the ornate printed labels and vases found on all but 21 of 348 plants.  Their use seems to have been a primarily Dutch fad in the first half of the 18^th century.  They definitely perk up a specimen, giving it a bit of class or status, showing the taste of the collector.  Carl Linnaeus’s patron in the Netherlands, the wealthy merchant and horticulturalist George Clifford used them; they befitted someone with a large garden, and a greenhouse each for Asian, American, European, and African plants.  Historians of botany have studied them and in some cases identified the printers and the years when they were produced.  In the case of the Zierikzee, the Naturalis group focused on labels with a putto on each side of a frame with the species information using pre-Linnaean nomenclature written, in the blank space between them. 

Both articles also researched old auction catalogues to attempt to find the answer to who created this work.  There is not direct evidence in the herbarium itself, which is the major mystery at the center of this controversy.  In the second article, Gerard Thijsse (2021) found a resemblance between an herbarium auctioned in 1790, that of Martin Wilhelm Schwencke.  It consisted of 10 volumes, and Thijsse thinks that six of them make up the herbarium in Zierikzee and that Schwencke collected the plants early in his long career.  Thijsse gives a detailed explanation as to why this makes more sense than the Naturalis team’s hypothesis, which links the herbarium to Jacob Ligtvoet  described in a 1752 auction catalogue shortly after his death.  Reading these two reports reminded me of opposing lawyers carefully laying out their arguments, using a variety of different kinds of evidence, some of it scientific some of it textual.

Both articles also discussed evidence from computer analyses of similarities between the information given in published botanical sources and the label data, which included references to several authors.  The two studies significantly diverge.  The Naturalis group sees some resemblances between the label data and the information in Herman Boerhaave’s two catalogues of the plants in the Leiden botanical garden, one published in 1720 and the other in 1730.  They suggest that the herbarium began to be put together between these two dates and consider the likely creator to be the gardener at Leiden, Jacob Ligtvoet who helped Boerhaave with the second catalogue.  They note that along with the preponderance of the plants from the Netherlands but there are exotic species that Boerhaave obtained from those associated with the Dutch East India Company. 

On the other hand, Thijsse found that the herbarium contains a “virtually” complete set of the plants mentioned by the unknown author of the 1738 Pharmacopoea Hagana.  He also argues that one of the vases used is similar to one the Georg Ehret designed in 1734, so that the Naturalis date for the herbarium is too early.  Since Thijsse’s article appeared later than the Naturalis one, he had the luxury of questioning some of its findings.  I wouldn’t be surprised if Naturalis is now working on a rebuttal.  Of course, there is much fertile ground for argument here.  Many pharmacopoeia of the time included similar species, and the plants could have been mounted earlier and then the vases added later as a decorative afterthought.  That’s the problem with history, it is impossible to know everything about the past, information is always limited and open to interpretation. 

This is very reminiscent of scientific controversies, where researchers ask similar questions but use different protocols or methods of attack and come up with opposite conclusions.  Is that because one of them is wrong or because they are really not looking at the same phenomenon?  Only more research will provide the answer, which may very well be the case here, or there will be a stalemate.  I’ve just finished rereading these articles, and my head hurts.  For once I have reached my herbarium saturation point.  I think there is more here than I want to know about this one collection.  One the other hand, I am not sorry that I spent time on this because it helped me to see how historians work.  Like scientists, they have to be creative in how they approach their problem, asking different kinds of questions so that eventually they may perhaps find a niche that opens up a new world of answers.

References

Post 2

Offerhaus, A., de Haas, E., Porck, H., Kardinaal, A., Ek, R., Pokorni, O., & van Andel, T. (2020). The Zierikzee Herbarium: Contents and origins of an enigmatic 18th century herbarium. Blumea, 66, 1–52. https://doi.org/10.3767/blumea.2021.66.01.01

Thijsse, G. (2021). The four W’s of two 18th century Dutch herbaria: The “Zierikzee Herbarium” and the herbarium of Simon D’Oignies. Blumea, 66, 263–274. https://doi.org/10.3767/blumea.2021.66.03.09

Herbaria: Pre-Linnaean

Specimen of Helichrysum arenarium from Breyne’s 1673 herbarium, Naturalis Biodiversity Center (Jong et al., 2022)

As I‘ve previously mentioned, older herbaria particularly pre-Linnaean ones are receiving increased attention.  In the past, the fact that they were of little use in investigating nomenclatural issues and often had scant label data meant they weren’t of much interest.  But historians are now finding them rich sources for studies on botanical inquiry in earlier centuries.  In addition, these collections are gold mines of information about the ecology of the areas where the plants were gathered and in some cases about agricultural and horticultural practices.  Historical research is often about solving mysteries of the past, and herbaria are full of clues.  This set of posts will deal with a few interesting collections.  The first is pre-Linnaean, that of the Prussian merchant and collector Jacob Breyne (1637-1697). 

In a previous series of posts on early modern herbaria (1,2,3,4) I discussed several in the Naturalis Biodiversity Center in Leiden, the Netherlands.  This institution not only has an outstanding collection but has supported a great deal of research on pre-Linnaean herbaria under the direction of Tinde van Andel, who holds appointments at both Leiden and Wageningen Universities.  She and her colleagues examined Jacob Breyne’s two book herbaria in Naturalis and set them in the context of what has been uncovered about his life (Jong et al., 2022).  With every personal herbarium there is an interplay between the plants and the maker in terms of where that person lived, where they might have traveled, and who else they knew in the botanical world.  In this case, Breyne was from a Dutch family that had moved to what is now Gdansk, Poland where they were traders in raw materials for medicines and dyes.  He learned botany in school and collected around Gdansk.  He then spent time in the Netherlands while he trained as a merchant with this uncle.  He also visited botanical gardens and studied botany at Leiden.  After returning to Gdansk he continued his interest in plants and corresponded with Dutch botanists and collectors connected to the Dutch East India Company, giving him access to plant material from South Africa and Asia.

Breyne’s two bound herbaria, one dated 1659 and the other 1673, indicate he had a sustained interest in plants.  In the two are a total of 105 specimens, some with original labels.  However there’s evidence of many missing specimens, often with the labels missing as well:  60 from the first, which only has 48 remaining, and 10 from the second, from which 10 pages were also cut out.  Most of the plants in these volumes are European species, many from the Gdansk area.  They give a picture of what was growing there at the time and how the flora has changed.  Breyne noted that the frog orchid Dactylorhiza viridis was abundant, though it’s now designated as regionally extinct for Gdansk and its vicinity.  Though there are few exotic specimens included, most plants in these collections are from this area.  Breyne may have collected them on his travels or gotten them from others; the labels contain little information on provenance.

There is another collection of Breyne specimens at Naturalis as part of the large Van Royen Herbarium, which was compiled by Adriaan van Royen (1704-1779) director of the Leiden botanic garden and his nephew David (1727-1799).  Van Andel and her colleagues examined the 89 specimens designated as from Breyne, but found that only 59 had labels with his handwriting, and some of the remainder had nothing indicating a connection to him (Jong et al., 2021).  They studied the 59 with the aim of finding specimens that might have been removed at some point from the two bound herbaria.  However, it became clear that there was probably no such link.  Most of unbound plants were from southern not northern Europe and specifically from the area around Montpelier in southern France, site of a university with a long tradition of botanical inquiry.  There are a couple of possibilities of how Breyne acquired this material since he was not known to have traveled that far.  Historians sift through a collector’s life experience to attempt to find links to collection specimens.  A contemporary of his, Ernst Gottfried Heyse, studied at Leiden and Montpelier before returning to Gdansk to teach and direct the botanical garden there.  Also Breyne corresponded and traded specimens with the French botanist Joseph Pitton de Tournefort.

There are limits to how many questions old collections can answer.  There is no good explanation as to why or when so much of Breyne’s older herbaria was removed.  Also, there’s no convincing hypotheses to explain how some of his specimens ended up in the van Royen Collection.  Breyne died several years before the elder van Royen was even born, but specimens do have a tendency to be passed from botanist to botanist.  The specimens in question appear to have been cut from larger sheets, perhaps from a bound volume.  After his father’s death, Breyne’s son was known to have sent parts of his collection to other botanists, and Leiden was a likely place to send them because of the family’s Dutch heritage.  Perhaps more information will turn up in the future, but even now a great deal has been learned recently about Breyne and his collections.  In the next post in this series, even deeper mysteries surround another old herbarium, and the results of the latest investigations are still open to debate.  

Reference

de Jong, M., Duistermaat, L., Stefanaki, A., & Andel, T. van. (2022). The book herbaria of Jacob Breyne (1637-1697) in the collection of Naturalis Biodiversity Center (Leiden, the Netherlands). Blumea, 67, 77-96.https://doi.org/10.21203/rs.3.rs-885416/v1

de Jong, M., Stefanaki, A., & van Andel, T. (2022). Mediterranean specimens of the Prussian Botanist Jacob Breyne (1637–1697) in the Van Royen Herbarium, Leiden, The Netherlands. Botany Letters, 169(2), 294-301. https://doi.org/10.1080/23818107.2022.2038667

Collections in the Future

Methods for measuring leaf shape in butterfly weed, Asclepias tuberosa, (Woodson 1947, p. 368)

This month’s series of posts will diverge from usual where I keep to one theme.  This time, I’ll seesaw between two topics that I, at least, see as related.  Each stems from a recent publication I found particularly noteworthy on how plant collections will be used and curated in the future.  The first is an article by Mason Heberling (2022) on “Herbaria as Big Data Sources of Plant Traits” and appeared in the International Journal of Plant Sciences.  It is a review of a topic that hasn’t received enough attention:  how herbaria can be used in functional trait analysis.  The other is Issue 8 of The Ethnobotany Assembly or T.E.A., a quarterly online journal about plant-people relationships.  It’s entitled Plant Humanities: Where Arts, Humanities, and Plants Meet and is edited and with a contribution by Felix Driver and Caroline Cornish.  There is a great deal coming out about the plant humanities and some of it I find disappointing, but not this publication.  The articles are varied and thought-provoking, but I’ll save further comments until the next post. 

Why am I juxtaposing such different types of work?  Precisely because they are so different.  Yet they both speak volumes about the possible future of herbaria if researchers of this caliber continue to give their attention to the amazing resources that plant specimens provide to so many fields.  I begin with plant traits because this topic fits squarely within biological inquiry, where recent reports like those on the Extended Specimen Network clearly put herbaria.  However, Heberling argues that specimens have been neglected by plant trait researchers who tend to look elsewhere for data.  He lays out his case in the first part of the article and also provides examples of where herbaria have made significant contributions to the field.  In the second part, which I’ll discuss later, he outlines how in the future herbaria and collecting might adapt to support this area of research. 

After his introduction, Heberling discusses community ecologists’ growing use of plant functional traits in their research during the past 20 years.  Functional traits include morphological, chemical, phenological, and physiological attributes that serve as surrogates in understanding individual fitness.  He uses as an example work on the leaf economics spectrum (LES) where characteristics such as leaf mass per area, construction costs, photosynthetic rate, and leaf life span have been found to relate to each other.  This quartet varies along a spectrum from long-lived, high construction costs, low photosynthetic rates, and large area to the other end with opposite traits.  Patterns falling outside the spectrum are thought to be maladaptive. 

Heberling notes that in most of this work “little or no explicit attention has been paid toward specimens as primary sources of trait data (p. 90).”  One reason he gives is that functional trait analyses are a recent development in plant science and preparing specimens is a technology that has been around for a long time.  In the past, noting traits like phenological status was not necessarily considered important, especially because it would be apparent to someone looking at the specimen.  This made sense until the age of digital data when researchers can be searching online databases for label information on phenological status and not finding it.  An image may not be available, and even if it were, it would be much more time efficient to simply search the data files.  Other changes in processing specimens that could aid trait research include preserving plants in different stages of development for life history research.

Also tackled are the arguments ecologists and evolutionary biologists have against using specimens, including issues of collection bias, such as toward ignoring young and immature plants and choosing those with flower and fruit.  This makes sense for taxonomic studies but not for plant life history work.  However, he contends that “we cannot assume the limits of herbaria without trying (p. 101).”  In this case, awareness of the issue could lead to changes in collection practices, with a broader selection of material chosen.  As for the idea that leaf area changes markedly over time in dried specimens, the assumption has been disproven in comparative studies.  Several traits, including amino acid and metal contents have also been validated for herbarium-based measurement.

Heberling provides an extensive table citing findings where specimens have been studied in trait research and describes many ingenious approaches used both in early studies and also more recently.  One of my favorites is a 2002 paper by Teece et al. on 11 Lewis and Clark specimens, among the earliest collected in the Western United States before development and industrialization caused substantive environmental changes.   Leaf fatty acid content was measured as was stable carbon isotope composition.  These results served as an important baseline for comparison to later specimens.  Heberling also discusses the vast literature on stomata, leaf area, herbivory, and other traits that is based on specimens.  It is a fascinating review.  However, he then notes that there is little herbarium specimen data in the two major trait databases TRY, begun in 2007 and BIEN, started in 2016.  With about 12 million records, TRY has only 10.4% of North American woody plants represented by even one specific leaf area measurement.  It is this dearth that Heberling addresses in the latter part of his article and that I’ll look at in the third post of this series. 

References

Heberling, J. M. (2022). Herbaria as Big Data Sources of Plant Traits. International Journal of Plant Sciences, 183(2), 87–118. https://doi.org/10.1086/717623.

Teece, M. A., Fogel, M., Tuross, N., McCourt, R. M., & Spamer, E. (2002). The Lewis and Clark Herbarium of The Academy of Natural Sciences. Part 3. Modern environmental applications of a historic nineteenth century botanical collection. Notulae Naturae, 477, 1–20.

Woodson, R. E. (1947). Some Dynamics of Leaf Variation in Asclepias tuberosa. Annals of the Missouri Botanical Garden, 34(4), 353. https://doi.org/10.2307/2394774.