Just three decades after Spanish conquistador Francisco Pizarro first encountered the potato in the high Andes of Peru in 1532, the potato was already being grown in the Canary Islands. And it found its way to mainland Europe via the Canaries shortly afterwards [1].

The potato was described by English herbalist John Gerard in his Herball published in 1597. In a revised version, published in 1633 over 20 years after his death, there is another beautiful woodcut of the potato, referred to Battata Virginiana or Virginian potatoes.

Potatoes became an important crop by the late 18th century, and particularly the staple of Ireland’s impoverished citizens in the years leading up to the Irish Potato Famine of the mid-1840s.

Today, potatoes are one of the world’s most important crops, grown in every continent except Antarctica. Known scientifically as Solanum tuberosum, it was given this name by the famous Swedish naturalist, Carl Linnaeus in his 1753 magnum opus, Species Plantarum.

The potato and its wild relatives must be one of the most studied groups of crop plants. Not that I’m biased (having researched potatoes for more than 20 years).

Potato diversity and germplasm collections
Its clear that there is a wealth of information about the diversity within the section of the genus Solanum that encompasses the potato. They have been studied extensively from a taxonomic point of view, breeding efforts worldwide have incorporated genes from many wild species to enhance productivity, and important germplasm collections were set up decades ago to preserve this important diversity, to study it, and use it in potato breeding.

Among the most important collections are held at:

• the International Potato Center in Peru (conserving the world’s largest collection of indigenous varieties from the Andes, as well as wild species);
• the Commonwealth Potato Collection at the James Hutton Institute in Scotland (a large collection of predominantly wild species);
• the NRSP-6 – United States Potato Genebank at Sturgeon Bay, Wisconsin (mainly wild species); and
• the CGN Potato Collection (formerly the Dutch-German Potato Collection maintained at Braunschweig, Germany) at Wageningen, the Netherlands.

The wild relatives of the potato have one of the broadest geographical and ecological ranges among species that have been domesticated for human consumption. While the various forms of cultivated potatoes were domesticated in the Andes of Peru and Bolivia, and on the coast of Chile, the wild species are found from the southwest USA (in the coniferous forests of Arizona, for instance) through Mexico and the countries of Central America to Panama, along the Andes south to Chile and northern Argentina, and south and east on to the plains of Argentina, Brazil, Paraguay and Uruguay. Wild species are found in the coastal desert of Peru, in the cloud forests of central America to almost 3000 m, at the highest altitudes of the Andes, well over 4000 m, and also growing in the highly humid transition zone on the eastern side of the Andes dropping down to the lowland forests (known as the ‘eyebrow of the mountain’ or ceja de la montaña).

Here is just a very small sample of the diversity—and beauty—of wild potato species (photos courtesy of my friends at the Commonwealth Potato Collection).

How many potato species are there?
Well, it depends, to some extent, on one’s perspectives as a taxonomist, use of different species concepts, and the methods used to study species diversity, and also on the work that earlier taxonomists published.

Essentially, there are three basic taxonomic approaches:

• Morphology: often based on the study of dried herbarium specimens collected in the wild. In the case of potatoes, this has led to the description of a multiplicity of species, with almost every variant being described as a separate species. This reliance on plant morphology was the approach taken by the 19th and early 20th century botanists.
• Biosystematics: takes an experimental view of species diversity, of breeding behaviour and relationships, and very much based on collections in the field and the study of ecology, and growing samples in a uniform environment such as the study one of my PhD students, Susan Juned, made of Solanum chacoense, a species from Argentina and Paraguay.
• Molecular biology: methods have become available in the last couple of decades to analyse the most basic variation in DNA, and helped to refine further how potato taxonomists view the diversity within the tuber-bearing Solanums, and the relationships between species.

While these different approaches still do not provide a definitive answer to the question of how many species there are, we know that taxonomists have described and named more than 200 species. To some extent it’s like asking how long is a piece of string. And that helps me to provide an analogy.

Take a piece of string. If you were to view this string along its length that, to your vision would be fore-shortened, it would be very difficult to say with any degree of certainty just how long the string actually was. However, if you increase the angle at which you view the string, until you are looking at right angles, your ability to estimate its length also increases. At right angles you can see the whole length, and measure it accurately in many different ways.

Taxonomic study is a bit like looking at the string from different angles. Each taxonomist builds on earlier studies, and describing new species or subsuming previously described ones into another species (as merely variants). This is one of the challenges of studying wild potato species: they are highly variable and show considerable phenotypic (or morphological) plasticity. It’s not always possible to study large numbers of plants under uniform conditions to reduce the variation caused by differences in habitats.

Furthermore potatoes have considerable chromosomal variation, with a base number of x=12, with diploids (2n=24) the most frequent, and mostly self-incompatible (i.e. they cannot self fertilise), infertile triploids (2n=36, including two cultivated species), tetraploids with 2n=48 (mostly self-fertile, and including the cultivated Solanum tuberosum of world-wide agriculture), some pentaploids (2n=60; including one cultivated form), and a few hexaploids with 2n=72. Wild potatoes are uncommonly promiscuous when grown together under experimental conditions, and will inter-cross readily (they are bee-pollinated), yet hybrids often do not survive beyond the second generation in the wild. Many species are separated by ecology, and generally do not come into contact with each other, thus maintaining their species identity.

Nevertheless, this is what makes the study of potatoes and wild species so very interesting, and that captured my interest directly for over two decades, and continues to do so, even though I moved on to the study of other crops like rice and grain legumes.

The potato taxonomists
Many botanists have taken an interest in wild potatoes. During the 19th century, the Swiss-French botanist Alphonse de Candolle (d. 1893) named a number of species, as did François Berthault (d. 1916). But the first decades of the 20th century leading up to the Second World War saw a lot of collecting and taxonomic description. In Germany, Friedrich August Georg Bitter, who specialised in the genus Solanum, described and named many species. However, it was the involvement of several Russian botanists and geneticists, under the leadership of Nicolai Vavilov, that saw an expansion in the collection of potatoes throughout the Americas, but a systematic evaluation of this germplasm leading to even more species being described.

Two names come to mind, in particular: SM Bukasov and VS Juzepczuk. They were active during the 1920s and 30s, taking part in several missions to South America, and developing further the concept of potato species. But much of their work was based on morphological comparison leading to the identification of even small variants as new species.

In August 1938, a young Cambridge graduate, Jack Hawkes, traveled to Leningrad in Russia to meet and discuss with Bukasov and Juzepczuk (and Vavilov himself) in preparation for the 1938-39 British Empire Potato Collecting Expedition to South America (which Jack has described in his 2004 memoir Hunting the Wild Potato in the South American Andes [2]).

That collecting expedition, and the subsequent studies (which led to Hawkes being awarded his PhD from the University of Cambridge in 1941 for a thesis Cytogenetic studies on South American potatoes supervised by renowned potato scientist Sir Redcliffe N Salaman), was the launch pad, so to speak, of potato taxonomy research for the rest of the 20th century, in which Hawkes became one of the leading exponents.

After Cambridge, Hawkes spent some years in Colombia (where he no doubt continued his studies of wild potatoes) but it was on his return to the UK in 1952 when appointed to a lectureship in the Department of Botany at The University of Birmingham (where he was to remain until his retirement in 1982) that his potato studies flourished, leading him to publish in 1956 his first taxonomic revision of the tuber-bearing Solanums (with a second edition appearing in 1963).

In 1990, he published his final synopsis of the tuber-bearing Solanums [3]; that taxonomic treatment is the one followed by the curators of the Commonwealth Potato Collection.

Jack’s approach to potato taxonomy was based on a thorough study of morphology backed up by rigorous crossing experiments, and a cytogenetic and sometimes serological evaluation of species relationships.

I first met Jack in February 1970 when he interviewed me for a place on his newly-founded MSc course on plant genetic resources, joining the course later that same year. In September 1971 I became one of Jack’s PhD students, joining others who were looking at the origin and evolution of the cultivated species [4].

Donovan S CorrellIn these revisions he was also taking into account the work of US botanist, Donovan S Correll who published his own potato monograph in 1962 [5], as well as three important South American botanists with whom he would collaborate from time-to-time: Professor César Vargas from the National University of Cuzco; Professor Martín Cárdenas from Cochabamba in Bolivia; and Professor Carlos Ochoa, originally from Cuzco, who was a professor at the Universidad Nacional Agraria (UNA) in La Molina, Lima and, around 1975 or so, joined the International Potato Center across the street from the UNA.

Vargas published a number of species descriptions in the 1950s, but made his most significant contribution in his two part monographs, Las Papas Sudperuanas published in 1949 and 1956. I met Vargas on a couple of occasions, first in January 1973 just after I’d joined CIP as Associate Taxonomist. And a second time in February 1974 when I was passing through Cuzco with Dr Peter Gibbs from the University of St Andrews in Scotland. Peter was making a study of incompatibility among different forms of the Andean tuber crop, oca (Oxalis tuberosa), and had joined me on an excursion to Cuyo-Cuyo in the Department of Puno. Vargas’s daughter Martha was studying for her MSc degree under Peter’s supervision at St Andrews.

It was Carlos Ochoa, however, whose studies of potatoes and their relatives rivalled (and in some respects eclipsed) those of Jack Hawkes. They were quite intense taxonomic rivals, with a not-altogether harmonious relationship at times. Carlos certainly played his taxonomic cards very close to his chest.

But the fact that he grew up in the Andes and had, from an early age, taken an interest in the diversity of this quintessential Andean crop and its wild relatives, led him to dedicate his life to uncovering the diversity of potatoes in his homeland. He was also a potato breeder and released some of the most important varieties in Peru, such as Renacimiento, Yungay, and Tomasa Condemayta.

In this video (in Spanish, and broadcast on Peruvian TV on his death in 2008) he talks about his early life in Cuzco, the pressures on him to study medicine or become a lawyer, and how he found his true vocation: the study of wild potatoes.

Setting potato taxonomy and germplasm exploration priorities at CIP
Forty-five years ago this week, CIP convened the first planning workshop on the exploration and taxonomy of potatoes [6], inviting a group of taxonomists and potato breeders to meet in Lima and mull over the ‘state of play’ taking into consideration what taxonomic research had already been accomplished, what was in the pipeline, and what CIP’s germplasm exploration policy (especially in Peru) should be. I attended that meeting (as an observer), having landed in Lima just a few days earlier.

On the taxonomic side were Jack Hawkes, Carlos Ochoa, and Donald Ugent who was a ethnobotany professor at Southern Illinois University in Carbondale. Richard Tarn, a potato breeder from Agriculture Canada at Fredericton, New Brinswick, had completed his PhD under Jack’s supervision at Birmingham. Frank Haynes, a professor of genetics and potato breeder at North Carolina State University (and long-time friend and colleague of CIP’s first Director General, Richard Sawyer) and Roger Rowe [7], then curator of the USDA’s potato collection at Sturgeon Bay (who would join CIP in July 1973 as the Head of Breeding and Genetics, and become my PhD co-supervisor) were the other participants.

In 1969, Jack had published (with his Danish colleague Peter Hjerting [8]) a monograph of the potatoes of southern cone countries of South America [9], and by the time of the CIP 1973 workshop was well into research on the potatoes of Bolivia [10], leading publication of a monograph in 1989.

What I’ve never been able to fathom after all these years is why Ochoa decided to write his own monograph of the Bolivian species rather than concentrating in the first instance on the Peruvian species. Nevertheless Ochoa did produce his own fine monograph in 1990 [11], beautifully illustrated with some fine watercolours by CIP plant pathologist Franz Frey. This was followed by an equally magnificent volume on the potatoes of Peru in 2004 [12], also illustrated by Frey.

Throughout his expeditions and research, Ochoa was supported by several assistants, the most notable being Ing. Alberto Salas. Now in his mid-70s, he has been collecting wild potatoes for five decades.

I knew Alberto when I first joined CIP in 1973, and it was a delight to meet him again (although he had retired) during my visit to CIP in July 2016.

Taking up the baton
With retirement, Hawkes and Ochoa passed the potato taxonomy baton to a new generation of researchers, principally David Spooner, a USDA scientist at the University of Wisconsin-Madison who made several collecting trips throughout the Americas.

David’s research took potato systematics to a new level, employing the developing molecular and genomic approaches, and use of different classes of markers to help him refine his understanding of the diversity of the tuber-bearing Solanums, building of course on the very solid Hawkes and Ochoa foundations.

Although no longer working on potatoes (his most recent focus on carrots supported the PhD thesis of Carlos Arbizu, Jr, the son of one of my PhD students at Birmingham in the 1980s), David’s scientific output on potatoes has been prodigious. With molecular insights supporting more traditional methods he has proposed a 50% reduction in the number of potato species from the more than 200 listed in Hawkes’s 1990 publication.

Is this the end of the potato taxonomy story? Probably for the time-being. It’s unlikely that anyone will pursue these studies to the same depth as Hawkes and Hjerting, Ochoa, or Spooner. Nevertheless, as the curators of the Commonwealth Potato Collection have done, most potato researchers will take a pragmatic approach and fix on a particular taxonomic treatment on which to base their management or use of germplasm. Taxonomy is one of those disciplines in which subjective interpretations (obviously based on empirical studies of diversity) can lead to contrary classifications. What is a distinct species to one taxonomist may be merely a variant to another. Undoubtedly these different taxonomic treatments of the tuber-bearing Solanums have permitted us to have a much better appreciation of just how long ‘the potato piece of string’ really is.

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[1] Hawkes, JG & J Francisco-Ortega, 1993. The early history of the potato in Europe. Euphytica 70, 1-7.

[2] Hawkes, JG, 2004. Hunting the Wild Potato in the South American Andes – Memories of the British Empire Potato Collectiing Expedition to South America 1938-1939. Wageningen, the Netherlands. ISBN: 90-901802-4.

[3] Hawkes, JG, 1990. The Potato – Evolution, Biodiversity and Genetic Resources. Belhaven Press, London.

[4] Since I was working on the origin and evolution of a cultivated species of potato for my PhD, I made only one short collecting trip for wild species with Jack in early 1975, to the Departments of Huanuco, Cerro de Pasco, and Lima. On his trips to Peru between 1973 and 1975 he would join me in the field to look at the germplasm I was studying and give me the benefit of his potato wisdom.

[5] Correll, DS, 1962. The Potato and its Wild Relatives. Contributions from the Texas Research Fiundation 4, pp. 606. Texas Research Foundation, Renner, Texas.

[6] International Potato Center, 1973. Report of the Workshop on Germplasm Exploration and Taxonomy of Potatoes. Lima, Peru. 35 pp.

[7] I’ve kept in touch with Roger and his wife Norma all these years. After I left CIP in 1981, Roger moved to East Africa to work with the animal diseases center that became ILRI after its merger with another CGIAR livestock center in Ethiopia. He was DDG-Research at CIMMYT in Mexico in the late 1980s and early 1990s. While I was at IRRI, he was based in Cairo working for the CGIAR center that became WorldFish (with its headquarters in Penang, Malaysia). Before it moved to Malaysia, ICLARM as it then was had its offices in Manila, and we would see Roger in the Philippines from time-to-time. It was great to meet up with Roger and Norma again in July 2016 when I was in Lima for the genebank review that I led.

[8] From what I can determine through a Google search, as of January 2018, Peter celebrated his 100th birthday in 2017. He has a Mexican tetraploid (2n=4x=48) species named after him, Solanum hjertingii. When I was at Birmingham in the 1980s I had two PhD students, Lynne Woodwards and Ian Gubb who studied this species because its tubers lack so-called enzymatic blackening, a trait that could be very useful in potato breeding.

[9] Hawkes, JG & JP Hjerting, 1969. The Potatoes of Argentina, Brazil, Paraguay, and Uruguay – A Biosystematic Study. Annals of Botany Memoirs No. 3. Clarendon Press, Oxford.

[10] Hawkes, JG & JP Hjerting, 1989. The Potatoes of Bolivia – Their Breeding Value and Evolutionary Relationships. Clarendon Press, Oxford.

[11] Ochoa, CM, 1990. The Potatoes of South America: Bolivia. Cambridge University Press, Cambridge.

[12] Ochoa, CM, 2004. The Potatoes of South America: Peru. International Potato Center, Lima, Peru.