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Maps of Model for Ullucus tuberosus domestication

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Semi-domesticated forms. Semi-domesticated forms of ulluco are thought to have long, trailing stems and smaller tubers similar to wild forms of ulluco, but palatable tubers similar to domesticated forms. Justin Haggard
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1. Maps of Model for Ullucus tuberosus domestication
Domesticated forms. Domesticated forms of ulluco have compact growth habits with upright stems and short internodes. Gaps in the range of this second step of domestication could explain why some semi-domesticated forms with longer stems continue to exist Justin Haggard
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1. Maps of Model for Ullucus tuberosus domestication

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Ulluco: Resilience and Preservation Through Time Justin Haggard Ullucus tuberosus (ulluco) is a crop grown throughout South America for its starchy tubers and spinach-like leaves. Grown alongside humans for thousands of years, the genetic diversity of ulluco along with indigenous knowledge of the crop has enabled it to survive amidst challenges of Spanish colonial rule. One challenge was competition from introduced European cultivars such as barley and carrots. Another challenge was that Indigenous groups who cultivativated ulluco faced hardships and challenges throughout colonization that echo into modern times. Despite these barriers, Indigenous Andean groups continued cultivating ulluco on smaller scales. This preserved Indigenous agricultural methods, and therefore ulluco biodiversity, still today. Now, ulluco has been proposed as a crop that could help solve diminishing food supplies amidst climate change. By Audrey Hernandez, Justin Haggard #Ulluco
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Ulluco: Resilience and Preservation Through Time Ullucus tuberosus (ulluco) is a crop grown throughout South America for its starchy tubers and spinach-like leaves. Grown alongside humans for thousands of years, the genetic diversity of ulluco along with indigenous knowledge of the crop has enabled it to survive amidst challenges of Spanish colonial rule. One challenge was competition from introduced European cultivars such as barley and carrots. Another challenge was that Indigenous groups who cultivativated ulluco faced hardships and challenges throughout colonization that echo into modern times. Despite these barriers, Indigenous Andean groups continued cultivating ulluco on smaller scales. This preserved Indigenous agricultural methods, and therefore ulluco biodiversity, still today. Now, ulluco has been proposed as a crop that could help solve diminishing food supplies amidst climate change. By Audrey Hernandez, Justin Haggard #Ulluco Nomenclature and Distribution
The Linnean binomial name is Ullucus tubersosus Caldas, which became official in 1809 in Nueva Granada (roughly encompasses present-day Colombia) (Royal Botanic Gardens Kew, n.d.). There are several common names for this plant. Historically, it was called the Quechua terms Ulluku or Ulluma, which means tuber (Slow Food Foundation for Biodiversity, n.d.). Today, in Peru and Argentina where it’s native, it’s largely referred to as ulluco (Kay, 1987). The Spanish name is papa lisa. In Ecuador and Colombia some common names are chigua, chuguas, melloco, and olloco. Today, ulluco is cultivated mostly in South America in Colombia, Ecuador, Chile, Venezuela, Bolivia, and Peru by families for sustenance or by farmers who sell their products in local markets (Specialty Produce, n.d.).

Morphology
There is significant morphological variation among specimens of Ullucus tuberosus across different growing regions (Parra-Quijano et al., 2012; Rousi et al., 1989). The simple leaves of ulluco can either be sagitate with acute apices, or reniform shaped with rounded apices (King, 1988). The leaves are smooth and succulent, and vary between lighter and darker shades of green or even reddish coloration (King, 1988). The leaves attach to the stem on a grooved petiole, and are arranged in an alternate pattern (King, 1988). The stems are ridged, and are either green, yellow, or reddish in coloration (King, 1988). Most domesticated ulluco has an upward and compact growth habit reaching 20 to 40 cm in height, but some varieties have trailing stems similar to the wild ancestor Ullucus aborigineus (Parra-Quijano et al., 2012; Rousi et al., 1989). Flowers are pedicellate, scentless, and typically shed shortly after the bud opens. Ovary is globose to somewhat pyriform and it’s slightly lobed (Royal Botanic Gardens Kew, n.d.). The stigma is weakly differentiated to obscurely 3-lobed and the plant has only 1 stamen (Royal Botanic Gardens Kew, n.d.).
When tubers are planted, inflorescences rise in nodes and open from base to apex of the plant (Pietila & Jokela, 1994). Often, ovule and embryo sac development begins but is slowed or stopped completely by the growth of the tuber. This is hypothesized to be the reason ulluco has several flowers but rarely produces seed and almost never produces viable seeds (Pietila & Jokela, 1994). Energy is used for tuber growth rather than flower development.

Evolutionary and Domestication History
For many years it was thought that there was no existing wild plant. Then in 1967, it was found and named Ullucus aborigineus. Its tubers are smaller, very bitter, and inedible. It has a thin, climbing, and long stem that can grow up to 2 meters (Parra-Quijano et al., 2012). This wild form is triploid, while cultivated forms are diploid (Parra-Quijano et al., 2012).
Ulluco coevolved alongside humans as they domesticated wild forms for cultivation. The growth forms of these domesticated crops vary widely between different growing sites (Parra-Quijano et al., 2012; Rousi et al., 1989). Different Columbian accessions, for example, showed large morphological variation between growing sites (Parra-Quijano et al., 2012). These variations included differences in tuber size, tuber color (including green, beige, magenta, and others), tuber shape, growth habit (erect or prostrate), stem color, and leaf shape (Parra-Quijano et al., 2012). Many similar morphological variations were also observed between different accessions in Peru (Rousi et al., 1989).
One interesting finding from the study of Columbian accessions is that the ulluco grown in the north-east Columbian Andes tended to have trailing stems and smaller tubers (Parra-Quijano et al., 2012). These traits make these accessions more similar to the wild ancestor, Ullucus aborigineus, and differentiate them from most other domesticated forms. Because of this, it seems that these specific accessions may be partially-domesticated, giving us insight into the transition between wild and domestic forms (Parra-Quijano et al., 2012).

Intersections with Human History
Ulluco is known for its high genetic diversity, which is largely due to traditional Indigenous cultivation practices that have been preserved for generations (Malice and Baudoin, 2009). This plant has not been introduced at a large scale in national markets partially due to its association with communities of low socio-economic status (Malice and Baudoin, 2009). From the Incan Empire through Spanish rule, ulluco has been cultivated by and sustained Indigenous Andean groups (Malice and Baudoin, 2009). Since it was domesticated and has been grown almost exclusively in the Andean highlands for thousands of years, it requires specific conditions and takes special knowledge to harvest. It must grow in cool, shady areas, and in rich soil over a long growing season (which spans from 5 to 9 months). Despite these restrictive conditions, recent successful attempts to grow ulluco outside of the Andes include New Zealand, some parts of Southern England, coastal areas of Europe, the Pacific Northwest, coastal areas of California, and the Appalachians in the United States (Specialty Produce, n.d.).
Human interaction with ulluco is largely confined to Indigenous groups in the high Andes. Botanical evidence of Ulluco has been dated as far back as 10,000 years ago (Spierling and King 1990). In coastal Peru, archaeologists found 5500 year old art depicting ulluco and containing traces of ulluco plant matter (Slow Food Foundation for Biodiversity, n.d.). Also, ulluco is depicted on ceremonial vessels from the Wari Empire and on qero ceremonial vessels from the post-Incan era (Arbizu and Tapia, 1994). Ulluco is one of the few crops that survived colonial rule (Slow Food Foundation for Biodiversity, n.d.).
Today, ulluco still has cultural importance for many people in South America. For example, in Bolivia, an important holiday is Semana Santa or Easter Holy Week. Many traditional dishes served during Semana Santa use ulluco (Specialty Produce, n.d.). In addition to feeding Andean communities, it is sometimes used medicinally. For example, native Ecuadoreans use it topically to treat skin conditions and consume it to treat digestive issues (Specialty Produce, n.d.).

Nutrition
Both the tubers and leaves of ulluco are consumed. The tubers are nutritionally similar to sweet potatoes or cassava (King 1988), and are a good source of carbohydrates (about 76 percent) and contain moderate amounts of protein (about 13 percent) but are low in calories (about 3.7 cal/g) (Nutritional analysis of three andean tubers). The tubers also contain pigments known as betalains, which are also found in beets and swiss chard, and have anti-inflammatory and antioxidant properties when ingested (Svenson et al., 2008).

Potential for Future
Recently, there has been an increase in scientific and agricultural interest in ulluco. As far as future applications, Ulluco has been identified as a possible resource for tackling diminishing food supplies in the face of climate change (Solá, 2021). To understand why, its ties to Indigenous communities should be highlighted. At the time of Spanish colonization in the 1500s, the Incan Empire had as many agricultural plant species as Europe and Asia combined, despite the lack of technologies such as iron, draft animals, and wheels that other civilizations had at the time (Solá, 2021). Thus, Andean civilizations were characterized as having incredibly high agrobiodiversity, largely due to polycultural practices and other traditional cultivation techniques. During Spanish rule, the plants of Andean communities were diminished as European crops replaced them (such as barley and carrots) (Solá, 2021). Despite this, traditional knowledge and practices have preserved ulluco agrobiodiversity, proving its resilience and strong ties to indigenous communities. Since agriculture relies heavily on a stable climate, and that stability is currently in jeopardy due to climate change, resilient crops like ulluco, as well as knowledge of ancient native cultivation practices, are important teachers (Solá, 2021).

Sources Cited
Arbizu, C., & Tapia, M. (1994). Andean tubers. Retrieved May 29, 2022, from https://hort.purdue.edu/newcrop/1492/tubers.html

Kay, D. E. (revised by Gooding, E. G. B.) (1987) Crop and Product Digest, No. 2- Root Crops, Second Edition. London: Tropical Development and Research Institute, xv & 380

King, S. R. (1988). Economic botany of the Andean tuber crop complex: Lepidium meyenii, Oxalis tuberosa, Tropaeolum tuberosum and Ullucus tuberosus (Order No. 8821097). Available from ProQuest Dissertations & Theses A&I; ProQuest Dissertations & Theses Global. (303662871). https://www.proquest.com/dissertations-theses/economi c-botany-andean-tuber-crop-complex/docview/303662871/se-2?accountid=14512

Malice, M. & Baudoin, J. (2009). Genetic diversity and germplasm conservation of three minor Andean tuber crop species. Biotechnologie, Agronomie, Société et Environnement. 13.

Parra-Quijano, M., Panda, S., Rodríguez, N. et al. Diversity of Ullucus tuberosus (Basellaceae) in the Colombian Andes and notes on ulluco domestication based on morphological and molecular data. Genet Resour Crop Evol 59, 49–66 (2012). https://doi.org/10.1007/s10722-011-9667-8

Pietilä, L., & Jokela, P. (1994). Developmental abnormalities in the ovule and embryo sac of Ulluco (Ullucus tuberosus) and their effect on seed set. Euphytica, 75(1-2), 31–39. https://doi.org/10.1007/bf00024529

Rousi, A., Jokela, P., Kalliola, R. et al. Morphological variation among clones of ulluco (Ullucus tuberosus, Basellaceae) collected in Southern Peru. Econ Bot 43, 58–72 (1989). https://doi.org/10.1007/BF02859325

Royal Botanic Gardens Kew. (n.d.). Ullucus tuberosus caldas. Plants of the World Online. Retrieved May 20, 2022, from https://powo.science.kew.org/taxon/urn:lsid:ipni.org:names:167633-1#other-data

Slow Food Foundation for Biodiversity. (n.d.). Nariño Ulloco - Ark of taste. Slow Food Foundation. Retrieved May 22, 2022, from https://www.fondazioneslowfood.com/en/ark-of-taste-slow-food/narino-olloco/

Solá , A. (2021, October 13). In Peruvian Andes, ancient crops hold promise for a climate-blighted future. Mongabay Environmental News. Retrieved May 25, 2022, from https://news.mongabay.com/2021/10/in-peruvian-andes-ancient-crops-hold-promise-for-a-climate-blighted-future/

Specialty Produce. (n.d.). Ulluco tubers. Specialty Produce. Retrieved May 20, 2022, from https://specialtyproduce.com/produce/Ulluco_Tubers_16986.php

Sperling, C.R. and S.R. King. 1990. Andean tuber crops: Worldwide potential. p. 428-435. In: J. Janick and J.E. Simon (eds.), Advances in new crops. Timber Press, Portland, OR.

Svenson, Johan et al. 2008. Betalains in Red and Yellow Varieties of the Andean Tuber Crop Ulluco (Ullucus tuberosus). 56, 17, 7730–7737. https://doi.org/10.1021/jf8012053