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Disturbances in tropical forests can have long-lasting ecological impacts, but their manifestations (ecological legacies) in modern forests are uncertain. Many Amazonian forests bear the mark of past soil modifications, species enrichments, and fire events, but the trajectories of ecological legacies from the pre-contact or post-colonial period remain relatively unexplored. We assessed the fire and vegetation history from 15 soil cores ranging from 0 to 10 km from a post-colonial Surinamese archaeological site. We show that (1) fires occurred from 96 bc to recent times and induced significant vegetation change, (2) persistent ecological legacies from pre-contact and post-colonial fire and deforestation practices were mainly within 1 km of the archaeological site, and (3) palm enrichment of Attalea, Oenocarpus and Astrocaryum occurred within 0, 1, and 8 km of the archaeological site, respectively. Our results challenge the notion of spatially extensive and persistent ecological legacies. Instead, our data indicate that the persistence and extent of ecological legacies are dependent on their timing, frequency, type, and intensity. Examining the mechanisms and manifestations of ecological legacies is crucial in assessing forest resilience and Indigenous and local land rights in the highly threatened Amazonian forests.

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This paper addresses an important debate in Amazonian studies; namely, the scale, intensity, and nature of human modification of the forests in prehistory. Phytolith and charcoal analysis of terrestrial soils underneath mature tierra firme (nonflooded, nonriverine) forests in the remote Medio Putumayo-Algodón watersheds, northeastern Peru, provide a vegetation and fire history spanning at least the past 5,000 y. A tree inventory carried out in the region enables calibration of ancient phytolith records with standing vegetation and estimates of palm species densities on the landscape through time. Phytolith records show no evidence for forest clearing or agriculture with major annual seed and root crops. Frequencies of important economic palms such as Oenocarpus, Euterpe, Bactris, and Astrocaryum spp., some of which contain hyperdominant species in the modern flora, do not increase through prehistoric time. This indicates pre-Columbian occupations, if documented in the region with future research, did not significantly increase the abundance of those species through management or cultivation. Phytoliths from other arboreal and woody species similarly reflect a stable forest structure and diversity throughout the records. Charcoal 14C dates evidence local forest burning between ca. 2,800 and 1,400 y ago. Our data support previous research indicating that considerable areas of some Amazonian tierra firme forests were not significantly impacted by human activities during the prehistoric era. Rather, it appears that over the last 5,000 y, indigenous populations in this region coexisted with, and helped maintain, large expanses of relatively unmodified forest, as they continue to do today.

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Maize (Zea mays ssp. mays) is among the world's most important and ancient domesticated crops. Although the chronology of its domestication and initial dispersals out of Mexico into Central and South America has become more clear due to molecular and multiproxy archaeobotanical research, important problems remain. Among them is the paucity of information on maize's early morphological evolution and racial diversification brought about in part by the poor preservation of macrofossils dating to the pre-5000 calibrated years before the present period from obligate dispersal routes located in the tropical forest. We report newly discovered macrobotanical and microbotanical remains of maize that shed significant light on the chronology, land race evolution, and cultural contexts associated with the crop's early movements into South America and adaptation to new environments. The evidence comes from the coastal Peruvian sites of Paredones and Huaca Prieta, Peru; dates from the middle and late preceramic and early ceramic periods (between ca. 6700 and 3000 calibrated years before the present); and constitutes some of the earliest known cobs, husks, stalks, and tassels. The macrobotanical record indicates that a diversity of racial complexes characteristic of the Andean region emerged during the preceramic era. In addition, accelerator mass spectrometry radiocarbon determinations carried out directly on different structures of preserved maize plants strongly suggest that assays on burned cobs are more reliable than those on unburned cobs. Our findings contribute to knowledge of the early diffusion of maize and agriculture and have broader implications for understanding the development of early preindustrial human societies.

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Questions that still surround the origin and early dispersals of maize (Zea mays L.) result in large part from the absence of information on its early history from the Balsas River Valley of tropical southwestern Mexico, where its wild ancestor is native. We report starch grain and phytolith data from the Xihuatoxtla shelter, located in the Central Balsas Valley, that indicate that maize was present by 8,700 calendrical years ago (cal. B.P.). Phytolith data also indicate an early preceramic presence of a domesticated species of squash, possibly Cucurbita argyrosperma. The starch and phytolith data also allow an evaluation of current hypotheses about how early maize was used, and provide evidence as to the tempo and timing of human selection pressure on 2 major domestication genes in Zea and Cucurbita. Our data confirm an early Holocene chronology for maize domestication that has been previously indicated by archaeological and paleoecological phytolith, starch grain, and pollen data from south of Mexico, and reshift the focus back to an origin in the seasonal tropical forest rather than in the semiarid highlands.

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Molecular evidence indicates that the wild ancestor of maize is presently native to the seasonally dry tropical forest of the Central Balsas watershed in southwestern Mexico. We report here on archaeological investigations in a region of the Central Balsas located near the Iguala Valley in Guerrero state that show for the first time a long sequence of human occupation and plant exploitation reaching back to the early Holocene. One of the sites excavated, the Xihuatoxtla Shelter, contains well-stratified deposits and a stone tool assemblage of bifacially flaked points, simple flake tools, and numerous handstones and milling stone bases radiocarbon dated to at least 8700 calendrical years B.P. As reported in a companion paper (Piperno DR, et al., in this issue of PNAS), starch grain and phytolith residues from the ground and chipped stone tools, plus phytoliths from directly associated sediments, provide evidence for maize (Zea mays L.) and domesticated squash (Cucurbita spp.) in contexts contemporaneous with and stratigraphically below the 8700 calendrical years B.P. date. The radiocarbon determinations, stratigraphic integrity of Xihuatoxtla's deposits, and characteristics of the stone tool assemblages associated with the maize and squash remains all indicate that these plants were early Holocene domesticates. Early agriculture in this region of Mexico appears to have involved small groups of cultivators who were shifting their settlements seasonally and engaging in a variety of subsistence pursuits.

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Previous research indicates that the Nanchoc Valley in northern Peru was an important locus of early and middle Holocene human settlement, and that between 9200 and 5500 (14)C yr B.P. the valley inhabitants adopted major crop plants such as squash (Cucurbita moschata), peanuts (Arachis sp.), and cotton (Gossypium barbadense). We report here an examination of starch grains preserved in the calculus of human teeth from these sites that provides direct evidence for the early consumption of cultivated squash and peanuts along with two other major food plants not previously detected. Starch from the seeds of Phaseolus and Inga feuillei, the flesh of Cucurbita moschata fruits, and the nuts of Arachis was routinely present on numerous teeth that date to between 8210 and 6970 (14)C yr B.P. Early plant diets appear to have been diverse and stable through time and were rich in cultivated foods typical of later Andean agriculture. Our data provide early archaeological evidence for Phaseolus beans and I. feuillei, an important tree crop, and indicate that effective food production systems that contributed significant dietary inputs were present in the Nanchoc region by 8000 (14)C yr B.P. Starch grain studies of dental remains document plants and edible parts of them not normally preserved in archaeological records and can assume primary roles as direct indicators of ancient human diets and agriculture.

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We examined pollen grains and starch granules from a large number of modern populations of teosinte (wild Zea spp.), maize (Zea mays L.), and closely related grasses in the genus Tripsacum to assess their strengths and weaknesses in studying the origins and early dispersals of maize in its Mesoamerican cradle of origin. We report new diagnostic criteria and question the accuracy of others used previously by investigators to identify ancient maize where its wild ancestor, teosinte, is native. Pollen grains from teosinte overlap in size with those of maize to a much greater degree than previously reported, making the differentiation of wild and domesticated maize in palynological studies difficult. There is presently no valid method for separating maize and teosinte pollen on a morphological basis. Starch grain analysis, a recently developed tool of archaeobotany, appears to be of significant utility in distinguishing the seeds of teosinte from maize. We propose that the differences in starch grain morphology and size between wild and domesticated maize defined in this study may be associated with domestication genes in Zea that have been documented in the starch biosynthesis pathway. As previously reported, phytoliths effectively discriminate the female reproductive structures of Tripsacum, teosinte, and maize. Multiproxy microfossil studies of archaeological and paleoecological contexts appear to be effective tools for investigating the earliest stages of maize domestication and dispersals.

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The history of maize (Zea mays L.) is one of the most debated topics in New World archaeology. Molecular and genetic studies indicate that maize domestication took place in tropical southwest Mexico. Although archaeological evidence for the evolution of maize from its wild ancestor teosinte has yet to be found in that poorly studied region, other research combining paleoecology and archaeology is documenting the nature and timing of maize domestication and dispersals. Here we report a phytolith analysis of sediments from San Andrés, Tabasco, that confirms the spread of maize cultivation to the tropical Mexican Gulf Coast >7,000 years ago ( approximately 7,300 calendar years before present). We review the different methods used in sampling, identifying, and dating fossil maize remains and compare their strengths and weaknesses. Finally, we examine how San Andrés amplifies the present evidence for widespread maize dispersals into Central and South America. Multiple data sets from many sites indicate that maize was brought under cultivation and domesticated and had spread rapidly out of its domestication cradle in tropical southwest Mexico by the eighth millennium before the present.

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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.

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Over the past decade, increasing attention to the recovery and identification of plant microfossil remains from archaeological sites located in lowland South America has significantly increased knowledge of pre-Columbian plant domestication and crop plant dispersals in tropical forests and other regions. Along the Andean mountain chain, however, the chronology and trajectory of plant domestication are still poorly understood for both important indigenous staple crops such as the potato (Solanum sp.) and others exogenous to the region, for example, maize (Zea mays). Here we report the analyses of plant microremains from a late preceramic house (3,431 +/- 45 to 3,745 +/- 65 14C bp or approximately 3,600 to 4,000 calibrated years bp) in the highland southern Peruvian site of Waynuna. Our results extend the record of maize by at least a millennium in the southern Andes, show on-site processing of maize into flour, provide direct evidence for the deliberate movement of plant foods by humans from the tropical forest to the highlands, and confirm the potential of plant microfossil analysis in understanding ancient plant use and migration in this region.

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Barley (Hordeum vulgare L.) and wheat (Triticum monococcum L. and Triticum turgidum L.) were among the principal 'founder crops' of southwest Asian agriculture. Two issues that were central to the cultural transition from foraging to food production are poorly understood. They are the dates at which human groups began to routinely exploit wild varieties of wheat and barley, and when foragers first utilized technologies to pound and grind the hard, fibrous seeds of these and other plants to turn them into easily digestible foodstuffs. Here we report the earliest direct evidence for human processing of grass seeds, including barley and possibly wheat, in the form of starch grains recovered from a ground stone artefact from the Upper Palaeolithic site of Ohalo II in Israel. Associated evidence for an oven-like hearth was also found at this site, suggesting that dough made from grain flour was baked. Our data indicate that routine processing of a selected group of wild cereals, combined with effective methods of cooking ground seeds, were practiced at least 12,000 years before their domestication in southwest Asia.

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Cucurbita (squash and gourd) phytoliths recovered from two early Holocene archaeological sites in southwestern Ecuador and directly dated to 10,130 to 9320 carbon-14 years before the present (about 12,000 to 10,000 calendar years ago) are identified as derived from domesticated plants because they are considerably larger than those from modern wild taxa. The beginnings of plant husbandry appear to have been preceded by the exploitation of a wild species of Cucurbita during the terminal Pleistocene. These data provide evidence for an independent emergence of plant food production in lowland South America that was contemporaneous with or slightly before that in highland Mesoamerica.

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Many angiosperms, both monocotyledons and dicotyledons, heavily impregnate their vegetative and reproductive organs with solid particles of silicon dioxide (SiO(2)) known as opaline phytoliths. The underlying mechanisms accounting for the formation of phytoliths in plants are poorly understood, however. Using wild and domesticated species in the genus Cucurbita along with their F(1) and F(2) progeny, we have demonstrated that the production of large diagnostic phytoliths in fruit rinds exhibits a one-to-one correspondence to the lignification of these structures. We propose that phytolith formation in Cucurbita fruits is primarily determined by a dominant genetic locus, called hard rind (Hr), previously shown to code for lignin deposition. If true, this evidence represents a demonstration of genetic control over phytolith production in a dicotyledon and provides considerable support to hypotheses that silica phytoliths constitute another important system of mechanical defense in plants. Our research also identifies Hr as another single locus controlling more than one important phenotypic difference between wild and domesticated plants, and establishes rind tissue cell structure and hardness under the effects of Hr as an important determinant of phytolith morphology. When recovered from pre-Columbian archaeological sites, Cucurbita phytoliths represent genetically controlled fossil markers of exploitation and domestication in this important economic genus.

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We have investigated the phylogenetic relationships among six wild and six domesticated taxa of Cucurbita using as a marker an intron region from the mitochondrial nad1 gene. Our study represents one of the first successful uses of a mtDNA gene in resolving inter- and intraspecific taxonomic relationships in Angiosperms and yields several important insights into the origins of domesticated Cucurbita. First, our data suggest at least six independent domestication events from distinct wild ancestors. Second, Cucurbita argyrosperma likely was domesticated from a wild Mexican gourd, Cucurbita sororia, probably in the same region of southwest Mexico that gave rise to maize. Third, the wild ancestor of Cucurbita moschata is still unknown, but mtDNA data combined with other sources of information suggest that it will probably be found in lowland northern South America. Fourth, Cucurbita andreana is supported as the wild progenitor of Cucurbita maxima, but humid lowland regions of Bolivia in addition to warmer temperate zones in South America from where C. andreana was originally described should possibly be considered as an area of origin for C. maxima. Fifth, our data support other molecular results that indicate two separate domestications in the Cucurbita pepo complex. The potential zone of domestication for one of the domesticated subspecies, C. pepo subsp. ovifera, includes eastern North America and should be extended to northeastern Mexico. The wild ancestor of the other domesticated subspecies, C. pepo subsp. pepo, is undiscovered but is closely related to C. pepo subsp. fraterna and possibly will be found in southern Mexico.

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