Early transatlantic movement of horses and donkeys at Jamestown

William Timothy Treal Taylor; Nicolas Delsol; Vicky M Oelze; Peter Mitchell; Leah Stricker; Michael Lavin; Akin Ogundiran; Lauren Hosek; Christina Isabelle Barrón-Ortiz; Olumide Ojediran; Diana Quintero-Bisono; Lucy Keith-Diagne; Dane Magoon; Matthew E Hill, Jr; Ariane E Thomas; Anna Waterman; David W Peate; Lorelei Chauvey; Stéphanie Schiavinato; Laure Calvière-Tonasso; Luís Borges; Aitor Brito-Mayor; Jonathan Santana; George Kamenov; Ludovic Orlando; John Krigbaum

doi:10.1126/sciadv.adw2595

. 2025 Sep 3;11(36):eadw2595. doi: 10.1126/sciadv.adw2595

Early transatlantic movement of horses and donkeys at Jamestown

William Timothy Treal Taylor ^1,^2,^*, Nicolas Delsol ^3,⁴, Vicky M Oelze ⁵, Peter Mitchell ^6,⁷, Leah Stricker ⁸, Michael Lavin ⁸, Akin Ogundiran ⁹, Lauren Hosek ¹, Christina Isabelle Barrón-Ortiz ¹⁰, Olumide Ojediran ¹, Diana Quintero-Bisono ², Lucy Keith-Diagne ¹¹, Dane Magoon ¹², Matthew E Hill Jr ¹³, Ariane E Thomas ¹³, Anna Waterman ¹⁴, David W Peate ¹⁵, Lorelei Chauvey ¹⁶, Stéphanie Schiavinato ¹⁶, Laure Calvière-Tonasso ¹⁶, Luís Borges ¹⁷, Aitor Brito-Mayor ¹⁸, Jonathan Santana ¹⁸, George Kamenov ¹⁹, Ludovic Orlando ¹⁶, John Krigbaum ⁴

^¹Department of Anthropology, University of Colorado–Boulder, 218 UCB, Boulder, CO 80309, USA.

^²Museum of Natural History, University of Colorado–Boulder, 218 UCB, Boulder, CO 80309, USA.

^³Université Laval, Laboratoire d’Archéologie, 300 rue Saint-Paul, Québec, QC G1K 7R1 Canada.

^⁴Department of Anthropology, University of Florida, Gainesville, FL 32611, USA.

^⁵Anthropology Department, University of California Santa Cruz, Santa Cruz, CA 95060, USA.

^⁶University of Oxford, School of Archaeology, 1 S Parks Rd, Oxford OX1 3TG, United Kingdom.

^⁷University of the Witwatersrand, Rock Art Research Institute, 1 Jan Smuts Ave, Braamfontein, Johannesburg 2017, South Africa.

^⁸Jamestown Rediscovery, Preservation Virginia, 1365 Colonial Parkway, Jamestown, VA 23081, USA.

^⁹Department of History, Northwestern University, 1881 Sheridan Rd, Evanston, IL 60208, USA.

^¹⁰Royal Alberta Museum, Quaternary Studies Program, 9810 103a Avenue NW, Edmonton, AB T5J 0G2 Canada.

^¹¹African Aquatic Conservation Fund, BP Joal, 23015, Senegal, West Africa.

^¹²University of Leicester, Archaeology and Ancient History, University Road, Leicester LE1 7RH, United Kingdom.

^¹³Department of Anthropology, University of Iowa, 114 Macbride Hall, Iowa City, IA 52242, USA.

^¹⁴Department of Natural and Applied Sciences, Mount Mercy University, Cedar Rapids, IA 52402, USA.

^¹⁵Department of Earth and Environmental Sciences, University of Iowa, 115 Trowbridge Hall, Iowa City, IA 52242, USA.

^¹⁶Université Paul Sabatier, Centre d’Anthropobiologie et de Génomique de Toulouse (CNRS UMR 5288), Toulouse 31000, France.

^¹⁷Direção Regional dos Assuntos Culturais dos Açores, Rua da Conceição, Palacete Silveira e Paulo, 9700-054 Angra do Heroísmo, Portugal.

^¹⁸University of Las Palmas de Gran Canaria, Department of Historical Sciences, Pérez del Toro 1, Las Palmas de G.C. E35003, Spain.

^¹⁹Department of Geological Sciences, University of Florida, Gainesville, FL 32611, USA.

Corresponding author. Email: william.taylor@colorado.edu

Roles

William Timothy Treal Taylor: Conceptualization, Data curation, Formal analysis, Funding acquisition, Investigation, Methodology, Project administration, Software, Supervision, Validation, Visualization, Writing - original draft, Writing - review & editing

Nicolas Delsol: Conceptualization, Investigation, Methodology, Writing - original draft, Writing - review & editing

Vicky M Oelze: Formal analysis, Resources, Visualization, Writing - review & editing

Peter Mitchell: Conceptualization, Data curation, Resources, Validation, Writing - review & editing

Leah Stricker: Conceptualization, Data curation, Resources, Validation, Writing - review & editing

Michael Lavin: Resources

Akin Ogundiran: Conceptualization, Resources, Writing - review & editing

Lauren Hosek: Formal analysis, Writing - review & editing

Christina Isabelle Barrón-Ortiz: Formal analysis, Investigation, Methodology, Validation, Visualization, Writing - original draft, Writing - review & editing

Olumide Ojediran: Conceptualization, Data curation, Formal analysis, Investigation, Methodology, Validation, Visualization, Writing - original draft, Writing - review & editing

Diana Quintero-Bisono: Writing - review & editing

Lucy Keith-Diagne: Data curation, Investigation, Resources, Validation, Writing - review & editing

Dane Magoon: Resources

Matthew E Hill Jr: Investigation, Writing - review & editing

Ariane E Thomas: Investigation, Writing - review & editing

Anna Waterman: Resources, Writing - review & editing

David W Peate: Resources, Writing - review & editing

Lorelei Chauvey: Investigation, Resources

Stéphanie Schiavinato: Resources

Laure Calvière-Tonasso: Resources

Luís Borges: Investigation, Resources, Validation, Writing - review & editing

Aitor Brito-Mayor: Investigation, Resources, Validation, Writing - review & editing

Jonathan Santana: Writing - review & editing

George Kamenov: Data curation, Formal analysis, Investigation, Methodology, Resources, Supervision, Validation, Visualization, Writing - original draft, Writing - review & editing

Ludovic Orlando: Conceptualization, Data curation, Formal analysis, Funding acquisition, Methodology, Project administration, Resources, Software, Supervision, Validation, Visualization, Writing - original draft, Writing - review & editing

John Krigbaum: Conceptualization, Data curation, Formal analysis, Funding acquisition, Investigation, Methodology, Resources, Supervision, Validation, Visualization, Writing - original draft, Writing - review & editing

PMCID: PMC12407088 PMID: 40901968

Abstract

Domestic equids were central to the initial colonization of the Atlantic coast of the Americas, a process partially chronicled by historical records. While Spanish colonists brought horses to the Caribbean decades earlier, settlement of the English colony at Jamestown, Virginia, was among the first dispersals of horses to the eastern seaboard. Archaeozoological analysis of identifiable domestic equid remains from two contexts associated with the initial occupation of Jamestown demonstrates intense processing and consumption of the first Jamestown horses during the “Starving Time” winter of 1609. Osteological and biomolecular study of these equid remains demonstrates their successful reproduction at the colony and use in transport activities and identifies an adult domestic donkey with mixed European and West African ancestry, possibly supplied through undocumented exchange during a transatlantic stopover. These results reveal the challenges of equid translocation in early settlement of eastern North America and the global connectivity of early transatlantic animal exchange.

Animal bones from Jamestown show that donkeys joined the colony's first Atlantic voyage and that starving colonists survived by eating their horses.

INTRODUCTION

The introduction of domestic horses into North America was among the most consequential events in the history of the Americas, with horses both transforming the lifeways of many Indigenous people as well as forming a core part of colonial infrastructure for European colonists (1). Although the first horses reached areas of western and southeastern North America through the activity of Spanish explorers and settlers, many of the first domestic equids to reach the eastern seaboard of the United States were those that accompanied English settlers. During later centuries, the expansion of these British colonial horses would come to shape the genetic landscape of both colonial and Indigenous horse populations across the continent (2). Jamestown, the first permanent English settlement in North America, represents an important snapshot in the early chapters of the human-equid story of the Americas.

British horses, donkeys, and early North America

Before the Industrial Revolution, horses provided the infrastructure for nearly all aspects of colonial life, from basic transport, communication, and trade to agriculture and warfare (3). In late Tudor and early Stuart England, changing land use and environment paired with rising demand for horses created a deficit of breeding animals (4), prompting new restrictions on equid export and trade (5, 6). Despite these complications, the ultimate success and expansion of British colonial horse populations in what is now the eastern United States shifted the genomic landscape of the early Americas (2). Today, most horses in the United States belong to breeds with some ancestral link to early British and American horses from the eastern seaboard. Donkeys, too, although often omitted from discussions around the impact of translocated domestic animals, had a tremendously important role in the cultural and agricultural enterprises of the early Americas (7). Today, both animals exist in large numbers as wild animals across the American West, where their ecological role and decisions around their management catalyze political and conservation debates.

Jamestown and the earliest British equids

The colony of Jamestown, located near the confluence of the James River and the Chesapeake Bay in the present-day state of Virginia, was initially settled by English colonists aboard a fleet of three ships that departed from London in 1606 (Fig. 1). First-person accounts document this voyage, which stopped in the Canary Islands, crossed the Atlantic to the Caribbean, and made stopovers throughout (Martinique, Dominica, Nevis, St. Croix, Puerto Rico, and the Mona and Monito Islands) before arriving at their final destination (8, 9) in 1607. Supply voyages in 1608 and 1609 likely followed a similar route. Documentary evidence of a third supply voyage suggests that “sixe Mares and two horses” were taken aboard in Plymouth, England before setting sail (10), and records imply that one male horse perished during the sea journey (8). Seven of the nine ships of the third supply reached Jamestown, while one ship was lost and another was stranded overwinter in Bermuda. Relations with the Indigenous Tsenacomocoan people living near the colony soured by the winter of 1609, resulting in a siege of the palisaded James Fort. This winter, known as the “Starving Time,” introduced extraordinary subsistence difficulties, and most of the colonists starved to death or perished from disease. The colonists were forced to consume animals not otherwise eaten, including horses, before eating dogs, rats, and even humans (11). In the spring of 1610, the stranded Bermuda colonists arrived in Jamestown, soon followed by a fourth supply expedition carrying Governor Lord De La Warr, who instituted a cleansing and rebuilding effort of James Fort.

Fig. 1. — (A) Study sites and localities mentioned in the Introduction, along with the route of the first colonial voyage to Jamestown. Map by Bill Nelson. (B) Excavation of the First Well at Jamestown, JR2718/Structure 185. Photo credit: Jamestown Rediscovery (Preservation Virginia). (C) Excavation of the Kitchen Cellar, JR3081/Structure 191. Photo credit: Jamestown Rediscovery (Preservation Virginia). (D) Site map with features highlighted. Analyzed structures are highlighted in red, excavated areas in yellow, identified structures in blue, and other archaeological features in orange. Location of the original 1607–1608 fort palisades are outlined in goldenrod. (E) Skeletal elements identified and present in the analyzed assemblage in red (left), and estimated age of horse specimens in the assemblage that could be aged using dentition (right). (F) Bayesian phase model on radiocarbon dates from select horse elements to infer horse activity in Structures 185 and 191 associated with the Starving Time at Jamestown. y, years.

Archaeological animal remains from two features linked to the Starving Time provide a unique opportunity to understand the role of domestic equids in early British settlement of the eastern seaboard of North America. The first was a barrel-lined well (“First Well”) located in the geographic center of the fort, constructed in late 1608 to early 1609 on the orders of John Smith (8). Designated JR2718/Structure 185, artifacts and stratigraphic relationships within the well feature suggest that the well was filled during spring 1610 “cleansing,” with successive filling episodes occurring in later years (12). Preliminary faunal analysis of material recovered from the well feature identified fish, crab, turtle, wild bird, and wild small mammals in addition to horse and dog remains (11). Just to the east of the First Well is a second, likely contemporaneous feature, excavated by Jamestown Rediscovery archaeologists in 2012. Designated JR3081/Structure 191, the feature was an L-shaped subterranean cellar with two brick-facade ovens. Likely the colony’s first kitchen described in early records as constructed in early 1608, archaeological evidence suggests that like the First Well, this feature was abandoned during the Starving Time winter and was filled with trash in the spring of 1610 (12). Prior faunal analysis of the assemblage from these two locations revealed a total of 77 specimens provisionally identified as “horse.” To assess the role of domestic equids during the earliest occupation at Jamestown, we conducted interdisciplinary archaeological, osteological, and biomolecular analysis (radiocarbon dating, isotopes, and DNA) of equid bones linked to Starving Time contexts from Jamestown.

RESULTS

Radiocarbon dating

Direct radiocarbon dating of five equid specimens produced three successful radiocarbon dates (Fig. 1), including two from adult domestic horses (catalog no. 117294 and no. 121460) and one from a specimen identified as an adult domestic donkey (no. 121161). After calibration, each of these specimens produced a nearly identical radiocarbon measurement consistent with an animal that died during the 16th or early 17th century (table S1). We assume that the deposition of the horses must have postdated the founding of the colony in 1607 but that the mineralization of the horse’s teeth (the event measured by ¹⁴C results) preceded both this date and the feature fill by several years (likely less than 10). We model this scenario using a modified version of the Charcoal Outlier model in OxCal, using a scaling parameter of zero. Results provide good measures of model agreement (A_model = 98.6, A_overall = 95.4). The feature fill of these two equid features is modeled with a start boundary of 1603–1615 (68%) or 1603–1618 (95%) and a median modeled date of 1610, while the end boundary is modeled at approximately 1609–1624 (68%) or 1603–1647 (95%), with a median modeled date of 1618. These results provide support for the association of these two features with the Starving Time winter of 1609–1610.

Elements present

Skeletal elements present in the assemblage include specimens from both appendicular and axial portions of the skeleton, including teeth, vertebrae, limbs, and pelvis. Given the highly processed nature of the identified elements and the lower caloric value of many of them (such as the bones of the lower limbs), it is highly likely that elements of the entire skeleton were originally represented among the full assemblage but that larger limb bones, ribs, and vertebrae have been processed so intensively as to render them difficult to identify as seen in other comparable assemblages (13). Alternatively, it may be that these lower-value elements were disposed of in the two depositional contexts analyzed here, the cellar and the well, while high-value components were curated and/or discarded in a different manner.

Species identification

For each element, species previously identified as Equus sp. in the collections at Jamestown Rediscovery were assessed against a comparative panel of three-dimensionally (3D) digitized skeletal elements of horse and donkey (reference collections from the Archaeozoology Laboratory at the University of Colorado). We excluded those that could not be positively confirmed as Equus from further analysis. Of these confirmed as Equus, 62 specimens were consistent with an identification as domestic horse (Equus caballus) or mule (E. caballus × Equus asinus), while one small upper cheek tooth (upper M2) was identified as domestic donkey (E. asinus).

The occlusal surface of the tooth (no. 121161) identified as domestic donkey was sufficiently well preserved for further assessment (Fig. 2A). This tooth shows dental features that are typically found in E. asinus. In particular, the distal length of the protocone is approximately equal to the mesial length of the protocone (i.e., the protocone is symmetrical), the pli caballin is absent, and the postprotoconal groove is deep. These features are found at a higher frequency in domestic donkeys than in domestic horses (section S2). The five samples analyzed for their DNA were identified as four horse mares and one donkey jack, which confirmed the morphological assessment (section S5).

Demographics

Age estimation of the identified horse specimens shows a profile of mostly so-called “prime age” adults between the ages of 6 and 10 years, with a minimum number of individuals (MNI) of at least two adult animals. This finding, along with the DNA-based determination of all analyzed horse teeth as female, is consistent with the assumption that the analyzed assemblage was derived from the horses known from first-person accounts, consisting of a total of eight animals, six females and two males. The estimated age of the specimens, with a central tendency of ~8 years, would indicate that the animals were young adults near maturity (~4 years) when they left England. We identified no canine teeth or other direct evidence of sex of the analyzed specimens but did identify the remains of a single juvenile specimen, a fractured phalanx of a young horse less than 1 year old. Although age determination using techniques like crown height is not reliable on the basis of available data for donkeys (14), the donkey remains also provide an MNI of at least one adult donkey of greater than ~2 years of age, based on its status as erupted and in wear with fully formed roots (15).

Pathology

Pathological features identified on the horse remains suggest that these animals were used in transport during their lifetime. Two second premolar specimens from the assemblage show exposure of the anterior enamel, a damage pattern often found in horses that have been ridden with a bit (16). Although anterior damage of the lower second premolars is not uniquely indicative of bridle use (17), in one case, the enamel is also fractured vertically in a manner consistent with traumatic contact from the anterior direction (Fig. 2D), consistent with the use of a bridle mouthpiece. The exposed cross-sectional profile of one horse metatarsal shows asymmetry that could be linked to transport or work-related activities (Fig. 2E) but may also reflect other aspects of activity patterns, such as gait (18, 19).

Several fractures in the assemblage are likely associated with kick injuries. In one case, a left metacarpal bone shows a healed fracture of the splint bone (Fig. 2F), which is commonly caused by kick injury from other animals. Similarly, the phalanx of the single foal identified exhibits a healed comminuted or crush fracture that reflects a high-energy injury (20). These patterns could reflect animals being held together in close confinement. One fragment of a mandible also showed an oval lytic lesion on the inferior lateral surface at the gonial angle; this feature likely represents a cyst with reactive bone formation along the floor of the cavity.

Taphonomy

Horse and donkey remains from Jamestown exhibit a high degree of cultural modification (Figs. 3 and 4), with all but 11 (82%) of the analyzed specimens exhibiting some clear form of modification [spiral fractures (n = 6), cut marks (n = 13), chop marks (n = 20), burning (n = 7), midshaft discoidal fractures (n = 2), and cortical bone removal or pulp extraction (n = 22)]. Most notable is the degree to which all marrow-bearing elements have been processed to open the marrow cavity and stripped of internal trabecular bone (Fig. 4A), even those typically considered to be low-yield (21). Many of the equid teeth in the assemblage have been split either in a longitudinal or transverse direction, presumably to expose the pulp cavity (Fig. 3). Limb bones and teeth in the assemblage commonly exhibit large or repeated impact fractures on lateral surfaces (Figs. 3 and 4). Under low-power microscopy, some tooth fragments even displayed embedded pieces of iron apparently linked to ax use (Fig. 3B). One equid tooth exhibited severe rounding that may relate to taphonomic impact of water within the well or other cultural or fluvial processes (Fig. 4D).

Fig. 3. — (A) A subset of split mandibular and maxillary cheek teeth in the Jamestown assemblage, including lateral impact fractures of tooth enamel and spiral fracturing. (B) Microscopy inset showing iron fragments embedded in some chopped specimens.

Fig. 4. — (A) Right radius with chop marks, impact fractures, and trabecular bone removal. (B and C) Phalanges with evidence of chop marks and splitting to access interior marrow cavity. (D) Single equid tooth showing unusual rounding.

DNA analysis

DNA preservation levels were compatible with the genome-wide characterization of the donkey jack specimen JT02/no. 121161 (Jamestown, hereafter) (section S5 and figs. S3 to S6). Neighbor-joining phylogenetic reconstructions supported the placement of the Jamestown specimen within a 100% bootstrap-supported monophyletic group including all the previously sequenced modern and ancient donkey specimens from Europe, the Canary Islands, and Brazil present in our comparative panel (Fig. 5C). Maximum likelihood phylogenetic reconstructions in TreeMix confirmed general genetic affinities of European origins in both the Chupaderos [17th to 19th century Mexico (2)] and Jamestown specimens, regardless of whether groups of modern accessions were stratified by country of origin or subcontinental region (Fig. 5D). This finding was consistent with the results of a principal components analysis (PCA), in which ancient European specimens, including those from Chupaderos and Jamestown, projected onto present-day European variation and toward positive values along the first component, which is characteristic of modern European populations (figs. S4 and S5). In this analysis, the placement of the Chupaderos donkey was closer to the modern populations from mainland Spain (ESP), the Canary Islands (CYK), and Brazil (BRA), relative to that of the Jamestown specimen. The latter was closer to the ancient European donkey from Western Europe, including medieval (Albufeira, Portugal: approximately 1228–1280 calibrated CE) and postmedieval (Fiumarella, Italy: approximately 1683–1936 CE), as well as from Classical Antiquity (Boinville, France: approximately 200–500 CE; Bourse, France: approximately 0–500 CE).

Fig. 5. — (A) The f₃-outgroup (f3O) statistics between a subset of ancient and modern donkeys. The group of *E. kiang* genome sequences (KIANG) present in the genome panel served as the outgroup. The f₃-outgroup statistics provides a measure for pairwise population genetic affinities. Modern donkey accessions were stratified according to the subcontinental groups of genetic affinities defined by Todd *et al.* (22), with Spanish accession subdivided to gain further resolution into possible genetic affinities in the region. (B) Phylogenetic relationships between the ancient donkey specimen from Jamestown (JT02/no. 121161) and a worldwide panel of modern donkeys, grouped by subcontinental groups of genetic affinities. The tree was rooted on the sequence data available for the *E. kiang* species, and node supports were assessed from 100 bootstrap pseudoreplicates. The tree models population genetic affinities, considering an optimal number of migration edges (Supplementary Materials). (C) Phylogenetic affinities between 223 donkey specimens distributed across the world. Node supports were estimated from 100 bootstrap pseudoreplicates and are provided in percentage. The labels of modern donkey accessions report the cluster of genetic affinities defined by Todd *et al.* (22), followed by the sample name. The labels of ancient donkey genomes indicate the sample name, followed by their country of origin (three-letter code) and their estimated radiocarbon age and/or the timeline of their associated archaeological context, except for the Jamestown JT02/no. 121161 and Chupaderos specimen from the 18th to 20th century. (D) f₄ statistics of the form f₄ [*Equus kiang*, population (Pop); Jamestown, Chupaderos]. This statistics measures whether the ancient donkey specimens from Jamestown and Chupaderos are genetically equally related to modern donkey accessions, grouped by country of origin or stratified by the subcontinental groups of genetic affinities defined by Todd *et al.* (22). Statistically significant positive (negative) values indicate extragenetic affinities between population and Chupaderos (Jamestown).

The historical specimen from Chupaderos, Mexico, placed within a monophyletic group of modern donkeys from the Canary Islands, while the specimen from Jamestown did not group with any modern or ancient donkey specimen. This was consistent in both neighbor-joining and TreeMix phylogenetic reconstructions and with f₃-outgroup statistics, which measure pairwise genetic affinities between those samples and a comparative panel of other ancient donkeys from Europe, and worldwide modern donkey groups stratified by country or subcontinent. The latter analyses revealed the different genomic makeup of both specimens, with Jamestown appearing closer to modern donkeys from the Balkan (YUM, North Macedonia) and Chupaderos also showing strong genetic affinities with Brazilian Pega donkeys and modern CYK donkeys. The presence of extragenetic affinities with Pega donkeys and the Canary Islands in the Chupaderos specimen, relative to Jamestown, was confirmed by statistically significant f₄ statistics of the form f₄(Equus kiang, X; Jamestown, Chupaderos, where X represents any modern donkey accession, grouped by country of subcontinental groups; fig. S5). This analysis also revealed an excess of genetic sharedness between modern accessions from West Africa (Mauritania), relative to Chupaderos. The presence of this extragenetic affinity was no longer supported when grouping all modern accessions from West Africa together (Mauritania as well as Senegal and Nigeria), suggesting specific contributions from Mauritania or closely related populations. This finding echoes the work from Todd et al. (22), which identified genetic contributions between Western Europe and West Africa in the Roman and Medieval time periods, which were replicated in our own TreeMix phylogenetic reconstructions.

Last, the genome sequenced from the Jamestown donkey was only compatible with one-way admixture modeling considering the medieval specimen from Albufeira, Portugal, or the Chupaderos, Mexico, specimen as possible sources (fig. S5). Multiple two-way qpAdm models confirmed that the Jamestown genome consisted mainly of Portuguese (PTG, Portugal), Mediterranean (TUN, Tunisia; YUMYUC, North Macedonia and Croatia), the Levant (SYR, Syria), or Anatolia (TUK, Turkey) genetic ancestry and ~8.4 to 13.4% from West African (MAU) ancestry, in line with TreeMix analyses and f₄ statistics (Fig. 5). No such models included genetic contributions from mainland Spain or northern Europe (IRE, Ireland; DEN, Denmark). qpAdm modeling indicated that the general affinity of the Chupaderos genome and modern populations from Europe and the TUK population from Anatolia, as most one-way models, could not be rejected. This was also true when considering the Roman specimens from Boinville, France, and the postmedieval specimen from Fiumarella, Italy, as a possible source. Other ancient specimens from Europe included a larger range of possible population sources. This indicates donkey genetic profiles changing through space and time during the past 2000 years in Europe, including variable degrees of West African ancestry, with some ancient specimens having a more pervasive and widespread genetic legacy in the region today.

Combined, our analyses indicate that the donkey specimen excavated at Jamestown exhibited a genetic profile most characteristic of European ancestry but also included West African ancestry, as the best model in our dataset depicts Mauritania as a source. The latter genetic contribution was absent in the ancient specimen from Chupaderos, Mexico, which indicates different population sources for the origin of the two donkey specimens. While the specimen from Chupaderos appeared genetically close to the Pega donkey from Brazil and modern donkeys from the Canary Islands, these specific genetic affinities were not found in the genome of the specimen from Jamestown.

Isotopic analysis

The two teeth subjected to isotope analysis (fig. S1 and tables S2 to S5), a horse upper third molar (M3) and a donkey upper second molar (M2), represent sequences that mineralized at different developmental stages, with the horse M3 mineralizing during the third and fourth year [25 to 55 months (23)] and the donkey M2 beginning mineralization toward the end of the first year [8 to 18 months (15)]. Hence, the isotopic profiles of both animals represent time averages encompassing multiple seasons.

The horse average enamel δ¹³C value of −13.2‰ indicates a C₃ plant–based diet typical for temperate climate and vegetation zones, whereas the donkey average enamel δ¹³C value of −8.8‰ suggests a mix of C₃ and C₄ plants, which could include wild C₄ grasses but also domesticated C₄ plants such as millet, sorghum, and maize, found in warmer, dryer regions. This finding of broad differences in fodder is supported by the disparate δ¹⁸O values in the two equids. The horse exhibits an average δ¹⁸O value of −6.0‰, which is ¹⁸O-depleted compared to that of the donkey, with an average δ¹⁸O value of −2.5‰. This again strongly suggests different points of origin for these two equids.

However, the radiogenic isotope ratios of strontium and lead are similar among the two equids sampled, and the respective averaged ⁸⁷Sr/⁸⁶Sr ratios of 0.7104 and 0.7100 are found in many regions of the world. The averaged Pb ratios of the horse and donkey enamel samples show a narrow range, with ²⁰⁶Pb/²⁰⁴Pb ranging from 18.410 to 18.504, ²⁰⁷Pb/²⁰⁴Pb from 15.625 to 15.636, and ²⁰⁸Pb/²⁰⁴Pb from 38.367 to 38.449 (table S2). These values are not typically found in the Americas (24) and likely derive from metal artifacts the two equids were exposed to either during life or through postmortem diagenesis (section S4). Matching these isotope values to available isoscape resources for Britain (25) suggests that, while the ⁸⁷Sr/⁸⁶Sr and ²⁰⁶Pb/²⁰⁴Pb ratios of the horse are present in the biosphere of modern Britain (fig. S2A), its calculated carbonate δ¹⁸O values converted to precipitation/drinking water δ¹⁸O values (−10.5‰) lie outside of the range of precipitation within Britain but align with published δ¹⁸O for both modern horses (26) and archaeological British horse tooth enamel from the late Medieval/Tudor period (27). Together, these data indicate that Britain is a plausible place of origin for the Jamestown horse. Similarly, the average ⁸⁷Sr/⁸⁶Sr ratios of the donkey match larger regions of Britain (fig. S2C) but only overlap with the modeled donkey’s drinking water δ¹⁸O values in the far southwestern coast and, hence, the warmest region of Britain (fig. S2D). However, the donkey’s δ¹⁸O (−2.5‰) is much higher than the published δ¹⁸O range in historical horses from Britain (26, 28). This implies that the donkey likely originated from a warmer climatic region.

Trace metal concentrations

The maximum threshold concentrations (MTC) of key trace elements and rare earth elements were analyzed (27) for all samples assayed (section S4). Although vanadium, uranium, and thorium in some of the samples are elevated above the MTC and indicate some degree of diagenetic alteration, the rest of the elements are within the possible in vivo range. One intriguing observation in the trace element data indicates a possible anthropogenic in vivo exposure to Ni for the animals. Both the donkey and horse tooth enamel show zones with extremely high Ni concentrations, well beyond values expected from natural diet and/or even diagenetic alteration. The horse enamel shows Ni as high as 1684 parts per million (ppm), and the donkey with an extremely high value (Ni = 6847 ppm; table S3).

DISCUSSION

Consumption of domestic horses

Archaeozoological and taphonomic results demonstrate previously undocumented levels of intensive processing and consumption of domestic equids at Jamestown. Taphonomic data show that adult horses were eaten, butchered, and cooked or boiled, with most elements split open to extract even the minutest nutritional resources including dental pulp. These findings mirror historic texts describing the Starving Time consumption of horses as a last resort. John Smith in his Generall Historie (1624) describes the dire situation of the colony after his October 1609 departure, writing based on the observations of a witness to the Starving Time, “as for our Hogs, Hens, Goats, Sheepe, Horse, or what lived, our commanders, officers, and salvages daily consumed them,” noting that even the skin of the horses was eaten (8).

Use of horses for transport

Osteological data indicate that either during the animals’ life at Jamestown or prior, at least some of the first domestic horses at the Jamestown colony were used for transport. Fractures to the enamel of a lower second premolar suggest bridling, while an asymmetric cross-sectional profile for a metapodial could reflect load and activity patterns linked with transport. These findings align well with recent investigation of equestrian artifacts from early James Fort–period contexts, including the First Well and Kitchen Cellar, in which seven sets of tack were also recovered. The two limb fractures identified are not necessarily linked to transport but might relate to confinement of animals during transatlantic transit or during daily life at Jamestown.

Presence of juvenile horse and donkey

Archaeozoological analyses reveal the presence of previously undocumented domestic equids during the earliest years at Jamestown. Although both male and female horses were mentioned in first-person accounts of the Third Supply that arrived at Jamestown in late summer 1609, the presence of a juvenile horse specimen shows that imported horses had successfully begun to breed and reproduce at the colony before their slaughter. Notably, archaeofaunal materials demonstrate the previously unknown presence of a domestic donkey during the initial settlement of the Jamestown colony. Although donkeys were not mentioned in chronicles of the first voyages, John Smith included both horses and asses in a list of animals that would be likely to thrive in Virginia, based on the success of “them that were carried thither,” suggesting that Smith had, in fact, witnessed donkeys in the colony prior to his departure in 1609 (8).

Biomolecular evidence for transatlantic exchange

Various lines of isotopic and genetic data support a British origin for the Jamestown horses but suggest that the newly identified donkey was acquired somewhere during transatlantic travel (Iberia, West Africa, or the Caribbean). Isotope data may be partly contradictory due to equifinality, as isotopic characteristics may overlap between geographic regions and most parts of the world are still not mapped for the complex isotopic variation found in the environment (29, 30). Nonetheless, these datasets can help exclude potential source regions, when considered in tandem and paired with knowledge of the Jamestown voyage, and provide important guidance for understanding areas of animal origin.

The analyzed horse sample has δ¹⁸O and ⁸⁷Sr/⁸⁶Sr values that fit within the range observed in British horses (26, 28). This is in agreement with the C₃ fodder this equid received for the temperate climate of the British Isles. Furthermore, the observed −13.2‰ for δ¹³C in the horse is close to the average −13.3‰ reported for British horses (28). A mismatch between the enamel δ¹⁸O values of archaeological horses and predictions for δ¹⁸O in modern precipitation was recently reported for horses from a medieval site in London (28), suggesting that the modern day meteoric δ¹⁸O values do not reflect the drinking water values of the historical past, a pattern perhaps associated in part with the Little Ice Age. After adjusting the δ¹⁸O values for an overall colder climate, the horse values appear consistent with this trend toward cooler conditions (section S4).

In contrast, the donkey’s δ¹⁸O values suggest provenience in warmer climates overall, fitting particularly well into δ¹⁸O isoscape predictions for the Mediterranean, including the Iberian Peninsula and North Africa (25) as well as the Caribbean (29), but not Britain. This is supported by the mixed C₃ and C₄ dietary signal in the donkey specimen, suggesting that the fodder of this animal derived from a drier, warmer environment, home to wild and domesticated C₄ grasses, including maize, sorghum, or millet, which were likely completely absent in 16th and early 17th century Britain. Hence, both the δ¹⁸O and δ¹³C data suggest possible origins for the donkey along the transatlantic voyage route.

Strontium isotope data from this donkey suggest that the animal may have had origins in either West Africa itself or the wider trade network associated with this region (31), stretching from mid-Atlantic to the Iberian Peninsula. The donkey’s ⁸⁷Sr/⁸⁶Sr ratios of 0.7100 exceed the range of published Sr values from the Canary Islands, which tend to lower values from the islands’ young volcanic geology (32). Similar values are rare in the Caribbean, outside of Trinidad, and are only found in biotic samples on metamorphic rocks in small parts of the islands, but more common in southern Portugal and to a lesser extent in western Spain. Bioavailable strontium isotope ratios vary greatly in continental West Africa [0.709 to 0.730 (33)] and the Iberian Peninsula [0.7046 to 0.735 (34, 35)], although less so in the Caribbean [0.706 to 0.711 (36)]. Although no bioavailable ⁸⁷Sr/⁸⁶Sr data are yet available for North Africa, a novel strontium isoscape of West Africa allows us to suggest that, within this region, the observed values are most similar to reference data from Senegambia and Sierra Leone (Fig. 6). Mauritania is dominated by the same Cenozoic/Quaternary sediments as Senegal and likely to have similar ⁸⁷Sr/⁸⁶Sr, consistent with values observed with the Jamestown donkey.

Fig. 6. — The probability of origin in West Africa based on ⁸⁷Sr/⁸⁶Sr and δ¹⁸O derived from tooth enamel, calculated and visualized for the donkey specimen from Jamestown. Highest probabilities of 80 to 90% can be found in coastal Senegambia and Sierra Leone. Presumably, the geology and climate of northern Senegal and Mauritania (not yet mapped for ⁸⁷Sr/⁸⁶Sr) would be an equally good fit.

Given these isotope values and the known trajectory of the Third Supply voyage (Fig. 1A), plausible scenarios for the donkey’s origin include acquisition of the donkey during an undocumented stop along the Iberian coast, members of the voyage trading for an animal raised in mainland West Africa during a documented stopover in the Canaries, or acquisition of a nonlocal animal raised in a more distant point of origin (e.g., Trinidad) during transit through the Caribbean.

When considered alongside genomic data showing a blend of West African and Iberian ancestry, these data appear to mirror patterns emerging from other taxa, such as cattle, showing a substantial contribution from African sources in the early colonial fauna from the Americas (37), and point toward a donkey drawn from emerging multicontinental and multicultural transatlantic trade networks, blending populations from Africa and Europe into complex biological and economic networks.

Scarcity and supply of early horses and donkeys after the Starving Time

After the Starving Time, little to no documentary evidence records horses or donkeys being present in the colony, although the 1610 Martial Laws stipulated that the killing of a mare or horse was punishable by death (38). Colonists expressed a preference for donkeys in correspondence, with one 1613 letter asking for “beasts to manure the land, either horses or asses, but indeed the latter is best for us” and for “100 she-asses” and several horses (39). The emerging demand for donkeys represents a departure from traditional English agricultural lifeways, which were best adapted for temperate landscapes and initially resulted in preferential success in places that recapitulated the climate of the British Isles (40, 41). Findings from the earliest layers at Jamestown demonstrate the social and logistic challenges of supply and survival associated with transatlantic transit and establishment of early equid populations in eastern North American colonies and point to other transatlantic faunal inputs into early British settlements on the eastern seaboard.

Interdisciplinary scientific study of faunal remains associated with the earliest English settlement of the eastern seaboard of the United States shows that, consistent with historic records, domestic horses were transported across the Atlantic from the British Isles where they were used for transport and successfully reproduced before meeting their end in the Starving Time winter of 1609. Modification of these horse remains shows that extraordinarily intensive processing was used to survive this winter, including butchering, cooking, and extracting marrow and pulp from nearly all the equids found at the site. Discoveries of donkey remains demonstrate a previously unknown dispersal of domestic donkeys into the colony, likely involving undocumented trade with emerging transatlantic networks of biological exchange linking Europe, Africa, and the Americas via the Caribbean. These results show the extraordinary diversity of emerging transatlantic trade networks in livestock during the early 17th century and demonstrate the impact of not only horse but also donkey in the earliest chapter of domestic livestock in the eastern seaboard of North America.

MATERIALS AND METHODS

Osteological study

To understand the role of domestic equids during the initial settlement of Jamestown, we conducted detailed osteological, taphonomic, and paleopathological study of 63 of the 77 specimens morphologically confirmed as members of the Equus genus. For each specimen, we first identified the animal to taxon based on size and morphological comparisons to reference data from specimens in the Archaeozoology Laboratory at the University of Colorado–Boulder. For specimens with intact epiphyses or visible dentition, we assessed the likely age of each identified specimen using eruption and wear tables from Levine (42) and Evans et al. (43), and for cheek teeth, crown height (42) was measured with Mitutoyo digital caliper. Although sex estimation is also sometimes possible based on pelvic morphology or the presence of intact mandibles and maxillae, the fragmented nature of the assemblage did not permit sex estimation in the assemblage.

Taphonomy

For each specimen, we assessed life history, health, and pathology information when possible, including evidence for fractures and disease (44), transport-linked damage to the dentition (16), and changes to the internal structure of the limbs (18, 19). We also assessed each specimen for evidence of nonhuman taphonomic processes such as weathering, root etching, and carnivore gnawing, as well as for cultural modifications including cut and chop marks, spiral fracturing, burning, pot polish, and other indicators of human activity. Each likely cut mark was also assessed under 20× and 200× microscopy using a Dino-Lite digital microscope, and marks lacking a distinct V-shaped profile were excluded from the analysis.

Radiocarbon dating

We selected five equid tooth specimens for direct radiocarbon dating using accelerator mass spectrometry (AMS) at the Keck-AMS facility at the University of California–Irvine, of which three produced successful measurements (section S2). Before conducting destructive analysis, each specimen was scanned using an EinScan-SE structured light scanner to produce a 3D model of morphological data. We then prepared a simple Bayesian uniform phase model estimate for horse activity at the initial Jamestown colony following the methods of Ramsey (45), using an initial bounding date of the arrival of 1607, and assumed the known historical date of the voyage arrival based on historic records.

Isotopic analysis

A suite of isotopic analyses were conducted on two specimens from the site, a horse (JR2718K, no. 117576) and the donkey (JR3081F, no. 121161), in the Bone Chemistry Laboratory in the Department of Anthropology, University of Florida. Each tooth was thoroughly cleaned, excess plaque and cementum were mechanically removed using a dental drill with a tungsten carbide burr bit, and the exposed surface was thoroughly cleaned to a polish. Sequential samples of tooth enamel were then drilled parallel to the growth axis, with bulk average data provided listed in table S2 (section S4). There was no pretreatment of collected powders analyzed for this study. All data are from tooth enamel structural carbonate and capture that point in time during the mineralization of each specimen. Two sets of analyses were conducted in the Department of Geological Sciences, University of Florida: (i) Light isotope analysis on nonpretreated tooth enamel (structural carbonate) was measured by isotope ratio mass spectrometry for carbon (δ¹³C) and oxygen (δ¹⁸O); (ii) Radiogenic isotope analysis on “pooled” lanes of tooth enamel powder was subject to ion chromatography, and separate samples were measured using multiple collector–inductively coupled plasma mass spectrometry for strontium (⁸⁷Sr/⁸⁶Sr) and lead (²⁰ⁿPb/²⁰⁴Pb) (section S4).

DNA

A total of five equid archaeological remains were analyzed for DNA content at the ancient DNA facilities of the Centre for Anthropobiology and Genomics of Toulouse (France). The experimental procedure followed the work of Librado et al. (46) and Todd et al. (22) (section S5). DNA screening revealed excellent DNA preservation in the donkey jack sample labeled JT02/no. 121161 (section S1, Fig. 5, and figs. S3 to S6). We then conducted phylogenetic analysis using neighbor-joining and TreeMix analyses and explored the relationship of this specimen to global modern and ancient population structure using PCA, Struct-f₄, and qpAdm (section S5). The sequence data generated in this study are available for download on the European Nucleotide Archive (accession no. PRJEB89422).

Acknowledgments

We thank J. Curtis (UF) for the light isotope analysis and J. Southon at UCI-Keck AMS facility for the radiocarbon dating.

Funding: This research was funded through an award by the National Science Foundation (NSF no. 1949305, “Horses and Human Societies in the American West”) and by France Génomique National Infrastructure, as part of “Investissement d’avenir” program managed by Agence Nationale pour la Recherche (ANR-10-INBS-09); the CNRS International Research Project AnimalFarm; the France Génomique Appel à Grand Projet (ANR-10-INBS-09-08, MARENGO project); and the European Research Council (ERC) under the European Union’s Horizon 2020 research and innovation programme (grant agreements 681605-PEGASUS and 101071707-HorsePower). Publication of this article was funded by the University of Colorado–Boulder Libraries Open Access Fund.

Author contributions: W.T.T.T. collected archaeozoological data and wrote the paper. N.D., V.M.O., G.K., L.O., and J.K. helped write the paper and collected/analyzed biomolecular data. P.M., M.L., and L.S. helped write the paper and analyzed historical and archaeological data. A.O., L.H., C.I.B.-O., O.O., D.Q.-B., L.K.-D., D.M., M.E.H., A.E.T., A.W., D.W.P., L.C., S.S., L.C.-T., L.B., A.B.-M., and J.S. contributed data and/or analyzed data and edited the manuscript.

Competing interests: The authors declare that they have no competing interests.

Data and materials availability: All data needed to evaluate the conclusions in the paper are present in the paper and/or the Supplementary Materials.

Supplementary Materials

The PDF file includes:

Supplementary Sections S1 to S6

Figs. S1 to S7

Tables S1 to S5

Legend for data S1

References

sciadv.adw2595_sm.pdf^{(1.4MB, pdf)}

Other Supplementary Material for this manuscript includes the following:

Data S1

sciadv.adw2595_data_s1.zip^{(12.8KB, zip)}

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Associated Data

This section collects any data citations, data availability statements, or supplementary materials included in this article.

Supplementary Materials

Supplementary Sections S1 to S6

Figs. S1 to S7

Tables S1 to S5

Legend for data S1

References

sciadv.adw2595_sm.pdf^{(1.4MB, pdf)}

Data S1

sciadv.adw2595_data_s1.zip^{(12.8KB, zip)}

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PERMALINK

Early transatlantic movement of horses and donkeys at Jamestown

William Timothy Treal Taylor

Nicolas Delsol

Vicky M Oelze

Peter Mitchell

Leah Stricker

Michael Lavin

Akin Ogundiran

Lauren Hosek

Christina Isabelle Barrón-Ortiz

Olumide Ojediran

Diana Quintero-Bisono

Lucy Keith-Diagne

Dane Magoon

Matthew E Hill Jr

Ariane E Thomas

Anna Waterman

David W Peate

Lorelei Chauvey

Stéphanie Schiavinato

Laure Calvière-Tonasso

Luís Borges

Aitor Brito-Mayor

Jonathan Santana

George Kamenov

Ludovic Orlando

John Krigbaum

Roles

Abstract

INTRODUCTION

British horses, donkeys, and early North America

Jamestown and the earliest British equids

Fig. 1. Jamestown site context, equid remains, and radiocarbon dating.

RESULTS

Radiocarbon dating

Elements present

Species identification

Fig. 2. Equid identifications and paleopathology.

Demographics

Pathology

Taphonomy

Fig. 3. Tooth modifications in the Jamestown Starving Time equid assemblage.

Fig. 4. Modified Starving Time–era horse bones from Jamestown.

DNA analysis

Fig. 5. Genomic analyses.

Isotopic analysis

Trace metal concentrations

DISCUSSION

Consumption of domestic horses

Use of horses for transport

Presence of juvenile horse and donkey

Biomolecular evidence for transatlantic exchange

Fig. 6. Strontium isotope comparisons, West Africa.

Scarcity and supply of early horses and donkeys after the Starving Time

MATERIALS AND METHODS

Osteological study

Taphonomy

Radiocarbon dating

Isotopic analysis

DNA

Acknowledgments

Supplementary Materials

The PDF file includes:

Other Supplementary Material for this manuscript includes the following:

REFERENCES AND NOTES

Associated Data

Supplementary Materials

ACTIONS

PERMALINK

RESOURCES

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