Transoceanic drift and the domestication of African bottle gourds in the Americas

Bottle gourd (Lagenaria siceraria) was one of the first domesticated plants, and the only one with a global distribution during pre-Columbian times. Although native to Africa, bottle gourd was in use by humans in east Asia, possibly as early as 11,000 y ago (BP) and in the Americas by 10,000 BP. Despite its utilitarian importance to diverse human populations, it remains unresolved how the bottle gourd came to be so widely distributed, and in particular how and when it arrived in the New World. A previous study using ancient DNA concluded that Paleoindians transported already domesticated gourds to the Americas from Asia when colonizing the New World [Erickson et al. (2005) Proc Natl Acad Sci USA 102(51):18315–18320]. However, this scenario requires the propagation of tropical-adapted bottle gourds across the Arctic. Here, we isolate 86,000 base pairs of plastid DNA from a geographically broad sample of archaeological and living bottle gourds. In contrast to the earlier results, we find that all pre-Columbian bottle gourds are most closely related to African gourds, not Asian gourds. Ocean-current drift modeling shows that wild African gourds could have simply floated across the Atlantic during the Late Pleistocene. Once they arrived in the New World, naturalized gourd populations likely became established in the Neotropics via dispersal by megafaunal mammals. These wild populations were domesticated in several distinct New World locales, most likely near established centers of food crop domestication.In independent centers of plant domestication worldwide, distinct suites of food crops tend to emerge from native flora under human selection. An exception to this is the bottle gourd (Lagenaria siceraria, Cucurbitaceae), which is native to Africa, but was used by diverse human cultures not only in Africa, but also across Eurasia, the Pacific Islands, and the New World during pre-Columbian times (1–4). Although bottle gourd fruits are edible, they are used by humans mostly for other purposes, including as lightweight, durable containers, fishnet floats, and musical instruments (5). This variety of utilitarian applications likely explains why bottle gourds are so globally pervasive.In the New World, bottle gourds appear in archaeological contexts as early as 10,000 BP (6) (9). Bottle gourds were long proposed to have arrived in the Americas via long-range dispersal on ocean currents (10–13). However, an analysis of DNA from living and archaeological gourds suggested that the bottle gourd may have been transported into the New World by the first colonizing humans (6). In this scenario, the bottle gourd, like the dog (14), crossed the Bering Land Bridge with colonizing humans already in its domestic form, making the bottle gourd one of the earliest domesticated species (1, 6).

Table 1.

Archaeological samples and associated AMS radiocarbon dates for gourd rind fragments used in this study

Mitochondrial diversity and the origins of African and European cattle.

The nature of domestic cattle origins in Africa are unclear as archaeological data are relatively sparse. The earliest domesticates were humpless, or Bos taurus, in morphology and may have shared a common origin with the ancestors of European cattle in the Near East. Alternatively, local strains of the wild ox, the aurochs, may have been adopted by peoples in either continent either before or after cultural influence from the Levant. This study examines mitochondrial DNA displacement loop sequence variation in 90 extant bovines drawn from Africa, Europe, and India. Phylogeny estimation and analysis of molecular variance verify that sequences cluster significantly into continental groups. The Indian Bos indicus samples are most markedly distinct from the others, which is indicative of a B. taurus nature for both European and African ancestors. When a calibration of sequence divergence is performed using comparisons with bison sequences and an estimate of 1 Myr since the Bison/Bos Leptobos common ancestor, estimates of 117-275,000 B.P. and 22-26,000 B.P. are obtained for the separation between Indians and others and between African and European ancestors, respectively. As cattle domestication is thought to have occurred approximately 10,000 B.P., these estimates suggest the domestication of genetically discrete aurochsen strains as the origins of each continental population. Additionally, patterns of variation that are indicative of population expansions (probably associated with the domestication process) are discernible in Africa and Europe. Notably, the genetic signatures of these expansions are clearly younger than the corresponding signature of African/European divergence.     

Reassessing Coxcatlan Cave and the early history of domesticated plants in Mesoamerica

Reanalysis and direct accelerator mass spectrometry radiocarbon dating of the cucurbit assemblage from Coxcatlan Cave provide information on the timing and sequence of the initial appearance of three domesticated plants in the Tehuacán Valley (Puebla, Mexico) and allow reassessment of the overall temporal context of plant domestication in Mexico. Cucurbita pepo is the earliest documented domesticate in the cave, dating to 7,920 calibrated calendrical (cal) years B.P. The bottle gourd (Lagenaria siceraria) is dated at 7,200 cal years B.P. Cucurbita argyrosperma does not appear until 2,065 cal years B.P. The earlier identification of Cucurbita moschata specimens is not confirmed. Seventy-one radiocarbon dates, including 23 accelerator mass spectrometry dates on cucurbits, provide ample evidence of postdepositional vertical displacement of organic materials in the western half of Coxcatlan Cave, but they also indicate that the eastern half of the cave was largely undisturbed.     

Why did modern human populations disperse from Africa ca. 60,000 years ago? A new model

Recent research has provided increasing support for the origins of anatomically and genetically "modern" human populations in Africa between 150,000 and 200,000 years ago, followed by a major dispersal of these populations to both Asia and Europe sometime after ca. 65,000 before present (B.P.). However, the central question of why it took these populations approximately 100,000 years to disperse from Africa to other regions of the world has never been clearly resolved. It is suggested here that the answer may lie partly in the results of recent DNA studies of present-day African populations, combined with a spate of new archaeological discoveries in Africa. Studies of both the mitochondrial DNA (mtDNA) mismatch patterns in modern African populations and related mtDNA lineage-analysis patterns point to a major demographic expansion centered broadly within the time range from 80,000 to 60,000 B.P., probably deriving from a small geographical region of Africa. Recent archaeological discoveries in southern and eastern Africa suggest that, at approximately the same time, there was a major increase in the complexity of the technological, economic, social, and cognitive behavior of certain African groups, which could have led to a major demographic expansion of these groups in competition with other, adjacent groups. It is suggested that this complex of behavioral changes (possibly triggered by the rapid environmental changes around the transition from oxygen isotope stage 5 to stage 4) could have led not only to the expansion of the L2 and L3 mitochondrial lineages over the whole of Africa but also to the ensuing dispersal of these modern populations over most regions of Asia, Australasia, and Europe, and their replacement (with or without interbreeding) of the preceding "archaic" populations in these regions.     

From the Cover: Initial formation of an indigenous crop complex in eastern North America at 3800 B.P

Although geneticists and archaeologists continue to make progress world-wide in documenting the time and place of the initial domestication of a growing number of plants and animals, far less is known regarding the critically important context of coalescence of various species into distinctive sets or complexes of domesticates in each of the world's 10 or more independent centers of agricultural origin. In this article, the initial emergence of a crop complex is described for one of the best-documented of these independent centers, eastern North America (ENA). Before 4000 B.P. there is no indication of a crop complex in ENA, only isolated evidence for single indigenous domesticate species. By 3800 B.P., however, at least 5 domesticated seed-bearing plants formed a coherent complex in the river valley corridors of ENA. Accelerator mass spectrometer radiocarbon dates and reanalysis of archaeobotanical assemblages from a short occupation of the Riverton Site in Illinois documents the contemporary cultivation at 3800 B.P. of domesticated bottle gourd (Lagenaria siceraria), marshelder (Iva annua var. macrocarpa), sunflower (Helianthus annuus var. macrocarpus), and 2 cultivated varieties of chenopod (Chenopodium berlandieri), as well as the possible cultivation of Cucurbita pepo squash and little barley (Hordeum pusillum). Rather than marking either an abrupt developmental break or a necessary response to population-packing or compressed resource catchments, the coalescence of an initial crop complex in ENA appears to reflect an integrated expansion and enhancement of preexisting hunting and gathering economies that took place within a context of stable long-term adaptation to resource-rich river valley settings.     

Sunflower (Helianthus annuus L.) as a pre-Columbian domesticate in Mexico

Mexico has long been recognized as one of the world's cradles of domestication with evidence for squash (Cucurbita pepo) cultivation appearing as early as 8,000 cal B.C. followed by many other plants, such as maize (Zea mays), peppers (Capsicum annuum), common beans (Phaseolus vulgaris), and cotton (Gossypium hirsutum). We present archaeological, linguistic, ethnographic, and ethnohistoric data demonstrating that sunflower (Helianthus annuus) had entered the repertoire of Mexican domesticates by ca. 2600 cal B.C., that its cultivation was widespread in Mexico and extended as far south as El Salvador by the first millennium B.C., that it was well known to the Aztecs, and that it is still in use by traditional Mesoamerican cultures today. The sunflower's association with indigenous solar religion and warfare in Mexico may have led to its suppression after the Spanish Conquest. The discovery of ancient sunflower in Mexico refines our knowledge of domesticated Mesoamerican plants and adds complexity to our understanding of cultural evolution.     

Genetic and archaeological perspectives on the initial modern human colonization of southern Asia

It has been argued recently that the initial dispersal of anatomically modern humans from Africa to southern Asia occurred before the volcanic “supereruption” of the Mount Toba volcano (Sumatra) at ∼74,000 y before present (B.P.)—possibly as early as 120,000 y B.P. We show here that this “pre-Toba” dispersal model is in serious conflict with both the most recent genetic evidence from both Africa and Asia and the archaeological evidence from South Asian sites. We present an alternative model based on a combination of genetic analyses and recent archaeological evidence from South Asia and Africa. These data support a coastally oriented dispersal of modern humans from eastern Africa to southern Asia ∼60–50 thousand years ago (ka). This was associated with distinctively African microlithic and “backed-segment” technologies analogous to the African “Howiesons Poort” and related technologies, together with a range of distinctively “modern” cultural and symbolic features (highly shaped bone tools, personal ornaments, abstract artistic motifs, microblade technology, etc.), similar to those that accompanied the replacement of “archaic” Neanderthal by anatomically modern human populations in other regions of western Eurasia at a broadly similar date.     

The origin of European cattle: evidence from modern and ancient DNA

Cattle domestication from wild aurochsen was among the most important innovations during the Neolithic agricultural revolution. The available genetic and archaeological evidence points to at least two major sites of domestication in India and in the Near East, where zebu and the taurine breeds would have emerged independently. Under this hypothesis, all present-day European breeds would be descended from cattle domesticated in the Near East and subsequently spread during the diffusion of herding and farming lifestyles. We present here previously undescribed genetic evidence in contrast with this view, based on mtDNA sequences from five Italian aurochsen dated between 7,000 and 17,000 years B.P. and >1,000 modern cattle from 51 breeds. Our data are compatible with local domestication events in Europe and support at least some levels of introgression from the aurochs in Italy. The distribution of genetic variation in modern cattle suggest also that different south European breeds were affected by introductions from northern Africa. If so, the European cattle may represent a more variable and valuable genetic resource than previously realized, and previous simple hypotheses regarding the domestication process and the diffusion of selected breeds should be revised.     

Eastern North America as an independent center of plant domestication

The status of eastern North America as an independent center of plant domestication has recently been called into question by a number of genetic and archaeological studies, which suggest that the region may not have witnessed the independent domestication of local crop plants, but rather may have been on the receiving end of domesticated crop plants introduced from Mexico. Here, I provide a synthesis of the currently available archaeological and genetic evidence from both eastern North America and Mexico regarding the spatial and temporal context of initial domestication of the four plant species identified as potential eastern North American domesticates: marshelder (Iva annua), chenopod (Chenopodium berlandieri), squash (Cucurbita pepo), and sunflower (Helianthus annuus). Genetic and archaeological evidence provides strong support for the independent domestication of all four of these plant species in the eastern United States and reconfirms the region as one of the world's independent centers of domestication.     

Meager genetic variability of the human malaria agent Plasmodium vivax

Malaria is a major human parasitic disease caused by four species of Plasmodium protozoa. Plasmodium vivax, the most widespread, affects millions of people across Africa, Asia, the Middle East, and Central and South America. We have studied the genetic variability of 13 microsatellite loci in 108 samples from 8 localities in Asia, Africa, South America, and New Guinea. Only one locus is polymorphic; nine are completely monomorphic, and the remaining three are monomorphic in all but one or two populations, which have a rare second allele. In contrast, Plasmodium falciparum displays extensive microsatellite polymorphism within and among populations. We further have analyzed, in 96 samples from the same 8 localities, 8 tandem repeats (TRs) located on a 100-kb contiguous chromosome segment described as highly polymorphic. Each locus exhibits 2-10 alleles in the whole sample but little intrapopulation polymorphism (1-5 alleles with a prevailing allele in most cases). Eight microsatellite loci monomorphic in P. vivax are polymorphic in three of five Plasmodium species related to P. vivax (two to seven individuals sampled). Plasmodium simium, a parasite of New World monkeys, is genetically indistinguishable from P. vivax. At 13 microsatellite loci and at 7 of the 8 TRs, both species share the same (or most common) allele. Scarce microsatellite polymorphism may reflect selective sweeps or population bottlenecks in recent evolutionary history of P. vivax; the differential variability of the TRs may reflect selective processes acting on particular regions of the genome. We infer that the world expansion of P. vivax as a human parasite occurred recently, perhaps <10,000 years ago.     

Ancient mitochondrial DNA analysis reveals complexity of indigenous North American turkey domestication

Although the cultural and nutritive importance of the turkey (Meleagris gallopavo) to precontact Native Americans and contemporary people worldwide is clear, little is known about the domestication of this bird compared to other domesticates. Mitochondrial DNA analysis of 149 turkey bones and 29 coprolites from 38 archaeological sites (200 BC–AD 1800) reveals a unique domesticated breed in the precontact Southwestern United States. Phylogeographic analyses indicate that this domestic breed originated from outside the region, but rules out the South Mexican domestic turkey (Meleagris gallopavo gallopavo) as a progenitor. A strong genetic bottleneck within the Southwest turkeys also reflects intensive human selection and breeding. This study points to at least two occurrences of turkey domestication in precontact North America and illuminates the intensity and sophistication of New World animal breeding practices.     

Microfossil evidence for pre-Columbian maize dispersals in the neotropics from San Andres, Tabasco, Mexico

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.     

Domestication of the donkey: timing, processes, and indicators

Domestication of the donkey from the African wild ass transformed ancient transport systems in Africa and Asia and the organization of early cities and pastoral societies. Genetic research suggests an African origin for the donkey, but pinpointing the timing and location of domestication has been challenging because donkeys are uncommon in the archaeological record and markers for early phases of animal domestication are hard to determine. We present previously undescribed evidence for the earliest transport use of the donkey and new paleopathological indicators for early phases of donkey domestication. Findings are based on skeletal data from 10 approximately 5,000-year-old ass skeletons recently discovered entombed in an early pharaonic mortuary complex at Abydos, Middle Egypt, and a concurrent study of 53 modern donkey and African wild ass skeletons. Morphometric studies showed that Abydos metacarpals were similar in overall proportions to those of wild ass, but individual measurements varied. Midshaft breadths resembled wild ass, but midshaft depths and distal breadths were intermediate between wild ass and domestic donkey. Despite this, all of the Abydos skeletons exhibited a range of osteopathologies consistent with load carrying. Morphological similarities to wild ass show that, despite their use as beasts of burden, donkeys were still undergoing considerable phenotypic change during the early Dynastic period in Egypt. This pattern is consistent with recent studies of other domestic animals that suggest that the process of domestication is slower and less linear than previously thought.     

Evidence for two independent domestications of cattle.

The origin and taxonomic status of domesticated cattle are controversial. Zebu and taurine breeds are differentiated primarily by the presence or absence of a hump and have been recognized as separate species (Bos indicus and Bos taurus). However, the most widely held view is that both types of cattle derive from a single domestication event 8000-10,000 years ago. We have examined mtDNA sequences from representatives of six European (taurine) breeds, three Indian (zebu) breeds, and four African (three zebu, one taurine) breeds. Similar levels of average sequence divergence were observed among animals within each of the major continental groups: 0.41% (European), 0.38% (African), and 0.42% (Indian). However, the sequences fell into two very distinct geographic lineages that do not correspond with the taurine-zebu dichotomy: all European and African breeds are in one lineage, and all Indian breeds are in the other. There was little indication of breed clustering within either lineage. Application of a molecular clock suggests that the two major mtDNA clades diverged at least 200,000, and possibly as much as 1 million, years ago. This relatively large divergence is interpreted most simply as evidence for two separate domestication events, presumably of different subspecies of the aurochs, Bos primigenius. The clustering of all African zebu mtDNA sequences within the taurine lineage is attributed to ancestral crossbreeding with the earlier B. taurus inhabitants of the continent.     

Molecular evidence for a single evolutionary origin of domesticated rice

Asian rice, Oryza sativa, is one of world's oldest and most important crop species. Rice is believed to have been domesticated ～9,000 y ago, although debate on its origin remains contentious. A single-origin model suggests that two main subspecies of Asian rice, indica and japonica, were domesticated from the wild rice O. rufipogon. In contrast, the multiple independent domestication model proposes that these two major rice types were domesticated separately and in different parts of the species range of wild rice. This latter view has gained much support from the observation of strong genetic differentiation between indica and japonica as well as several phylogenetic studies of rice domestication. We reexamine the evolutionary history of domesticated rice by resequencing 630 gene fragments on chromosomes 8, 10, and 12 from a diverse set of wild and domesticated rice accessions. Using patterns of SNPs, we identify 20 putative selective sweeps on these chromosomes in cultivated rice. Demographic modeling based on these SNP data and a diffusion-based approach provide the strongest support for a single domestication origin of rice. Bayesian phylogenetic analyses implementing the multispecies coalescent and using previously published phylogenetic sequence datasets also point to a single origin of Asian domesticated rice. Finally, we date the origin of domestication at ～8,200-13,500 y ago, depending on the molecular clock estimate that is used, which is consistent with known archaeological data that suggests rice was first cultivated at around this time in the Yangtze Valley of China.     

Global archaeological evidence for proboscidean overkill

One million years ago, proboscideans occupied most of Africa, Europe, Asia, and the Americas. Today, wild elephants are only found in portions of sub-Saharan Africa and South Asia. Although the causes of global Pleistocene extinctions in the order Proboscidea remain unresolved, the most common explanations involve climatic change and/or human hunting. In this report, we test the overkill and climate-change hypotheses by using global archaeological spatiotemporal patterning in proboscidean kill/scavenge sites. Spanning approximately 1.8 million years, the archaeological record of human subsistence exploitation of proboscideans is preferentially located on the edges of the human geographic range. This finding is commensurate with global overkill, suggesting that prehistoric human range expansion resulted in localized extinction events. In the present and the past, proboscideans have survived in refugia that are largely inaccessible to human populations.     

Mitochondrial DNA "clock" for the Amerinds and its implications for timing their entry into North America.

Students of the time of entry of the ancestors of the Amerinds into the New World are divided into two camps, one favoring an "early" entry [more than approximately 30,000 years before the present (YBP)], the other favoring a "late" entry (less than approximately 13,000 YBP). An "intermediate" date is unlikely for geological reasons. The correlation of the appropriate data on mtDNA variation in Amerinds with linguistic, archaeological, and genetic data offers the possibility of establishing a time frame for mtDNA evolution in Amerinds. In this paper, we estimate that the separation of the Chibcha-speaking tribes of Central America from other linguistic groups/nascent tribes began approximately 8000-10,000 YBP. Characterization of the mtDNA of 110 Chibcha speakers with 14 restriction enzymes leads on the basis of their time depth to an estimated mtDNA nucleotide substitution rate for Amerinds of 0.022-0.029% per 10,000 years. As a first application of this rate, we consider the mtDNA variation observed in 18 Amerind tribes widely dispersed throughout the Americas and studied by ourselves with the same techniques, and we estimate that if the Amerinds entered the New World as a single group, that entry occurred approximately 22,000-29,000 YBP. This estimate carries a large but indeterminate error. The mtDNA data are thus at present equivocal with respect to the most likely times of entry of the Amerind into the New World mentioned above but favor the "early" entry hypothesis.     

The goat domestication process inferred from large-scale mitochondrial DNA analysis of wild and domestic individuals

The emergence of farming during the Neolithic transition, including the domestication of livestock, was a critical point in the evolution of human kind. The goat (Capra hircus) was one of the first domesticated ungulates. In this study, we compared the genetic diversity of domestic goats to that of the modern representatives of their wild ancestor, the bezoar, by analyzing 473 samples collected over the whole distribution range of the latter species. This partly confirms and significantly clarifies the goat domestication scenario already proposed by archaeological evidence. All of the mitochondrial DNA haplogroups found in current domestic goats have also been found in the bezoar. The geographic distribution of these haplogroups in the wild ancestor allowed the localization of the main domestication centers. We found no haplotype that could have been domesticated in the eastern half of the Iranian Plateau, nor further to the east. A signature of population expansion in bezoars of the C haplogroup suggests an early domestication center on the Central Iranian Plateau (Yazd and Kerman Provinces) and in the Southern Zagros (Fars Province), possibly corresponding to the management of wild flocks. However, the contribution of this center to the current domestic goat population is rather low (1.4%). We also found a second domestication center covering a large area in Eastern Anatolia, and possibly in Northern and Central Zagros. This last domestication center is the likely origin of almost all domestic goats today. This finding is consistent with archaeological data identifying Eastern Anatolia as an important domestication center.     

Dog domestication and the dual dispersal of people and dogs into the Americas

Advances in the isolation and sequencing of ancient DNA have begun to reveal the population histories of both people and dogs. Over the last 10,000 y, the genetic signatures of ancient dog remains have been linked with known human dispersals in regions such as the Arctic and the remote Pacific. It is suspected, however, that this relationship has a much deeper antiquity, and that the tandem movement of people and dogs may have begun soon after the domestication of the dog from a gray wolf ancestor in the late Pleistocene. Here, by comparing population genetic results of humans and dogs from Siberia, Beringia, and North America, we show that there is a close correlation in the movement and divergences of their respective lineages. This evidence places constraints on when and where dog domestication took place. Most significantly, it suggests that dogs were domesticated in Siberia by ∼23,000 y ago, possibly while both people and wolves were isolated during the harsh climate of the Last Glacial Maximum. Dogs then accompanied the first people into the Americas and traveled with them as humans rapidly dispersed into the continent beginning ∼15,000 y ago.     

Chronic hepatitis B: current epidemiology in the Americas and implications for management

Hepatitis B virus (HBV) remains a serious health threat in many parts of the world. Although its prevalence is lower in the Americas than in Asia, Africa and the Middle East, it is responsible for significant morbidity and mortality in North, Central and South America. There is a nonuniform pattern of distribution throughout this region, with HBV prevalence related to geographical, social and cultural factors that predispose certain individuals to infection. This report details the incidence, modes of viral transmission of hepatitis B in the Americas and clinical course of disease in different regions of the Americas. Additionally, the implications for management focusing on issues predominant in high-risk populations are presented.