The colonial-era remains raised deep questions. Who were they? Were they related? Where were they from?

Now, DNA analysis published Monday in the Proceedings of the National Academy of Sciences offers new layers to their complex story, which has been slowly unfolding thanks to an ongoing collaboration between anthropological geneticists and the Gullah Society, a nonprofit focused on preserving African American burial grounds. The Gullah Society was dissolved in 2021 after its founder, Ade Ofunniyin, died, but the work continues under the Anson Street African Burial Ground Project.

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The first round of research published three years ago presented detailed studies of the bones and an analysis of their mitochondrial DNA, which is inherited from mothers. That work revealed their approximate ages, sexes and their maternal African ancestry. The researchers concluded that these nameless people — they dubbed them the Ancestors — were probably enslaved people.

It also inspired a ceremony in which Yoruba priests gave each person an honorary name, guided in part by what could be gleaned from scientific analysis of their remains.

The ancestors were mostly male, ranging in age from infants to older adults. Six of them — Banza, Kuto, Zimbu, Daba, Ganda and Talata — were probably abducted in Africa and brought to Charleston or born along the journey. The others were born in the Lowcountry around Charleston but traced their ancestry to diverse parts of West and Central Africa. Coosaw, a female adolescent, was partly Native American.

The newest results confirm those findings and expand on them. Mapping their genomes gave researchers more insight into their ancestry and helped answer the question of whether they were related to one another. They weren’t, with the exception of Isi and Welela. The 36 people appear to have been buried as they died, rather than in family groups or in a mass grave.

“The ancestors represent thousands and thousands of people whose history and existence is largely unrecorded, if not entirely unrecorded,” said Theodore Schurr, an anthropologist at the University of Pennsylvania and one of the leaders of the work. “They represent in spiritual, as well as historical and biological ways, the ancestors of people of African descent living in Charleston and elsewhere today.”

The insight into forgotten lives is possible because of serendipity, advocacy and a culturally sensitive approach. Scientists went into the work with their own questions about what they could learn, but they also asked the African American community in Charleston what they wanted to know about these remains.

The community had specific questions. Were women and children buried there? Were they related? Where were they from?

The physical study of the bones themselves could only get so far, and the first round of analysis of mitochondrial DNA gave a limited window into their maternal ancestry. With the permission of the community, the scientists extracted DNA from fragments of a skull bone and teeth to learn more. Such analyses must be performed under clean-room conditions to avoid contamination with modern DNA.

Raquel Fleskes, an anthropological geneticist at the University of Connecticut, wore a GoPro camera to document the process of extracting DNA from the bone samples, to share the experience with the community.

The results offer a mix of revelations. Most of the people buried in the graveyard were born in Charleston, but they would have traced their African roots to very different regions and cultures. By comparing their DNA to modern-day populations, the researchers found that three matched closely with people from Gabon. Four bore close ties to people from Ghana, Ivory Coast and Sierra Leone. One person, Lisa, appeared to have ties to Gambia.

“These ancestors are really diverse individuals, they’re coming from all parts of Africa,” Fleskes said.

Next, the researchers hope to analyze samples from their teeth — sampling the oral microbiome — to see what they can discern about the ancestors’ diet, and perhaps find clues about any diseases they may have suffered.

Anne Stone, an anthropological geneticist at Arizona State University who edited the paper for PNAS but was not involved in the research, said that the genetic study adds a dimension of knowledge that could only be hinted at by detailed study of the bones.

“I think that this work is important because it provides another source of information about the lives of enslaved Africans, and particularly helps shed light on the relationships among individuals and their ancestry,” Stone said in an email.

Although the insight into the ancestors’ lives is fragmented, Schurr said the diversity of the people in the graveyard speaks to the brutality of their lives. If these were enslaved people, they would have been separated from family and friends indiscriminately, in part because connections that might have helped foster resistance were usually stamped out.

“It speaks to the structural violence of slavery, and the demeaning of humanity of these individuals, not allowing them to be with kin folk, or people from the same cultural groups, the same ethnic populations,” Schurr said.

Despite the lack of any clear genetic connections between most of the ancestors, the burials also demonstrate a high level of care. There were nails and brass pins in the soil, suggesting that the bodies had been interred in caskets or burial shrouds. Tokens scattered among the remains appeared to be a sign of honor from the community.

“It was done so respectfully, so honorably and well cared for, that you could tell that the people were buried by — I don’t know if they were relatives,” said La’Sheia Oubré, who leads education and community engagement efforts for the Anson Street African Burial Ground Project.

“In the African American community in Charleston, you don’t have to be related to someone by blood to care for them.”