Why are there no persisting hybrids of humans with Denisovans, Neanderthals, or anyone else?

April 1, 2016
113 (17) E2354
Research Article
Nuclear and mitochondrial DNA sequences from two Denisovan individuals
Susanna Sawyer, Gabriel Renaud [...] Svante Pääbo
The tour-de-force report of Sawyer et al. (1) on genomes of two Denisovans and the accompanying editorial and figure (2) support the notion of “a web of now-extinct populations linked by limited, but intermittent…gene flow” (3): involving multiple hominin lineages for thousands of years, before the mysterious disappearance of all taxa other than us “behaviorally modern humans” (BMHs). Although attention focuses on rare introgressions of non-BMH alleles facilitating adaptation of invading BMHs to ecological challenges, there is a bigger elephant in the room.
Current genomic and archaeological data indicate that BMHs arose in Africa ∼100,000–200,000 y ago and spread across the planet (including the rest of Africa), encountering other extant hominins like Neanderthals, Denisovans, archaic African hominins, and possibly other lineages from earlier diasporas of Homo erectus. Although genomic evidence indicates interbreeding, the number of functional genes incorporated is limited, resulting in a “leaky replacement” (3), without persistence of true hybrids. Thus, our single BMH (sub)species was the “winner” in every contact/replacement event, spanning tens of thousands of years. I cannot find any other example wherein a single (sub)species from one geographic origin completely replaced all extant cross-fertile (sub)species in every planetary location, with limited introgression of functional genetic material from replaced taxa, and leaving no hybrid species. Typically, one instead finds multiple cross-fertile (sub)species, with hybrid zones in between.
Although this apparent one-of-a-kind phenomenon could have occurred by chance, the singularity allows one to posit a uniquely complex genetic/biological/cultural transition of BMHs. As Pääbo suggested (3), adaptive accumulation of an “explosive constellation” of genetic variants (alleles) could have endowed BMHs with an unparalleled combination of cognitive features, guaranteeing success at every subsequent encounter with other hominins.
Why did hybrid species not persist, at least at the geographical extremes of BMH expansion? Assuming that hundreds of new alleles comprised the BMH genotype, F1 hybrids with other hominins would likely lack the complete cognitive package required to compete for mating within BMH groups. Tellingly, 10 of 10 non-BMH mitochondrial sequences are outside the current BMH range (1), suggesting that mating of BMH males with non-BMH females generated progeny that were not included within BMH groups. In contrast, progeny of female BMHs and non-BMH males may have had the opportunity to survive within BMH groups, with sufficient mating success rates to allow transmissions of a few alleles valuable to the newcomers, but related to ecological adaptation, not cognition.
Such “human exceptionalism” is currently frowned upon, as are extraordinary explanations of evolutionary events. However, unless there are other clear examples of such complete replacement of all related taxa by one single (sub)species, BMHs may indeed be a rare exception. Although environmental factors such as climate or infectious disease (4) could have generated the initial African bottleneck, the critical BMH phenotype was likely cognitive. This fits ecocultural models predicting Neanderthal extinction through competition with modern humans (5) and suggests an improbable BMH transition through a long-standing “psychological evolutionary barrier”––possibly involving initially maladaptive features such as reality denial and mortality salience, which conspired to generate the winning combination (6).


I acknowledge valuable comments from Pascal Gagneux.


S Sawyer, et al., Nuclear and mitochondrial DNA sequences from two Denisovan individuals. Proc Natl Acad Sci USA 112, 15696–15700 (2015).
CB Stringer, I Barnes, Deciphering the Denisovans. Proc Natl Acad Sci USA 112, 15542–15543 (2015).
S Pääbo, The diverse origins of the human gene pool. Nat Rev Genet 16, 313–314 (2015).
X Wang, et al., Specific inactivation of two immunomodulatory SIGLEC genes during human evolution. Proc Natl Acad Sci USA; NISC Comparative Sequencing Program 109, 9935–9940 (2012).
W Gilpin, MW Feldman, K Aoki, An ecocultural model predicts Neanderthal extinction through competition with modern humans. Proc Natl Acad Sci USA 113, 2134–2139 (2016).
A Varki, Human uniqueness and the denial of death. Nature 460, 684 (2009).

Information & Authors


Published in

Go to Proceedings of the National Academy of Sciences
Go to Proceedings of the National Academy of Sciences
Proceedings of the National Academy of Sciences
Vol. 113 | No. 17
April 26, 2016
PubMed: 27044111


Submission history

Published online: April 1, 2016
Published in issue: April 26, 2016


I acknowledge valuable comments from Pascal Gagneux.



University of California, San Diego/Salk Institute Center for Academic Research and Training in Anthropogeny (CARTA), La Jolla, CA 92093


Author contributions: A.V. wrote the paper.

Competing Interests

The author declares no conflict of interest.

Metrics & Citations


Note: The article usage is presented with a three- to four-day delay and will update daily once available. Due to ths delay, usage data will not appear immediately following publication. Citation information is sourced from Crossref Cited-by service.

Citation statements



If you have the appropriate software installed, you can download article citation data to the citation manager of your choice. Simply select your manager software from the list below and click Download.

Cited by


    View Options

    View options

    PDF format

    Download this article as a PDF file


    Get Access

    Login options

    Check if you have access through your login credentials or your institution to get full access on this article.

    Personal login Institutional Login

    Recommend to a librarian

    Recommend PNAS to a Librarian

    Purchase options

    Purchase this article to get full access to it.

    Single Article Purchase

    Why are there no persisting hybrids of humans with Denisovans, Neanderthals, or anyone else?
    Proceedings of the National Academy of Sciences
    • Vol. 113
    • No. 17
    • pp. 4543-E2471







    Share article link

    Share on social media