Botany Today: Specimen Photographs

In this last of a series of posts (1,2,3) on the future of botany, I want to take up a topic that seems to be receiving more attention lately:  the relationships between specimens and photographs.  Even as stout a defender of herbarium specimens as myself has to admit that there is a considerable difference between them and living plants, between brown and virtually two-dimensional as opposed to colorful and in 3-D.  Photographs have the storage advantages of specimens while providing at least some of the information that specimens lack.  In addition, digital photography has made it cheaper to take photographs and easier to store them. 

Yet despite their usefulness, plant photographs, at least for some species, are not easy to come by.  In a 2021 article in Nature Plants, Pitman et al. reported on a survey of 25 online databases of field photographs and found that they held only about 53% of the 125,000 vascular plant species from the Americas.  The databases they searched included those hosted by botanic gardens, natural history museums, and other entities such as Flora of the World and  Also, three social media and community science platforms were surveyed: iNaturalist, Pl@ntNet and Flickr.  These three accounted for 37.8% of the species, and photos of about a third of these species were not found on any other sites.  Flickr and iNaturalist also had the best geographic coverage. 

Not surprisingly, North American species were more likely to be photographed than those of central and South America.  But there were wide variations in coverage, with 71.9 % of Bolivian species pictured versus 56.8% of those in Brazil.  There were also differences among plant families with 98.1% of the Liliaceae captured, but only 18.7% of the Piperaceae, which have less showy flowers.  Not surprisingly, rare plants were less likely to be photographed as were relatively new species.  In addition, the researchers found that when they rechecked their results after several months that a number of species were represented that hadn’t been there earlier.  So this is definitely a changing landscape, but one that’s troubling to those attempting to document and study biodiversity. 

The Americas are not the only areas that have this problem.  A new Australian study of (Mesaglio et al., 2023) revealed that 17.6% of the Australian flora of 21,077 vascular plant species are not represented by any photographs.  Here 33 online resources were used, again including social media and community science platforms.  Obviously the coverage is much better than for the Americas.  Australia has a well-organized system of state herbaria, was an early adopter of digitization, and is able to draw upon an up-to-date list of its species.  Still there are gaps, again related to the rarity of species or their recent discovery, and there are also geographic disparities.  While Western Australia is the most species-diverse state there are some areas that are very difficult to access, hence less photography has been done.  Other reasons for gaps cited by the authors were for plants that are difficult to identify or that lack “charisma.” 

Both articles stress the importance not only of the number of photographs but their quality.  These need to be high-resolution and capture as many of the plant’s identifying characteristics as possible.  Thus the best photographs are likely to be those taken by individuals who know enough about plant systematics to literally focus on identifying traits.  This is a topic that Carlos Gómez-Bellver (2019) and his colleagues in Barcelona address in an article where they propose standards for photographs to complement herbarium vouchers.  However, the guidelines would also be useful for photographs not physically attached to specimens, but linked to them as part of the extended specimen concept. 

The authors list six cases where photos would considerably increase a specimen’s taxonomic value.  First are large species such as a palm, where photos of the plant as a whole and its habitat would help to put the specimen into context.  Succulent or spiny plants, that are difficult to prepare would also benefit from such photos, with only small pieces of the plant itself attached; the same is true of plants with toxic substances.  Then there are species for which little remains but seeds and withered material, also species that lose important morphological traits when they are pressed and dried.  There might be cases where the specimen is the only one in the vicinity, so the photo would serve as a voucher; the same would be true of plants at sacred sites where collection might be considered disrespectful. 

The authors call for the development of a protocol with standards that photographs must meet to be considered as reference material to be tied to a voucher.  I can’t go into all the elements here, but they include reflecting the entire size of the plant and its habitat.  The photo must also include the standard metadata about date and location, the collector, and precise geolocation.  What they term “fusion vouchers” are those that include both a specimen and photos; these would be stored in herbaria.  Photo vouchers alone could be filed in a variety of ways which are discussed.  At this time, there is no standardization.  But as the other articles I’ve discussed here make clear, it would be important for the photos to be accessible digitally.


Gómez‐Bellver, C., Ibáñez, N., López‐Pujol, J., Nualart, N., & Susanna, A. (2019). How photographs can be a complement of herbarium vouchers: A proposal of standardization. TAXON, 68(6), 1321–1326.

Mesaglio, T., Sauquet, H., Coleman, D., Wenk, E., & Cornwell, W. K. (2023). Photographs as an essential biodiversity resource: Drivers of gaps in the vascular plant photographic record. New Phytologist, 238(4), 1685–1694.

Pitman, N. C. A., Suwa, T., Ulloa Ulloa, C., Miller, J., Solomon, J., Philipp, J., Vriesendorp, C. F., Derby Lewis, A., Perk, S., Bonnet, P., Joly, A., Tobler, M. W., Best, J. H., Janovec, J. P., Nixon, K. C., Thiers, B. M., Tulig, M., Gilbert, E. E., Campostrini Forzza, R., … Hilo de Souza, E. (2021). Identifying gaps in the photographic record of the vascular plant flora of the Americas. Nature Plants, 7(8), 1010–1014.

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 21st 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.


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.

Davis, C. C. (2023). The herbarium of the future. Trends in Ecology & Evolution, 38(5), 412–423.

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.

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.

Taxon and Nomenclature

Turland et al., 2018, published by Koeltz Botanical Books

In my last post in this series (1,2,3) on articles in Taxon, the journal of the International Association for Plant Taxonomy, I want to discuss a group of “perspective” articles on a thorny nomenclatural issue.  The first, by Gideon Smith and Estrela Figueiredo (2022) of Nelson Mandela University in South Africa, dealt with the scientific plant names that refer to people or ideas that can be considered offensive, particularly in a post-colonial context.  The example they use is the root “rhodes-“ to commemorate Cecil Rhodes who made a fortune from diamond mining in South Africa and was prime minister of the Cape Colony from 1890-1896.  He was a symbol of British imperialism and exploitation of indigenous people.  The authors used this example because it was related to a South African movement begun in 2015 called “Rhodes Must Fall,” referring to a statue on the University of Cape Town campus that was eventually removed.  However, the movement developed beyond that and came to embody disposing of lingering reminders of colonialism in other contexts, including botany.  They also mentioned that the Portuguese dictator Antonio Salazar is commemorated in the name, Kalanchoe salazarii, native to the former Portuguese colony of Angola.  In addition they cited an earlier article (Knapp et al., 2020) that brought up the problematic word, caffra, derived from the Arabic for infidel, that is considered an awful racial slur in Africa yet appears in various forms in many botanical epithets. 

Smith and Figueiredo note that the present International Code of Nomenclature for algae, fungi, and plants (Turland and Wiersema, 2018) does not allow for a name change for reasons of offensive language.  They call for a change in the code to address this problem, and in fact, a proposal for such a change was published in the December 2021 issue of Taxon.  (This article was published online after the Rhodes article, but there were difficulties publishing paper editions during COVID, so not all the paper issues were published in sequence).  It was written by two Australia systematists, Timothy Hammer and Kevin Thiele.  They cited the examples of caffra and of hibbertia, commemorating George Hibbert a British slave trader and owner who was also a strident anti-abolitionist.  They proposed that the language in the code stating that a name or epithet cannot be rejected “merely because it is inappropriate or disagreeable, or because another is preferable” be changed.  Also, a new article should be added stating that a legitimate name or its epithet may be rejected as culturally offensive or inappropriate.  The actual proposal gives more detail but that is its essence. 

In the April 2022 issue of Taxon, Sergei Mosyakin, director of the Kholodny Institute of Botany in Ukraine wrote a rebuttal to Smith and Figueiredo.  He notes that his country has suffered from colonialism and ethnic oppression, but argues that dealing with such history through nomenclatural change is fraught with difficulties.  Allowing changes could lead to a “slippery slope” and looks like “a new form of politically motivated scientific totalitarianism and censorship” (p. 251).  Mosyakin goes on at some length about the difficulties in evaluating what is inappropriate and how this is to be decided.  He makes valid points but overall his language is more strident than that in the other articles, and needless to say it provoked a response.

The next article in what was becoming a series was published in the December 2022 issue of Taxon and was written by Smith, Figueiredo, Hammer and Thiele.  It was brief, measured, and to the point, though they do write that Mosyakin “severely misrepresented” their views and proposals.  They address three of his contentions, the first being the slippery slope argument.  The Hammer and Thiele proposal for amending the Code included the creation of a permanent committee, like several others within the Nomenclatural Section, to deal with proposed changes in an orderly fashion.  This would be in keeping with the standard way changes are handled.  The authors admit that there will be some proposed changes that might be considered in a “gray area” between extremes, but contend that this is true of most complex issues and shouldn’t be an argument against dealing with them at all.

The authors also reject the idea that their proposal involves politically motivated censorship.  They see as “far-fetched” the view that the Nomenclatural Section will be conducting “purges” or become a totalitarian regime:  “In our view, if a community of end-users formally decides that a mechanism should be established to restrict the use of some scientific names and epithets for the greater good, this process is not ‘censorship’” (p. 934).  Finally, they don’t think their proposals “erase history” because of how nomenclatural change works.  The previous names do not disappear, but rather, move into synonymy.  This move doesn’t expunge the name but acknowledges that it is no longer considered culturally acceptable.  I think this is the strongest of their arguments.  Whether the change will lead to a slippery slope and what some would consider censorship will only be determined if the Code is amended and proposals for name rejections considered.  In the meantime, this discussion is a fruitful one; it fits well into the much larger conversation about efforts to move toward decolonial natural history collections and beyond that to decolonial societies. 

Since I wrote the first draft of this post, I’ve come upon two more articles on this subject in the December 2022 issue of Taxon, one by Mosyakin and one by Thiele et al.  Not surprisingly they don’t change their positions but do elaborate on them, especially Mosyakin.  He gives several examples of where name change could lead and what new problems it could produce in the future.


Knapp, S., Vorontsova, M. S., & Turland, N. J. (2020). Indigenous Species Names in Algae, Fungi and Plants: A Comment on Gillman & Wright (2020). TAXON, 69(6), 1409–1410.

Smith, G. F., & Figueiredo, E. (2022). “Rhodes-” must fall: Some of the consequences of colonialism for botany and plant nomenclature. TAXON, 71(1), 1–5.

Hammer, T. A., & Thiele, K. R. (2021). (119–122) Proposals to amend Articles 51 and 56 and Division III, to allow the rejection of culturally offensive and inappropriate names. TAXON, 70(6), 1392–1394.

Mosyakin, S. L. (2022). If “Rhodes-” must fall, who shall fall next? TAXON, 71(2), 249–255.

Smith, G. F., Figueiredo, E., Hammer, T. A., & Thiele, K. R. (2022). Dealing with inappropriate honorifics in a structured and defensible way is possible. TAXON, 71(5), 933–935.

Mosyakin, S. L. (2022). Defending Art. 51 of the Code: Comments on Smith & al. (2022). TAXON, 71(6), 1141–1150.

Thiele, K. R., Smith, G. F., Figueiredo, E., & Hammer, T. A. (2022). Taxonomists have an opportunity to rid botanical nomenclature of inappropriate honorifics in a structured and defensible way. TAXON, 71(6), 1151–1154.

*The references are given in the order in which they were first published that is, online.

Taxon and the Flora of Madeira

Map of Madeira, 1904, Edward Stanford; David Rumsey Historical Map Collection

This series of posts deals with the articles found in the systematic botany journal Taxon that deal with topics beyond conventional taxonomic treatments.  Among my favorites are those with an historical slant, like a recent one dealing with the plant collections of Richard Thomas Lowe (1802-1874) on the island of Madeira (Mesquita et al., 2022).  To put his work in context Joseph Banks and Daniel Solander visited there and made collections in 1768, early in James Cook’s first voyage around the world.  Francis Masson also did so a decade later.  Lowe was the next botanist to make significant collections on Madeira and the other islands of its archipelago, but his work was not confined to a brief visit.  He lived on Madeira from 1826 until 1852, much of that time as a clergyman.  He spent the rest of his life in England, but returned to Madeira for several months almost yearly.  When he died in a shipwreck in 1874, he was still working on his flora of Madeira that was published in several volumes. 

The Taxon paper covers the authors’ research on 2,280 of Lowe’s specimens that they were able to georeference, most now at the herbaria of the Natural History Museum, London and the Royal Botanic Gardens, Kew.  Positional uncertainty was noted.  Lowe’s locale descriptions varied in specificity but more than half rated as very high or high.  During his years of collecting, he had managed to visit most parts of the island, though not surprisingly, areas with steep slopes, of which there are many, were not visited.  His later work often involved returning to areas where he could find plant groups that posed taxonomic problems.  He was someone who came to know his research area well as is revealed in his writings.  His specimens were from 1350 locations and represented about 800 different taxa.  Lowe visited many locations a number of times, including at different times of the year.  He sent duplicates to several other botanists with exchanges of information on many taxa.  His correspondents included William Jackson Hooker, John Henslow, Robert Brown, Augustin de Candolle, and Philip Webb.  The researchers conclude:  “As a result of Lowe’s sustained and systematic approach, he is the single most prolific contributor to the study of Madeira’s endemic flora (p. 876).” 

Lowe’s work is an important contribution to biodiversity research because oceanic islands like Madeira have high proportions of endemic species and provide examples of rapid evolutionary radiations.  Also, because of the island’s size, populations are relatively low for many species, so having a historical record of occurrence in the past is helpful for present-day conservation efforts.  The fact that there were areas that Lowe found too remote or impossible to explore, including the many areas of cliffs, mean that these are good places in which to search for new species.  Equipment for scaling rock faces has improved, and even drones can be employed in survey work.   

There is much more to the article than I can recount in this post.  The number and content of the figures indicate how much analysis went into this paper and thus how much it says about Lowe’s contributions.  Maps are key, including the first figure, a topographic map with place names for Madeira and indicating just how much elevation variability exists there.  Next are more detailed historical maps and then a series of maps showing where Lowe’s georeferenced specimens were collected noting first locations, then precision of locations, followed by vegetation zones, and slope.  For slope, there is also a bar graph showing the relationship between slope and the amount of area at that slope. 

Then comes my two favorite graphics, or at least the ones I found most telling.  Figure 9 shows six maps of the island representing the itineraries for extended trips in six different years ranging from 1827 to 1860, including two Lowe made when he was no longer living there.  These are color-coded to show the months when each location was visited.  This is a good example of a well-designed graphic, as is figure 10, a graph that tracks with a line the number of specimens collected each year.  Then for each year it also gives bars indicating the percent of specimens from the six most common families on the island.  In most years, Lowe collected in all these families, but there are indications that, as mentioned in the text, he was focused on particular groups at certain points.  For example, in 1872, near the end of his collecting, Poaceae and Lamiaceae specimens were particularly well-collected.  Not coincidently they were the two families that had yet to be published in A Manual Flora of Madeira, which was left unfinished at the time of his death in a shipwreck that occurred when he was again bound for Madeira. 

I had never heard of Richard Lowe before I read this article, but it pains me that his flora was left unfinished.  The researchers who produced this work used the extensive data they generated from painstaking georeferencing and analysis to create not only a work of science but of history.  They created a portrait of a botanist and of work that will inform biodiversity research in the future and also support further study of the history of botany in Madeira.  They used specimen data and also delved deeply into Lowe’s correspondence and notes in a beautiful example of bioinformatics meeting the digital humanities.


Mesquita, S., Carine, M., Castel-Branco, C., & Menezes de Sequeira, M. (2022). Documenting the flora of a diversity hotspot: Richard Thomas Lowe (1802–1874) and his botanical exploration of Madeira island. TAXON, 71(4), 876–891.

Taxon and Digitization

Madhuca longifolia from Singapore, Royal Botanic Garden Edinburgh

This series of posts is looking at articles that have appeared in Taxon and deal with broader issues than the journal’s main fare of taxonomic treatments.  A timely article appeared a few months ago measuring how effective digital specimen images are in taxonomic research (Phang et al., 2022).  This study grew out of the COVID pandemic when access to collections was almost nonexistent in many parts of the world.  The authors were working on the genus Madhuca (Sapotaceae) for the Flora of Singapore.  Two were based in Scotland and one in Singapore, but all had the same access problem.  In this report they evaluated images of Madhuca collections from both Singapore and the adjacent Malaysian state of Johore.  The images were found in a number of JSTOR Global Plants and herbarium databases.  Another major source was the Royal Botanic Garden Edinburgh’s Sapotaceae Resource Centre (SRC) database, which also stores specimen and field images taken by researchers, often of material not otherwise available online. 

The overall result of the study was that while specimen images were valuable research tools, they could not provide all the information needed for a thorough taxonomic analysis.  In many cases, micro-morphological characters could not be seen clearly in digital images, even at high resolution, and these are precisely the characters often needed in defining the boundaries among species.  To provide quantitative results, the researchers rated the images as of high, medium, or low utility.  For the high ranking, an image needed to capture at least 5 qualitative and 3 quantitative macro-molecular characters.  Medium needed to meet the first criterion but not the second, and low had to have 4 qualitative characters. 

The report provides in-depth analysis of the results that I’ll just briefly recap here.  Not surprisingly, the specimen images found in herbarium databases ranked more highly than those in the SRC that were taken by researchers.  It wasn’t always a matter of the image quality that was the problem, but the absence of a ruler tool, like the one found in JSTOR Global Plants, or at least a measurement bar as a standard.  The authors also reported:  “Of the 219 specimen images examined, 125 (comprising 103 researcher images and 22 institutional images) had macromorphological characteristics hidden from view due to the low resolution of the image, the way the specimen had been mounted onto the herbarium sheet or had portions placed in an unopened capsule on the sheet” (p. 1068).  Herbarium databases varied in terms of the image resolution available.  Better quality images could probably be obtained by contacting the institution, but this often wasn’t possible during COVID, and in any case, would add steps to the taxonomist’s work. 

Other findings were that fruit and seed measurements were difficult for all images, with very few fruiting specimens available.  This was in part because there were usually only a few specimens for each species under study, a reminder of the crying need for continued collection, particularly in biodiverse areas with many species having either small populations or limited ranges.  Over all, the taxonomists were only able to identify 22% of the Madhuca species from researcher images, that number rose to 34% with institutional ones, and to 94% with physical examination of the specimen when the Singapore herbarium was again accessible.  This last figure resulted not only from microscopic examination of specimens, but from being able to closely examine flowers and fruits and open fragment packets.  The major message of the study is that online resources are very valuable for taxonomic investigations, but don’t come close to replacing specimens themselves.

It’s important to remember that there are many uses for online collections that don’t necessarily require such close study.  Virtual access is sufficient for many uses, especially when the access is through an information-rich database that’s easy to use.  Usability was the focus of a post on the Natural Sciences Collection Association website written by Teagan Reinert and Karen Bacon of the National University of Ireland, Galway.  It is a brief, but valuable recap of what determines a database’s rating anywhere from “very easy” to “usable but frustrating.”  It articulates what many of us experience subliminally as we search for specimens. 

To take the frustrating end of the spectrum first, there are sites that may have long loading time, low-quality images, return many irrelevant results, or “just don’t work.”  Sometimes a keyword search is handy, but the advanced search should be easy to find, and it’s great if searches by date range or cultivated species are easy to do.  Databases like those of New York Botanical Garden or the Royal Botanic Garden Edinburgh are given high marks because all the basic information on a specimen is shown without having to click further or open several screens.  The latter is particularly cumbersome if many specimens need to be accessed.  As for images, good quality is definitely a plus; also useful is an easy way to tell if there are differently sized images available.  For each image it should be clear what the license status is, such as public domain or creative commons license.  I find this very helpful, as is the last suggestion in the post:  “How the image or specimen data should be cited should be stated very clearly on the website either on its own easily accessed or clear labelled page, or on the specimen’s landing page. . . . But that information can sometimes be hidden in Frequently Asked Questions or on the bottom of a page that isn’t entirely relevant.”  Amen.


Phang, A., Atkins, H., & Wilkie, P. (2022). The effectiveness and limitations of digital images for taxonomic research. TAXON, 71(5), 1063–1076.

Taxon and the Flora of Brazil

Title page of first part of Flora Brasiliensis (1840-1906), Biodiversity Heritage Library

I belong to the International Association for Plant Taxonomy (IAPT) not because I am a plant taxonomist, but because I want to learn about the field.  Its journal Taxon is particularly helpful in this regard, though I can’t say that I read it cover to cover.  The articles I find most interesting take a broad view of the field, delve into its history, or deal with nomenclatural issues.  In this series of posts, I’ll highlight a few recent items I found particularly informative, beginning with one having a hefty 980 contributors, the Brazil Flora Group (2022).  The author list is shorter, but still lengthy, and suggests the massive collaboration underlying the creation of a Brazilian flora.

The impetus for the project began in response to the Global Strategy for Plant Conservation (GSPC), adopted in 2002 by the parties to the Convention on Biological Diversity.  The plan’s first target was to publish a list of the world’s plants by 2010, with plants broadly defined as including algae and fungi.  In 2010 Brazil published an online “List of the Species of the Brazilian Flora” and a “Catalogue of Brazilian Plants and Fungi,” which documented 40,989 species of algae, fungi, and plants.  By that time the second target of an online World Flora by 2020 was looming.  Since Brazil is a large country with great biodiversity, these tasks were themselves correspondingly massive, especially since the last Flora Brasiliensis was published from 1840-1906 and ran to 15 volumes, documenting 19,629 species in Brazil. 

An online information system was created for the Brazilian flora species list, and it was further developed for the task of constructing an online flora.  Between 2010 and 2015, 430 specialists were involved in adding new species to the list, updating determinations, and contributing descriptive data.  In the following five years, 554 more taxonomists joined the project, then called Brazilian Flora 2020.  The Taxon paper is essentially a review of the results of this work, including what it revealed about future needs in discovering and protecting Brazil’s biodiversity and supporting the taxonomic work necessary to accomplish these goals.  Meanwhile there was another project called “Plants of Brazil: Historic Rescue and a Virtual Herbarium for Knowledge and Conservation of the Brazilian Flora—Reflora.”   It’s aim was to develop a virtual herbarium that included specimens not only from the large collection at the Rio de Janeiro Botanic Garden, but also from the Royal Botanic Gardens, Kew and the National Museum of Natural History, Paris.  These are among the many European institutions with significant collections of tropical plant specimens because of their former colonial enterprises.  The Reflora infrastructure made it possible to upload images, curate specimen records with updated identifications, and add geographic and distribution data.  As this work progressed specimens from many more collections were added so that researchers now have access to millions of specimens through the Reflora Virtual Herbarium

As a result of this work, the Brazil Flora Group was able to report that by December 31, 2020 there were 46,975 known algae, fungi, and plants in Brazil, with 19,669 endemics.    These include 6,320 fungi, 4,993 algae, 1,610 bryophytes, 1,403 ferns and lycophytes, 114 gymnosperms, and 35,549 angiosperms.  This is hardly a complete count; some areas are under collected.  The most substantial collections have come from the Cerrado and also the Atlantic Rainforest, an area that has suffered from overdevelopment with loss of native vegetation.  Regions like the Caatinga and Pantanal are less well sampled.  There was also great disparity in the rates of increase in different types of species.  Amazingly, there was a 75% rise in the number of known fungal species between 2010 and 2020, an indication of the fungal richness yet to be discovered.  Not coincidentally the largest mycological collections are in the three states where the greatest number of mycologists are located.  Angiosperm numbers, on the other hand, only increased by 7%.  Interestingly, the number of known species in the heavily sampled Brazilian Atlantic Rainforest and Cerrado actually deceased between 2015 and 2020.  Yes, new species were named, but identification of synonymies and deletion of erroneous records more than offset this increase.

The article, as befits the massive size of the project it describes, is filled with data and insights.  The Brazil Flora Group focused on a number of areas that need attention if future GSPC targets are to be met.  One major issue is the need to build a stronger taxonomic infrastructure in the country, concomitant with its biodiversity.  With almost 1,000 taxonomists involved in the flora, expertise from around the world has been marshalled and will continue to support Brazil’s efforts, but it is no substitute for expertise within the country.  What is called the “taxonomic impediment,” lack of facilities and taxonomists, is a worldwide problem, as is the second area of concern: georeferencing.  Only about half the occurrence records in the Global Biodiversity Information Facility (GBIF) have coordinates and only a third of these have uncertainty information, which is essential for spatial analyses.  Geographic data are particularly important in conservation efforts.  As was mentioned earlier, also of concern is the issue of under-sampled areas, and along with this, species and families that have been neglected taxonomically.  So there is much work to do, but still, this report is also a celebration of wonderful accomplishments.


Group, T. B. F., Gomes-da-Silva, J., Filardi, F. L. R., Barbosa, M. R. V., Baumgratz, J. F. A., Bicudo, C. E. M., Cavalcanti, T. B., Coelho, M. A. N., Costa, A. F., Costa, D. P., Dalcin, E. C., Labiak, P., Lima, H. C., Lohmann, L. G., Maia, L. C., Mansano, V. F., Menezes, M., Morim, M. P., Moura, C. W. N., … Zuntini, A. R. (2022). Brazilian Flora 2020: Leveraging the power of a collaborative scientific network. TAXON, 71(1), 178–198.

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.

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.


Moraes, P. L. R. (2009). The Brazilian herbarium of Maximilian, Prince of Wied. Neodiversity, 4(2), 16–51.

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).

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.

Humanistic Uses of Herbaria

Secret of the Ferns by Anselm Kiefer in the Margulies Collection

This series of posts (1, 2) is highlighting projects sponsored by the Dumbarton Oaks Museum and Research Library.  I’ve already mentioned their Zoom presentations.  One particularly notable one was held in March 2021 and dealt with Humanistic Uses of Herbaria.  It was hosted by New York Botanical Garden Humanities Institute along with Dumbarton which is home of the Plant Humanities Initiative along with JSTOR Labs.  There were four speakers that day, all stars in their respective fields.  First was Barbara Thiers, now Director Emerita of the NYBG herbarium, leader in many endeavors to digitize herbarium collections, and author of Herbarium (see earlier post).  She gave a great introduction to the history and importance of herbaria and was followed by Pam Soltis, curator at the University of Florida’s Museum of Natural History, one of the leads on the iDigBio project to digitize specimens, and an expert on evolutionary genetics and ecology.  She presented on the future of research using herbarium specimens.  Anatole Tchikine, curator of rare books at Dumbarton, was the next speaker and discussed herbaria in the collection including a very interesting manuscript by the landscape architect Samuel Parsons, Jr., Leaves of Hardy Oaks and Maples.  He created it early in his career as a way to study leaf form and how this might affect the shade a tree produced. 

Yota Batsaki, executive director of Dumbarton Oaks, gave the last presentation on “The Apocalyptic Herbarium: Anselm Kiefer’s Secret of the Ferns (2007).”  She dealt with the herbarium as metaphor for the deterioration of life, particularly plant life, on earth.  Kiefer is among the most noted postwar German artists and often uses plants in his works.  He was born in 1945 in a Berlin bunker and played in ruble as a child, so it is not surprising that the war and the holocaust have been among his major themes.  Many of his paintings, often multimedia works, are devastated landscapes, with thick layers of paint, sometimes splashed with molten lead and embedded with dried plants (Biro, 2013). 

As Batsaki noted, more recently Kiefer has turned to other forms of devastation, including thoughtless abuse of the earth.  In dealing with this theme he uses some of the same tropes he employed in earlier work.  In particular, she discusses a large installation, Secret of the Ferns, that fills a room at the Margulies Collection in Miami.  At the center of the room are two concrete bunkers that have seen much wear and tear.  A two-story one is at the back and in front of it is one with a pile of coal at its entrance.  Along the left and right walls are hung large framed works in two ranks on each side, 48 in all towering over the viewer. 

Kiefer’s installations are complex; there are many layers to them and many details.  I was intrigued by Batsaki’s presentation but I knew I was missing some of the nuances.  I hoped that she would publish on this topic, and she has, in an article in Environmental Humanities (2021).  Through the text and images of the work, I was able to dig more deeply into it.  While I was somewhat familiar with Thiers and Solitis’s work, this was another realm.  Here were plants being used in a very different way, not to reveal information about genetics or environmental change, but to get at deep questions of what humans value and how they relate to other forms of life on earth.  Batsaki does a great job of “interrogating” the work.  This is a term that scholars in the humanities use regularly, but it is sort of foreign to me.  Interrogating living things seems rather aggressive and is a verb seldom used by biologists, though many of their techniques can be quite aggressive.  It is an example of how science and the humanities have to learn each other’s language and attitudes if they are to do more than just meet occasionally as at the March seminar.

Hanging to the left and right of the bunkers, many of the frames hold large pressed fern fronds against a dark background, in some cases, with the stipe appearing to rise from dried, cracked earth.  Each is encased in what Batsaki describes as a vitrine and framed in lead, a common material for Kiefer, who is intrigued with its role in alchemy.  The ferns relate to the installation’s title which is from a Paul Celan poem.  Celan’s work, often dealing with aspects of German history and the holocaust, has proved a rich reservoir of inspiration for Kiefer.  The artist is drawn to ferns because of their long history on earth as the earliest vascular plants and one source of the organic material in coal.  Their presence in the frames is tied to the coal by the bunker:  a reminder that burning coal has led to disastrous changes in the earth’s atmosphere that threatens the long-resilient ferns and all life on earth, what Batsaki describes as the “slow violence of extinction.” (p. 394). 

In this short post, it’s impossible to do justice either to the artwork or to the essay.  There is a great deal here as Batsaki investigates a variety of themes including that of transformation, examining how ferns were long thought to be mysterious because they did not form seeds and so their mode of reproduction was unknown until their tiny spores were studied in the mid-19th century.  One line from her essay that I find particularly memorable is: “If the herbarium started as an aide to memory [in the early modern era], the installation transforms it into a vehicle for memorialization.” (p. 409).  Unfortunately too many sheets in herbaria serve the same function for extinct species.


Batsaki, Y. (2021). The Apocalyptic Herbarium: Mourning and Transformation in Anselm Kiefer’s Secret of the Ferns. Environmental Humanities, 13(2), 391–413.

Biro, M. (2013). Anselm Kiefer. New York: Phaidon.

Botany and Art: Intimacies

Lupinus argenteus, Silver Lupine” by Susan Rubin

The last post discussed how herbarium sheets are sometimes collages with illustrations of different kinds attached along with the plant material.  There was an interesting case in Taxon recently of an illustration used to identify a type specimen (Fleischmann and Gonella, 2020).  The species in question is Drosera intermedia, an insectivorous plant found from eastern North America, through the Caribbean to tropic South America.  As with many plants, particularly those with a relatively long botanical history, nailing down the first publication of a name and the type specimen can be complicated.  The authors here wade through the literature and cite a 1798 publication by Johann Dreves and Friedrich Hayne, though a 1800 publication by Hayne is usually given.  Why I find this case interesting is that Fleischmann and Gonella argue that a specimen in the Munich herbarium is the lectotype because it so closely resembles the illustration of the plant in the 1798 publication.  It is known that Haynes himself did the drawing on which it is based. 

This seems relatively straightforward, except for the fact that there is no indication on the sheet linking the specimen to Haynes.  The handwriting on the label is that of Johann Christian von Schreber, who traded and bought plants from a number of botanists.  This sheet is part of a Schreber collection acquisitioned in 1813 by the herbarium in Munich’s Bavarian Natural History Collections.  Also on the sheet is a not in the handwriting of Albrecht Roth, who was an early proponent of the idea that plants could attract and digest insects and thus derive nourishment from them.  Schreber thought this outlandish.  Sending the plant to Schreber was less about taxonomy and more about plant physiology.  In the note Roth writes that “the incurved leaves [of the specimen] hold dead insects.”  Roth published an article in which he remarked that he had received Drosera from Haynes with insects trapped in the leaves, providing evidence for linking Haynes’s illustration to Schreber’s specimen through Roth. 

This is a case of what I would call investigative botany, practiced by those taxonomists who also have a love of history.  The “excuse” is to find type specimens for species that are untypified or mis-typified, but it is also a way to satisfy an urge to solve a mystery.  Here the hunt was made more challenging, and perhaps therefore more intriguing, because the fate of the bulk of Haynes’ herbarium is unknown, and a search of what does exist turned up nothing related to the Drosera.  It’s suggestive of the more casual attitude toward specimens used in describing a species at that time that Haynes sent at least one of them on to Roth, and then Roth passed it on to Schreber in service of his insectivore argument.  It took dogged work to link the specimen’s provenance to the illustration in the original description, which is very similar.

My other two examples of intimate relationships between specimens and art are of a different kind and definitely tend toward the artistic rather than scientific end of the spectrum.  The first is a painting I saw on the web some time ago, and it keeps coming to mind.  It is “Lupinus argenteus, Silver Lupine” by Susan Rubin.  It won the Group Gold Medal from the Royal Horticultural Society London in 2017.  It’s a work of trompe-l’oeil and shows a herbarium specimen of the lupine, with faded colors and all the associated trappings of such a sheet.  This one is stamped from the Denver Botanic Gardens (where Rubin teaches) and includes a typed label, accession number, and barcode sticker.  Overlaid on it is a fresh lupine flower with its beautiful blue-purple inflorescence and green leaves.  The cutting has a small paper label and casts a shadow on the sheet suggesting it has merely been placed there for a moment to compare the live and dead specimens. 

Not surprisingly, Rubin is a botanical artist and much of her work is more traditional, though tending toward the artistic rather than the documentary.  She has done a series of trompe-l’oeil paintings, but none of the others have a herbarium specimen.  They show illustrations, sometimes taped or pinned to an artist’s table along with notes, preparatory sketches, a pencil or two, and other tools of the trade.  Somehow, these additions make the work more lively as it seems in the act of becoming.  The lupine is an indication of the accuracy of her work, and how it is grounded in the plant itself. 

Finally, I want to mention a rather odd convergence of art and science.  This was brought to my attention by the Swedish historian of science Anna Svensson, whose dissertation is a wonderful example of how history, botany, art, and the digital environment can be interwoven.  Anna spent some time at the Botanical Garden in Florence hunting among its treasures.  One that she found was a small bound herbarium where some of the flowers were painted over to give them more color.  I’ve written about early herbaria where missing petals or leaves were painted in, but the plants themselves were unadorned.  The Florence example went a step further.  It’s definitely at the far, far end of the scientific/artistic spectrum and a very unscientific move, but fascinating nonetheless. 


Fleischmann, A., & Gonella, P. M. (2020). Typification and authorship of Drosera intermedia (Droseraceae). Taxon, 69(1), 153–160.

Note: I would like to thank Susan Rubin for allowing me to use her art in this post.