Bound by nature - Cultural evolution has not freed hunter-gatherers from environmental forcing

Cultural evolution has made humans enormously potent ecosystem engineers and has enabled us to survive and flourish under a variety environmental conditions.

Even hunter-gatherers, who obtain their food from wild plant and animal resources using seemingly simple technologies, have been able to extract energy in harsh arctic and desert conditions and compile vast knowledge on medical plants to fight against pathogens in the tropics. Because of culture, humans are often considered to be divorced from the environment and not under the same ecological forcing as other species.

Hunter-gatherer population density is constrained by environmental conditions

However, in a new paper published in Proceedings of the National of Sciences USA (PNAS) scientists from the University of Helsinki, Faculty of Science, show that key environmental parameters, namely climate-related primary productivity, biodiversity, and pathogen stress have strong influence on the global pattern of population densities of ethnographically documented hunter-gatherers.

“These results demonstrate that cultural evolution has not freed human hunter-gatherers from strong environmental forcings”, says Dr Miikka Tallavaara.

Downside of the tropical affluence: abundance of pathogens

Interestingly, the importance of the three environmental variables varies between different parts of the globe: Productivity and biodiversity exert the strongest influence on population density in high- and mid-latitudes, whereas pathogen stress becomes more important in the tropics.

“Our results suggest that temperate and subtropical biomes provide the most suitable conditions for pre-agricultural humans due to the optimum between the positive effects of resource availability and the negative effects of pathogens”, adds professor Miska Luoto.

The role of productive environments in the beginning of agriculture

A large fraction of the area of these highly suitable biomes has nevertheless been occupied by agricultural populations during the last 6000 years. Competing theories about the origins of agriculture emphasize the role of either productive and stable environments or population packing among hunter-gatherers as prime movers behind the transition from hunting and gathering to farming.

Dr Tallavaara says: “Our study highlights the difficulty in judging between environmental and demographic causes of the transition, because resource availability and hunter-gatherer population density appear to be closely linked”.

Environment has affected humans throughout our evolutionary history

Most of the history of our species, humans have been hunters and gatherers. Thus, the results of this study imply that human populations have been influenced by environmental conditions throughout their evolutionary history.

“Environment has been crucial driver of human population dynamics up to the end of the pre-industrial era but we still are heavily depended on natural conditions and life-support systems, although modern technology has made it more invisible to most people in daily life”, says Docent Jussi T. Eronen.

The paper, published in the journal Proceedings of the National Academy of Sciences of the United States (PNAS) is entitled “Productivity, biodiversity, and pathogens influence the global hunter-gatherer population density”.

More information:

Dr Miikka Tallavaara, University of Helsinki, email:

Professor Miska Luoto, University of Helsinki, +358 44 272 7327, email:

Docent Jussi T. Eronen, University of Helsinki, email:

Climate driven net primary productivity, biodiversity, and pathogens constrain the global hunter-gatherer population density

However, the most influential drivers of population density are context-dependent. Biodiversity affects population density mostly in the low-productivity environments, whereas pathogen stress, which has insignificant effects in areas of low productivity, is a crucial constraint on population density in the high-productivity settings, especially the tropics. These findings highlight the key role that environmental factors play in shaping the global population density patterns of pre-agricultural humans. Blue, red, and grey arrows indicate positive, negative and statistically insignificant effect of a variable. Thickness of an arrow indicate the strength of the effect of a variable. Net primary productivity has also indirect effects on population density mediated by biodiversity and pathogens.