were green where the fruit were shaded and frequently produced excessive runners. Not until strawberries from Virginia became widely distributed did F. virginiana really make an impact on the horticultural industry. What became known as the ‘Scarlet strawberry’ was favoured for its large fruit size, high yields and deep red colour. They were particularly enjoyed in jam, because of their persistent colour, acid flavour, high aroma and retention of shape (Darrow, 1966).
It is not clear when the first Scarlet strawberries arrived in Europe, but in the garden of de La Brosse there was an item called F. americana magno fruto rubro, which may have been a large-fruited, scarlet type. The surviving colonists from Plymouth may also have brought back Scarlet strawberries when they returned in 1586. Regardless, during the late 1500s and 1600s, all kinds of native plants of North America appeared in the gardens of Europe, making multiple introductions of the Scarlet strawberry likely.
The early cultivar development of F. virginiana was primarily conducted by growers who found raising seed imported from North America often resulted in horticulturally important variations. The numbers of varieties available increased dramatically from three to about 26, over a period of a few decades at the turn of the 17th century (Darrow, 1966). Some of the most important early cultivars were ‘Oblong Scarlet’, ‘Grove End Scarlet’, ‘Duke of Kent’s Scarlet’, ‘Knight’s Large Scarlet’, ‘Wilmot’s Late Scarlet’, ‘Morrisana Scarlet’, ‘Common Scarlet’, the ‘Australian Scarlet’ and ‘Hudson Bay Scarlet’. They came from all over the New World including Nova Scotia, Virginia and New York. The early improvements were modest, however, and generally did not yield any substantial advancements from the best of the earlier imported types such as ‘Large Early Scarlet’.
Garden culture of strawberries began in North America in the middle of the 17th century with varieties imported from England. Early garden calendars listed three types of strawberries: (i) the hautboy (F. moschata); (ii) the Chili (Fragaria chiloensis, originally from Chile); and (iii) the Redwood (F. vesca from Europe) (Fletcher, 1917). Little cultivation of F. virginiana was undertaken until the importation of ‘Large Early Scarlet’ in the late 1700s, even though native strawberry populations abounded and ‘Large Early Scarlet’ had actually been sent to England a century earlier from the wilds of North America.
The first native American clone of F. virginiana to be propagated for sale in North America in 1791 was called ‘Hudson’. It was very vigorous and had soft, scarlet fruit with high flavour. It was cultivated well into the 1800s and can probably be considered the first important American strawberry (Hedrick, 1925). The first commercial strawberry plantings were established around Boston, New York, Philadelphia and Baltimore in the early 1800s. Until that time, most commercial strawberries had been gathered from the extensive wild populations that were springing up in conjunction with the clearing of forests and the abandonment of worn-out agricultural sites.
‘Red Wood’, an English variety of F. virginiana, was probably the most important variety grown in the early part of the 18th century in North America, along with a number of Scarlet varieties (Fletcher, 1917). ‘Red Wood’ was thought to be inferior in flavour to the older Scarlet varieties but had a longer season of production. Other popular cultivars during the first half of the 1800s were ‘White Wood’ (a white-fruited type), ‘Early Hudson’, ‘Old Scarlet’, ‘Crimson Cone’, ‘Large Early Scarlet’, ‘Hudson Bay Scarlet’, ‘Methven Scarlet’, ‘Ross Phoenix’ and ‘Early Virginia’. All of these except for ‘Ross Phoenix’ and ‘Hudson Bay Scarlet’ were imported from England.
DOMESTICATION OF F. CHILOENSIS IN THE NEW WORLD
The cultivated strawberry of South America, F. chiloensis, has a long and rich history (Hancock et al., 1999; Lavín and Maureira, 2000). It was utilized well over 1000 years ago by the indigenous Mapuches between the rivers Biobio and Tolten in south-central Chile, and by the more northern Picunches tribe between the rivers Itata and Biobio. The Picunches had contact with the northern agrarian Inca invaders and were probably the first to transport elite plants from the wild to their home gardens. The Mapuches were primarily hunters and gatherers but learned about agriculture from the Picunches.
The indigenous Chileans used strawberry fruits fresh, dried, as a fermented juice or as medicinal infusions against indigestion, diarrhoea and bleeding (de Moesbach, 1992). The Mapuches made many kinds of fermented juices, but the one from the llahuen or lahueñe small red-fruited wild strawberry was their favourite, called lahueñe mushca (Labarca, 1994).
Most evidence indicates that the primary domesticants were the larger white-fruited forms, called kallén or quellghen by the Mapuches. Albino-fruited types are rare in nature but have been found at three southern locations. Some red-fruited forms may also have been domesticated but reports of their existence are sketchy. Darrow (1957) described Chilean large red-fruited forms from around Santiago in the middle of the 20th century, although there are no earlier reports of cultivated lahueñe. Wild red-fruited forms were abundant from Santiago southward, so the pressure to cultivate them was probably minimal. It is reported, however, that the Mapuches planted small plots of the wild red forms in open spaces in the forests as a trap for the Spanish soldiers. When the soldiers dropped their arms to pick the fruit, the fierce Indians attacked and killed them (Gonzalez de Nájera, 1866).
Strawberry cultivation by the Mapuches was mostly limited to garden plots. After the conquest by the Spaniards, larger commercial plantings of 1–2 ha began to appear in the coastal areas from north of the Itata River to Chiloé Island. These traditional plantings of F. chiloensis flourished until the 1950s, when they began to be mixed with Northern Hemisphere cultivars of F. × ananassa. Cultivation of F. chiloensis has now largely disappeared, but small plantings can still be found along the traditional area of cultivation from Iloca on the coast of Curicó province to Chiloé Island (Hancock et al., 1999). Recently, there has been a burst of interest in redeveloping the commercial F. chiloensis industry in Chile, particularly around the white-fruited forms (Retamales et al., 2005; Céspedes, 2018; Morales-Quintana and Ramos, 2019).
SPREAD OF F. CHILOENSIS OUTSIDE CHILE
During their period of exploration and conquest in the mid- to late 1500s, the Spanish spread F. chiloensis throughout north-western South America. Major industries developed around Cuzco (Peru), Bogotá (Colombia) and Ambato (Ecuador) (Popenoe, 1921, 1926; Darrow, 1957). The source of these plants is not known; however, the variability
