Lidar reveals pre-Hispanic low-density urbanism in the Bolivian Amazon.

Archaeological remains of agrarian-based, low-density urbananism1-3 have been reported to exist beneath the tropical forests of Southeast Asia, Sri Lanka and Central America4-6. However, beyond some large interconnected settlements in southern Amazonia7-9, there has been no such evidence for pre-Hispanic Amazonia. Here we present lidar data of sites belonging to the Casarabe culture (around AD 500 to AD 1400)10-13 in the Llanos de Mojos savannah-forest mosaic, southwest Amazonia, revealing the presence of two remarkably large sites (147 ha and 315 ha) in a dense four-tiered settlement system. The Casarabe culture area, as far as known today, spans approximately 4,500 km2, with one of the large settlement sites controlling an area of approximately 500 km2. The civic-ceremonial architecture of these large settlement sites includes stepped platforms, on top of which lie U-shaped structures, rectangular platform mounds and conical pyramids (which are up to 22 m tall). The large settlement sites are surrounded by ranked concentric polygonal banks and represent central nodes that are connected to lower-ranked sites by straight, raised causeways that stretch over several kilometres. Massive water-management infrastructure, composed of canals and reservoirs, complete the settlement system in an anthropogenically modified landscape. Our results indicate that the Casarabe-culture settlement pattern represents a type of tropical low-density urbanism that has not previously been described in Amazonia.

Early Holocene crop cultivation and landscape modification in Amazonia.

The onset of plant cultivation is one of the most important cultural transitions in human history1-4. Southwestern Amazonia has previously been proposed as an early centre of plant domestication, on the basis of molecular markers that show genetic similarities between domesticated plants and wild relatives4-6. However, the nature of the early human occupation of southwestern Amazonia, and the history of plant cultivation in this region, are poorly understood. Here we document the cultivation of squash (Cucurbita sp.) at about 10,250 calibrated years before present (cal. yr BP), manioc (Manihot sp.) at about 10,350 cal. yr BP and maize (Zea mays) at about 6,850 cal. yr BP, in the Llanos de Moxos (Bolivia). We show that, starting at around 10,850 cal. yr BP, inhabitants of this region began to create a landscape that ultimately comprised approximately 4,700 artificial forest islands within a treeless, seasonally flooded savannah. Our results confirm that the Llanos de Moxos is a hotspot for early plant cultivation and demonstrate that-ever since their arrival in Amazonia-humans have markedly altered the landscape, with lasting repercussions for habitat heterogeneity and species conservation.

Impact of pre-Columbian "geoglyph" builders on Amazonian forests.

Over 450 pre-Columbian (pre-AD 1492) geometric ditched enclosures ("geoglyphs") occupy ∼13,000 km2 of Acre state, Brazil, representing a key discovery of Amazonian archaeology. These huge earthworks were concealed for centuries under terra firme (upland interfluvial) rainforest, directly challenging the "pristine" status of this ecosystem and its perceived vulnerability to human impacts. We reconstruct the environmental context of geoglyph construction and the nature, extent, and legacy of associated human impacts. We show that bamboo forest dominated the region for ≥6,000 y and that only small, temporary clearings were made to build the geoglyphs; however, construction occurred within anthropogenic forest that had been actively managed for millennia. In the absence of widespread deforestation, exploitation of forest products shaped a largely forested landscape that survived intact until the late 20th century.

Environmental impact of geometric earthwork construction in pre-Columbian Amazonia.

There is considerable controversy over whether pre-Columbian (pre-A.D. 1492) Amazonia was largely "pristine" and sparsely populated by slash-and-burn agriculturists, or instead a densely populated, domesticated landscape, heavily altered by extensive deforestation and anthropogenic burning. The discovery of hundreds of large geometric earthworks beneath intact rainforest across southern Amazonia challenges its status as a pristine landscape, and has been assumed to indicate extensive pre-Columbian deforestation by large populations. We tested these assumptions using coupled local- and regional-scale paleoecological records to reconstruct land use on an earthwork site in northeast Bolivia within the context of regional, climate-driven biome changes. This approach revealed evidence for an alternative scenario of Amazonian land use, which did not necessitate labor-intensive rainforest clearance for earthwork construction. Instead, we show that the inhabitants exploited a naturally open savanna landscape that they maintained around their settlement despite the climatically driven rainforest expansion that began ∼2,000 y ago across the region. Earthwork construction and agriculture on terra firme landscapes currently occupied by the seasonal rainforests of southern Amazonia may therefore not have necessitated large-scale deforestation using stone tools. This finding implies far less labor--and potentially lower population density--than previously supposed. Our findings demonstrate that current debates over the magnitude and nature of pre-Columbian Amazonian land use, and its impact on global biogeochemical cycling, are potentially flawed because they do not consider this land use in the context of climate-driven forest-savanna biome shifts through the mid-to-late Holocene.

Preceramic maize from Paredones and Huaca Prieta, Peru.

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.

Fire-free land use in pre-1492 Amazonian savannas.

The nature and scale of pre-Columbian land use and the consequences of the 1492 "Columbian Encounter" (CE) on Amazonia are among the more debated topics in New World archaeology and paleoecology. However, pre-Columbian human impact in Amazonian savannas remains poorly understood. Most paleoecological studies have been conducted in neotropical forest contexts. Of studies done in Amazonian savannas, none has the temporal resolution needed to detect changes induced by either climate or humans before and after A.D. 1492, and only a few closely integrate paleoecological and archaeological data. We report a high-resolution 2,150-y paleoecological record from a French Guianan coastal savanna that forces reconsideration of how pre-Columbian savanna peoples practiced raised-field agriculture and how the CE impacted these societies and environments. Our combined pollen, phytolith, and charcoal analyses reveal unexpectedly low levels of biomass burning associated with pre-A.D. 1492 savanna raised-field agriculture and a sharp increase in fires following the arrival of Europeans. We show that pre-Columbian raised-field farmers limited burning to improve agricultural production, contrasting with extensive use of fire in pre-Columbian tropical forest and Central American savanna environments, as well as in present-day savannas. The charcoal record indicates that extensive fires in the seasonally flooded savannas of French Guiana are a post-Columbian phenomenon, postdating the collapse of indigenous populations. The discovery that pre-Columbian farmers practiced fire-free savanna management calls into question the widely held assumption that pre-Columbian Amazonian farmers pervasively used fire to manage and alter ecosystems and offers fresh perspectives on an emerging alternative approach to savanna land use and conservation that can help reduce carbon emissions.

Adaptation strategies of giant viruses to low-temperature marine ecosystems.

Microbes in marine ecosystems have evolved their gene content to thrive successfully in the cold. Although this process has been reasonably well studied in bacteria and selected eukaryotes, less is known about the impact of cold environments on the genomes of viruses that infect eukaryotes. Here, we analyzed cold adaptations in giant viruses (Nucleocytoviricota and Mirusviricota) from austral marine environments and compared them with their Arctic and temperate counterparts. We recovered giant virus metagenome-assembled genomes (98 Nucleocytoviricota and 12 Mirusviricota MAGs) from 61 newly sequenced metagenomes and metaviromes from sub-Antarctic Patagonian fjords and Antarctic seawater samples. When analyzing our data set alongside Antarctic and Arctic giant viruses MAGs already deposited in the Global Ocean Eukaryotic Viral database, we found that Antarctic and Arctic giant viruses predominantly inhabit sub-10°C environments, featuring a high proportion of unique phylotypes in each ecosystem. In contrast, giant viruses in Patagonian fjords were subject to broader temperature ranges and showed a lower degree of endemicity. However, despite differences in their distribution, giant viruses inhabiting low-temperature marine ecosystems evolved genomic cold-adaptation strategies that led to changes in genetic functions and amino acid frequencies that ultimately affect both gene content and protein structure. Such changes seem to be absent in their mesophilic counterparts. The uniqueness of these cold-adapted marine giant viruses may now be threatened by climate change, leading to a potential reduction in their biodiversity.

Hydrographic shifts in coastal waters reflect climate-driven changes in hydrological regimes across Northwestern Patagonia.

Climate-driven changes in freshwater inputs have been shown to affect the structure and function of coastal ecosystems. We evaluated changes in the influence of river runoff on coastal systems of Northwestern Patagonia (NWP) over recent decades (1993-2021) by combined analysis of long-term streamflow time series, hydrological simulation, satellite-derived and reanalysis data on sea surface conditions (temperature, turbidity, and salinity). Significant decreases in minimum streamflow across a zone spanning six major river basins were evident at weekly, monthly, and seasonal scales. These changes have been most pronounced in mixed-regime northern basins (e.g., Puelo River) but appear to be progressing southward to rivers characterised by a nival regime. In the adjacent two-layer inner sea, reduced freshwater input corresponds with a shallower halocline and increased surface temperatures across northern Patagonia. Our results underscore the rapidly evolving influence of rivers on adjacent estuarine and coastal waters in NWP. We highlight the need for cross-ecosystem observation, forecasting, mitigation and adaptation strategies in a changing climate, together with corresponding adaptive basin management of systems that supply runoff to the coastal marine waters.

Pre-Columbian agricultural landscapes, ecosystem engineers, and self-organized patchiness in Amazonia.

The scale and nature of pre-Columbian human impacts in Amazonia are currently hotly debated. Whereas pre-Columbian people dramatically changed the distribution and abundance of species and habitats in some parts of Amazonia, their impact in other parts is less clear. Pioneer research asked whether their effects reached even further, changing how ecosystems function, but few in-depth studies have examined mechanisms underpinning the resilience of these modifications. Combining archeology, archeobotany, paleoecology, soil science, ecology, and aerial imagery, we show that pre-Columbian farmers of the Guianas coast constructed large raised-field complexes, growing on them crops including maize, manioc, and squash. Farmers created physical and biogeochemical heterogeneity in flat, marshy environments by constructing raised fields. When these fields were later abandoned, the mosaic of well-drained islands in the flooded matrix set in motion self-organizing processes driven by ecosystem engineers (ants, termites, earthworms, and woody plants) that occur preferentially on abandoned raised fields. Today, feedbacks generated by these ecosystem engineers maintain the human-initiated concentration of resources in these structures. Engineer organisms transport materials to abandoned raised fields and modify the structure and composition of their soils, reducing erodibility. The profound alteration of ecosystem functioning in these landscapes coconstructed by humans and nature has important implications for understanding Amazonian history and biodiversity. Furthermore, these landscapes show how sustainability of food-production systems can be enhanced by engineering into them follows that maintain ecosystem services and biodiversity. Like anthropogenic dark earths in forested Amazonia, these self-organizing ecosystems illustrate the ecological complexity of the legacy of pre-Columbian land use.

The combined impact of land use change and aquaculture on sediment and water quality in oligotrophic Lake Rupanco (North Patagonia, Chile, 40.8°S).

Water and sediment quality in North Patagonia's large, oligotrophic lakes are expected to suffer as native forest continues to be fragmented and degraded by its conversion to cropping and pasture land uses. These changes in land use are expected to increase diffuse nutrient loads to the region's lakes. In addition, these lakes are home to the world's second largest salmon aquaculture industry which provides additional point sources of nutrients within the lakes. We studied the combined influences of land use change and salmon farming on the nutrient concentrations in a North Patagonian lake (Lake Rupanco, 233 km(2) water surface, 163 m average depth) in four sub-watersheds ranging in disturbance from near-pristine forest to 53% converted to cropping and pasture. Nitrogen exports from the tributary sub-watersheds increased from 33 kg TN/km(2)/y to 621 kg TN/km(2)/y as the proportion of crop and pasture land increased. The combined nutrient load from land use change and salmon farming has led to significant differences in the nitrogen concentrations of the lake's water column and sediments in the near-shore zones across the lake. Total nitrogen concentrations in the sediments varied from 37 ± 18 mg/kg in near-pristine sub-watersheds without salmon farming to 6400 ± 698 mg/kg where the sub-watershed was dominated by crop and pasture lands combined with the presence of salmon farming. These results demonstrate the importance of considering the impacts of both salmon farming and land use on water and sediment quality for future environmental planning, management and decision making.

Assessing the micro-phytoplankton response to nitrate in Comau Fjord (42°S) in Patagonia (Chile), using a microcosms approach.

Anthropogenic (aquaculture) changes in environment nutrient concentrations may affect phytoplankton (biomass and taxa composition) in marine coastal waters off the Chilean Patagonia. The effects of adding nitrate (NO3(-)) to natural phytoplankton assemblages were evaluated considering biomass, cell abundance, and taxonomic composition. Microcosm experiments were performed in the spring, summer, and winter in the Comau Fjord located in Subantarctic Patagonia. At the end of the experiments, NO3(-) decreased rapidly and was undetectable in treatments, indicating a strong NO3(-) deficiency associated with an exponential increase in Chl-a concentrations, particulate organic nitrogen, and carbon in these treatments. Moreover, given the depleted nitrate concentrations of the spring and summer experiments, the micro-phytoplankton taxa structure shifted from mixed diatom and dinoflagellate assemblages (Ceratium spp., Dinophysis spp., Coscinodiscus sp., Rhizosolenia pungens) to assemblages dominated by blooms of the classic chain-forming diatoms found in temperate and cold waters such as Chaetoceros spp., Skeletonema spp., and Thalassiosira spp. Thus, nitrogen sources (i.e., nitrate, ammonia) may influence phytoplankton abundance and biomass accumulation dynamics in the northern section of Patagonia. It also emphasizes the importance of diatom taxa in regards to the short-term response of phytoplankton to changing environmental nutrient conditions due to natural (decreasing freshwater stream flow) and anthropogenic (aquaculture) events. This situation may be one of the future scenarios in the Patagonian fjords, thus stressing the needs for active environmental monitoring and impact assessment.

Corrigendum: Biogeography of southern ocean active prokaryotic communities over a large spatial scale.

[This corrects the article DOI: 10.3389/fmicb.2022.862812.].

More than 10,000 pre-Columbian earthworks are still hidden throughout Amazonia.

Indigenous societies are known to have occupied the Amazon basin for more than 12,000 years, but the scale of their influence on Amazonian forests remains uncertain. We report the discovery, using LIDAR (light detection and ranging) information from across the basin, of 24 previously undetected pre-Columbian earthworks beneath the forest canopy. Modeled distribution and abundance of large-scale archaeological sites across Amazonia suggest that between 10,272 and 23,648 sites remain to be discovered and that most will be found in the southwest. We also identified 53 domesticated tree species significantly associated with earthwork occurrence probability, likely suggesting past management practices. Closed-canopy forests across Amazonia are likely to contain thousands of undiscovered archaeological sites around which pre-Columbian societies actively modified forests, a discovery that opens opportunities for better understanding the magnitude of ancient human influence on Amazonia and its current state.

Starch grain and phytolith evidence for early ninth millennium B.P. maize from the Central Balsas River Valley, Mexico.

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.

The cultural and chronological context of early Holocene maize and squash domestication in the Central Balsas River Valley, Mexico.

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.

Ice Age megafauna rock art in the Colombian Amazon?

Megafauna paintings have accompanied the earliest archaeological contexts across the continents, revealing a fundamental inter-relationship between early humans and megafauna during the global human expansion as unfamiliar landscapes were humanized and identities built into new territories. However, the identification of extinct megafauna from rock art is controversial. Here, we examine potential megafauna depictions in the rock art of Serranía de la Lindosa, Colombian Amazon, that includes a giant sloth, a gomphothere, a camelid, horses and three-toed ungulates with trunks. We argue that they are Ice Age rock art based on the (i) naturalistic appearance and diagnostic morphological features of the animal images, (ii) late Pleistocene archaeological dates from La Lindosa confirming the contemporaneity of humans and megafauna, (iii) recovery of ochre pigments in late Pleistocene archaeological strata, (iv) the presence of most megafauna identified in the region during the late Pleistocene as attested by archaeological and palaeontological records, and (v) widespread depiction of extinct megafauna in rock art across the Americas. Our findings contribute to the emerging picture of considerable geographical and stylistic variation of geometric and figurative rock art from early human occupations across South America. Lastly, we discuss the implications of our findings for understanding the early human history of tropical South America. This article is part of the theme issue 'Tropical forests in the deep human past'.

Biogeography of Southern Ocean Active Prokaryotic Communities Over a Large Spatial Scale.

The activity of marine microorganisms depends on community composition, yet, in some oceans, less is known about the environmental and ecological processes that structure their distribution. The objective of this study was to test the effect of geographical distance and environmental parameters on prokaryotic community structure in the Southern Ocean (SO). We described the total (16S rRNA gene) and the active fraction (16S rRNA-based) of surface microbial communities over a ~6,500 km longitudinal transect in the SO. We found that the community composition of the total fraction was different from the active fraction across the zones investigated. In addition, higher α-diversity and stronger species turnover were displayed in the active community compared to the total community. Oceanospirillales, Alteromonadales, Rhodobacterales, and Flavobacteriales dominated the composition of the bacterioplankton communities; however, there were marked differences at the order level. Temperature, salinity, silicic acid, particulate organic nitrogen, and particulate organic carbon correlated with the composition of bacterioplankton communities. A strong distance-decay pattern between closer and distant communities was observed. We hypothesize that it was related to the different oceanic fronts present in the Antarctic Circumpolar Current. Our findings contribute to a better understanding of the complex arrangement that shapes the structure of bacterioplankton communities in the SO.

Legacies of Indigenous land use and cultural burning in the Bolivian Amazon rainforest ecotone.

The southwestern Amazon Rainforest Ecotone (ARE) is the transitional landscape between the tropical forest and seasonally flooded savannahs of the Bolivian Llanos de Moxos. These heterogeneous landscapes harbour high levels of biodiversity and some of the earliest records of human occupation and plant domestication in Amazonia. While persistent Indigenous legacies have been demonstrated elsewhere in the Amazon, it is unclear how past human-environment interactions may have shaped vegetation composition and structure in the ARE. Here, we examine 6000 years of archaeological and palaeoecological data from Laguna Versalles (LV), Bolivia. LV was dominated by stable rainforest vegetation throughout the Holocene. Maize cultivation and cultural burning are present after ca 5700 cal yr BP. Polyculture cultivation of maize, manioc and leren after ca 3400 cal yr BP predates the formation of Amazonian Dark/Brown Earth (ADE/ABE) soils (approx. 2400 cal yr BP). ADE/ABE formation is associated with agroforestry indicated by increased edible palms, including Mauritia flexuosa and Attalea sp., and record levels of burning, suggesting that fire played an important role in agroforestry practices. The frequent use of fire altered ADE/ABD forest composition and structure by controlling ignitions, decreasing fuel loads and increasing the abundance of plants preferred by humans. Cultural burning and polyculture agroforestry provided a stable subsistence strategy that persisted despite pronounced climate change and cultural transformations and has an enduring legacy in ADE/ABE forests in the ARE. This article is part of the theme issue 'Tropical forests in the deep human past'.