A revisited history of cacao domestication in pre-Columbian times revealed by archaeogenomic approaches.

Humans have a long history of transporting and trading plants, contributing to the evolution of domesticated plants. Theobroma cacao originated in the Neotropics from South America. However, little is known about its domestication and use in these regions. In this study, ceramic residues from a large sample of pre-Columbian cultures from South and Central America were analyzed using archaeogenomic and biochemical approaches. Here we show, for the first time, the widespread use of cacao in South America out of its native Amazonian area of origin, extending back 5000 years, likely supported by cultural interactions between the Amazon and the Pacific coast. We observed that strong genetic mixing between geographically distant cacao populations occurred as early as the middle Holocene, in South America, driven by humans, favoring the adaptation of T. cacao to new environments. This complex history of cacao domestication is the basis of today's cacao tree populations and its knowledge can help us better manage their genetic resources.

The use and domestication of Theobroma cacao during the mid-Holocene in the upper Amazon.

Cacao (Theobroma cacao L.) is an important economic crop, yet studies of its domestication history and early uses are limited. Traditionally, cacao is thought to have been first domesticated in Mesoamerica. However, genomic research shows that T. cacao's greatest diversity is in the upper Amazon region of northwest South America, pointing to this region as its centre of origin. Here, we report cacao use identified by three independent lines of archaeological evidence-cacao starch grains, absorbed theobromine residues and ancient DNA-dating from approximately 5,300 years ago recovered from the Santa Ana-La Florida (SALF) site in southeast Ecuador. To our knowledge, these findings constitute the earliest evidence of T. cacao use in the Americas and the first unequivocal archaeological example of its pre-Columbian use in South America. They also reveal the upper Amazon region as the oldest centre of cacao domestication yet identified.

Paleoindian settlement of the high-altitude Peruvian Andes.

Study of human adaptation to extreme environments is important for understanding our cultural and genetic capacity for survival. The Pucuncho Basin in the southern Peruvian Andes contains the highest-altitude Pleistocene archaeological sites yet identified in the world, about 900 meters above confidently dated contemporary sites. The Pucuncho workshop site [4355 meters above sea level (masl)] includes two fishtail projectile points, which date to about 12.8 to 11.5 thousand years ago (ka). Cuncaicha rock shelter (4480 masl) has a robust, well-preserved, and well-dated occupation sequence spanning the past 12.4 thousand years (ky), with 21 dates older than 11.5 ka. Our results demonstrate that despite cold temperatures and low-oxygen conditions, hunter-gatherers colonized extreme high-altitude Andean environments in the Terminal Pleistocene, within about 2 ky of the initial entry of humans to South America.

Directly dated starch residues document early formative maize (Zea mays L.) in tropical Ecuador.

The study of maize (Zea mays L.) domestication has advanced from questions of its origins to the study-and debate-of its dietary role and the timing of its dispersal from Mexico. Because the investigation of maize's spread is hampered by poor preservation of macrobotanical remains in the Neotropics, research has focused on microbotanical remains whose contexts are often dated by association, leading some to question the dates assigned. Furthermore, some scholars have argued that maize was not introduced to southwestern Ecuador until approximately 4150-3850 calendar years before the present (cal B.P.), that it was used first and foremost as a fermented beverage in ceremonial contexts, and that it was not important in everyday subsistence, challenging previous studies based on maize starch and phytoliths. To further investigate these questions, we analyzed every-day cooking vessels, food-processing implements, and sediments for starch and phytoliths from an archaeological site in southwestern Ecuador constituting a small Early Formative village. Employing a new technique to recover starch granules from charred cooking-pot residues we show that maize was present, cultivated, and consumed here in domestic contexts by at least 5300-4950 cal B.P. Directly dating the residues by accelerator mass spectrometry (AMS) radiocarbon measurement, our results represent the earliest direct dates for maize in Early Formative Ecuadorian sites and provide further support that, once domesticated approximately 9000 calendar years ago, maize spread rapidly from southwestern Mexico to northwestern South America.

Starch fossils and the domestication and dispersal of chili peppers (Capsicum spp. L.) in the Americas.

Chili peppers (Capsicum spp.) are widely cultivated food plants that arose in the Americas and are now incorporated into cuisines worldwide. Here, we report a genus-specific starch morphotype that provides a means to identify chili peppers from archaeological contexts and trace both their domestication and dispersal. These starch microfossils have been found at seven sites dating from 6000 years before present to European contact and ranging from the Bahamas to southern Peru. The starch grain assemblages demonstrate that maize and chilies occurred together as an ancient and widespread Neotropical plant food complex that predates pottery in some regions.