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Demic diffusion

Demic diffusion is a demographic model describing the spread of cultural traits, such as agriculture, through the migration, reproduction, and interbreeding of populations, resulting in significant genetic contributions to recipient groups.^[1] This process contrasts with cultural diffusion, which involves the transmission of ideas or practices without substantial population movement, such as local hunter-gatherers adopting farming techniques from neighboring groups.^[2] Originally proposed to explain the Neolithic transition in Europe, the model posits that farming expanded from the Near East around 10,000 years ago via successive waves of farmer dispersal into hunter-gatherer territories, creating clinal gradients in genetic and archaeological data.^[1] The concept was first formalized by Albert J. Ammerman and Luigi Luca Cavalli-Sforza in their 1971 analysis of archaeological site dates, which estimated the rate of Neolithic spread across Europe at approximately 1 km per year using a wave-of-advance mathematical framework.^[1] Their subsequent 1984 book expanded this into a comprehensive demic diffusion hypothesis, integrating genetic, linguistic, and archaeological evidence to argue for a major influx of Near Eastern farmers through admixture with indigenous populations.^[1] This model highlights how demographic pressures, such as higher reproductive rates among farmers, drove iterative short-range colonizations, rather than long-distance leaps or purely ideational transfers.^[3] Genetic studies have provided strong support for demic diffusion, particularly in southern and central Europe, where ancestry components show a southeast-to-northwest cline indicative of Neolithic farmer input.^[1] For instance, ancient DNA analyses estimate an average ~40% Neolithic farmer autosomal ancestry in modern Europeans, higher in the south and decreasing northward due to later admixtures.^[4] Regional variations exist: demic processes dominated in the Balkans and Central Europe with advance speeds of 0.68–1.48 km/year, while cultural diffusion was more prominent in northern and Alpine regions at slower rates below 0.66 km/year.^[2] In Scandinavia, demic diffusion accounted for over 50% of the Neolithic spread, with hunter-gatherer incorporation limited to fewer than 8 individuals per 10 pioneering farmers.^[3] Recent 2025 studies using advanced modeling and ancient DNA further confirm the predominance of demic diffusion, while highlighting increasing local admixture with hunter-gatherers during the expansion.^[5]^[4] These findings underscore demic diffusion's role in shaping Europe's genetic landscape, though ongoing research refines the balance with cultural mechanisms.^[3]

Overview

Definition

Demic diffusion refers to the spread of cultural, technological, or genetic traits through the physical migration and subsequent reproduction of human populations, rather than through the transmission of ideas or practices alone.^[6] This process emphasizes the role of population movements in facilitating the dissemination of innovations, such as agriculture, by enabling the establishment of new settlements and intermixing with local groups. Key characteristics of demic diffusion include gradual population expansion driven by short-distance migrations, often over generations, which result in gene flow, partial replacement of indigenous populations, or cultural admixture.^[7] These migrations typically occur as small groups move into adjacent territories, reproduce, and establish daughter populations that continue the pattern, creating a wavefront of trait propagation. The basic mathematical model for demic diffusion is represented as a wave of advance, described by the Fisher-Kolmogorov equation:

\frac{\partial p}{\partial t} = D \nabla^2 p + r p (1 - p)

where p is the population density, D is the diffusion coefficient reflecting migration rates, and r is the intrinsic growth rate.^[8] This equation captures the interplay between diffusive spread and logistic population growth, predicting a constant velocity for the advancing front.^[2] The concept of demic diffusion was initially conceptualized by Albert J. Ammerman and Luigi Luca Cavalli-Sforza in 1971, in the context of the Neolithic farming spread across Europe.^[9]

Distinction from Cultural Diffusion

Cultural diffusion refers to the transmission of cultural elements, such as ideas, technologies, beliefs, or practices, from one society or group to another primarily through mechanisms like imitation, trade, social interaction, or exchange, without requiring substantial population relocation.^[10]^[11] This process emphasizes the movement of intangible traits across existing populations, often preserving the genetic makeup of the adopting groups while altering their behaviors or material culture. In distinction from demic diffusion, cultural diffusion does not inherently involve the migration of people or the introduction of new genetic lineages, focusing instead on the adoption of innovations through learning and contact.^[2] Demic diffusion, by contrast, entails the physical dispersal of human populations carrying both cultural traits and genetic markers, potentially leading to genetic continuity with source populations, admixture, or even partial replacement of indigenous groups in the target areas.^[1] This fundamental difference highlights demic diffusion's demographic dimension—driven by population growth, movement, and reproduction—versus cultural diffusion's reliance on social and economic networks that facilitate idea propagation over distances without mass relocation.^[12] A clear example of this distinction appears in the Neolithic spread of agriculture across Europe, where demic diffusion manifests as early farmers migrating from the Near East, establishing settlements, and interbreeding with local hunter-gatherers, resulting in observable genetic signatures of admixture.^[2] In a cultural diffusion scenario, the same hunter-gatherer populations might acquire farming techniques through trade or observation from distant groups, integrating these practices into their lifestyles without undergoing demographic shifts or genetic influx from migrants.^[13] These mechanisms often operate on a spectrum rather than as mutually exclusive processes, ranging from pure demic diffusion (fully migration-driven, with 100% population movement) to pure cultural diffusion (0% migration, entirely idea-based), though empirical studies of historical expansions frequently reveal hybrids.^[12] For instance, earlier models of the European Neolithic transition estimated that demic diffusion contributed approximately 60% to the overall spread, complemented by 40% cultural diffusion,^[14] while recent studies as of 2025 suggest predominant demic diffusion with minimal cultural contribution (~0.5%).^[15] This hybrid nature, first systematically explored in the demic framework by Ammerman and Cavalli-Sforza, allows for nuanced interpretations of archaeological and genetic data.^[1]

Historical Development

Origins of the Concept

The concept of demic diffusion emerged in the early 1970s as a model to explain the spread of agriculture and associated population movements in prehistoric Europe, first articulated by Albert J. Ammerman and Luigi Luca Cavalli-Sforza in their population genetics research.^[16] In their 1971 paper, they estimated the rate of Neolithic spread across Europe at approximately 1 km per year using a wave-of-advance mathematical framework based on archaeological site dates.^[9] This was expanded in a seminal 1973 book chapter, where they adapted mathematical diffusion equations to simulate how farming communities could expand demographically, leading to gradual population replacement rather than mere idea transmission.^[17] This framework built directly on earlier ecological models, particularly Ronald A. Fisher's 1937 theory of the "wave of advance" for gene propagation in populations, which described how advantageous traits spread spatially through migration and selection at a constant rate.^[2] Ammerman and Cavalli-Sforza modified this to human demography, positing that increased population densities from agricultural innovations drove successive generations to migrate into adjacent territories, creating predictable expansion fronts.^[7] A key expansion of these ideas appeared in their 1984 book, The Neolithic Transition and the Genetics of Populations in Europe, which integrated archaeological data with genetic simulations to quantify demic processes over millennia. Here, the model predicted an advance rate of about 1 km per year for Neolithic farming frontiers, aligning with radiocarbon-dated site distributions across the continent.^[18] This work formalized demic diffusion as a mechanism distinct from cultural exchange, emphasizing gene flow via human mobility. The theoretical foundations gained empirical traction through genetic analyses, notably in a 1978 study by Paolo Menozzi, Alberto Piazza, and Cavalli-Sforza, which constructed "synthetic maps" of allele frequencies across European populations. These maps revealed smooth clines—gradual gradients—in genetic markers, such as blood group variants, radiating from the southeast (Near East origins) toward northern and western Europe, consistent with demic waves of migrating farmers rather than localized adaptations. Developed amid debates on whether agriculture spread primarily through migration or imitation, this approach provided a quantitative tool to test hypotheses about prehistoric population dynamics, influencing subsequent interdisciplinary research in anthropology and genetics.^[19]

Key Contributors

Luigi Luca Cavalli-Sforza, an Italian geneticist, pioneered the demic diffusion model through his integration of population genetics, archaeology, and linguistics to explain the spread of Neolithic farming and associated genetic patterns across Europe. In collaboration with archaeologist Albert Ammerman, Cavalli-Sforza developed the foundational "wave of advance" model in their 1971 paper and 1973 book chapter, which mathematically described the gradual expansion of farming populations as a demic process rather than mere cultural transmission.^[20] This work formalized demic diffusion as a hypothesis testable via genetic and archaeological data, emphasizing population movements from the Near East.^[6] Cavalli-Sforza further advanced the model in subsequent publications, including the 1978 study with Paolo Menozzi and Alberto Piazza, which used synthetic genetic maps of European populations to support demic expansion patterns aligned with Neolithic migrations. Their seminal 1994 book, The History and Geography of Human Genes, synthesized global genetic data to argue that demic diffusion accounted for major clines in human genetic variation, particularly in Europe, linking it to prehistoric population dispersals. Paolo Menozzi, an Italian geneticist, contributed significantly as a co-author in these efforts, providing statistical analyses that reinforced the genetic evidence for demic processes in human prehistory. His work with Cavalli-Sforza and Piazza helped establish demic diffusion as a cornerstone of interdisciplinary human evolutionary studies. Archaeologist Colin Renfrew extended the model's application by linking demic diffusion of early farmers to the spread of Indo-European languages in his 1987 book Archaeology and Language: The Puzzle of Indo-European Origins, proposing an Anatolian origin for these languages via population movements around 7000 BCE. Renfrew's synthesis of archaeological evidence with the demic framework influenced debates on linguistic and cultural dispersals, building on the genetic foundations laid by Cavalli-Sforza and colleagues.^[21] Together, these contributors' interdisciplinary approach transformed demic diffusion from a speculative idea into a rigorously supported model, fostering ongoing research in population history.^[17]

Theoretical Mechanisms

Population Dynamics

Demic diffusion involves the gradual expansion of populations through repeated short-range migrations, typically spanning 15-30 km per generation, which collectively propagate a wavefront of settlement into unoccupied or sparsely populated territories. This process relies on local movements by small family groups or communities rather than long-distance leaps, enabling the steady advancement of demographic frontiers over centuries. Such dynamics were first modeled for the Neolithic spread of farming in Europe, where iterative migrations from established settlements created a continuous gradient of population density.^[22] The driving force behind this expansion lies in differential population growth rates, particularly the elevated fertility among migrant groups like early farmers, who benefited from resource-rich agricultural practices that supported larger family sizes compared to indigenous hunter-gatherers. These higher growth rates—estimated at 0.01-0.04 per year in various models for Neolithic farmers—allowed migrants to outcompete locals demographically, leading to genetic admixture through intermarriage or, in some cases, partial population replacement as expanding groups filled new ecological niches. This reproductive advantage created a self-reinforcing cycle, where increased densities in core areas prompted further outward dispersal.^[2] To simulate these processes, researchers employ stochastic reaction-diffusion models that account for random variations in migration and growth, predicting the wavefront's advance velocity as v = 2 \sqrt{D r}, where v represents the speed of expansion, D is the diffusion coefficient (derived from migration distance and rate, often calibrated to 1-10 km² per generation), and r is the intrinsic growth rate. These models demonstrate how even modest parameters—such as a 5 km standard deviation in dispersal and around 2% annual growth—yield observed Neolithic spread rates of approximately 1 km per year, highlighting the model's robustness in replicating archaeological timelines without invoking mass relocations.^[3] Threshold effects further constrain sustainable expansion, requiring a minimum number of migrants (often tens to hundreds per settlement) to establish viable populations beyond which stochastic extinction risks diminish, modulated by the local environmental carrying capacity that limits density before further dispersal occurs. In low-capacity regions, insufficient initial colonists can halt wavefront propagation, emphasizing the interplay between demographic viability and habitat suitability in demic processes.^[12]

Genetic and Demographic Processes

In demic diffusion, gene flow occurs as expanding populations introduce novel alleles into recipient gene pools, resulting in clinal gradients of genetic variation along the direction of migration. This process dilutes the source population's genetic signature over distance due to recurrent admixture with local groups, producing smooth transitions in allele frequencies rather than sharp boundaries. For instance, theoretical models predict a gradual westward decrease in Near Eastern-derived ancestry across Europe during the Neolithic expansion, reflecting the progressive integration of migrant farmers' genes into indigenous hunter-gatherer populations. Uniparental markers such as Y-chromosome and mitochondrial DNA (mtDNA) often exhibit signatures of demic diffusion influenced by sex-biased migration patterns. Patrilineal inheritance via the Y-chromosome facilitates the spread of male-mediated haplogroups from the source population, leading to elevated Y-haplogroup diversity and frequency clines in the expansion direction, particularly in cases of male-biased dispersal. In contrast, mtDNA patterns may show less pronounced changes if female migration rates are lower, highlighting how demographic asymmetries shape uniparental genetic legacies.^[1] Admixture in demic diffusion is modeled through hybrid zones where incoming migrants interbreed with residents, resulting in overall contributions of 20-50% incoming ancestry in simulations of population wavefronts. These zones act as buffers, blending gene pools and stabilizing the expansion front while generating intermediate genetic profiles. Such models emphasize partial replacement rather than complete displacement, allowing for sustained gene flow that attenuates with distance from the origin.^[23] Coalescent theory provides a framework to connect demographic processes in demic diffusion to observable genetic patterns, enabling estimates of migration timing and rates from allele frequency distributions. By tracing lineages backward to common ancestors, it accounts for how serial founder effects and admixture alter coalescence times, offering insights into the pace and scale of population movements without relying on direct archaeological data. This approach has been applied to reconstruct the temporal dynamics of expansions, linking effective population sizes and migration events to genetic diversity gradients.^[24]^[1]

Supporting Evidence

Archaeological Findings

Archaeological evidence for demic diffusion in the Neolithic expansion of Europe is prominently illustrated by the linear distribution of early farming sites, pottery styles, and tools originating from Anatolia and spreading westward. Sites associated with the Linearbandkeramik (LBK) culture, dating to approximately 5500–4500 BCE, reveal a pattern of farming villages that emerged rapidly across Central Europe, with characteristic incised pottery and ground stone tools appearing in a wavefront-like progression from southeastern origins toward the Rhine and Danube regions.^[25] This distribution aligns with demic processes, as the uniformity in material culture suggests population movements rather than sporadic local inventions.^[26] Strontium isotope analysis of tooth enamel from LBK skeletons provides direct evidence of migration among early farmers in Central Europe. Examinations of remains from sites like Vedrovice and Nitra, for instance, show elevated strontium ratios (87Sr/86Sr) indicating that up to 20–30% of individuals originated from non-local areas, likely from the southeast, supporting the influx of farming populations into hunter-gatherer territories.^[27] Further studies confirm greater mobility at the onset of farming, with isotope signatures in western European sites revealing that early Neolithic communities incorporated migrants who brought agricultural practices over distances of hundreds of kilometers.^[28] Settlement patterns of the LBK further corroborate demic diffusion through the establishment of dense, successive villages along fertile river valleys, such as the Danube and Elbe, rather than scattered or isolated adopters of farming. These nucleated settlements, often comprising longhouses and enclosures, exhibit a clustered layout that advanced steadily, consistent with population-led wavefront migration where groups relocated en masse to exploit new arable lands.^[29] Radiocarbon dating of LBK sites demonstrates a uniform temporal progression, with calibrated dates indicating a spread rate of approximately 1 km per year from the Balkans to Central Europe, matching predictions from demic models of population dispersal.^[25]

Genetic Studies

Genetic studies have provided compelling molecular evidence for demic diffusion through the analysis of ancient and modern DNA, revealing substantial population replacements associated with the spread of farming practices. A landmark study by Haak et al. in 2015 sequenced genomes from 69 ancient Europeans spanning 8,000 to 3,000 years ago, demonstrating that Early European Farmers (EEF) carried ancestry primarily derived from Neolithic populations in Anatolia. This EEF component replaced approximately 75% of the Western Hunter-Gatherer (WHG) genomes in Central and Northern Europe during the Neolithic transition, indicating large-scale migration rather than solely cultural adoption of agriculture.^[30] Y-chromosome haplogroup data further supports demic diffusion by tracing male-mediated population movements. Semino et al. in 2000 analyzed 22 binary markers on the non-recombining Y chromosome from over 1,000 European males, identifying haplogroup G (specifically G2a) as originating from the Near East and correlating strongly with the geographic distribution of Neolithic archaeological sites across Europe. The spread of G2a lineages aligns with the timing and routes of farming expansion, suggesting that male farmers migrated in groups sufficient to establish these genetic signatures in recipient populations.^[31] Autosomal DNA analyses have elucidated admixture patterns and continuity in ancestry clines. Lazaridis et al. in 2014 sequenced high-coverage genomes from a 7,000-year-old farmer and eight 8,000-year-old hunter-gatherers, modeling present-day Europeans as deriving from three ancestral sources: WHG, EEF from Anatolia, and Ancient North Eurasians. Their analysis revealed smooth clines in EEF ancestry proportions decreasing from southeast to northwest Europe, consistent with demic diffusion involving ongoing gene flow and admixture gradients during the Neolithic.^[32] Recent studies extend this evidence to other regions, confirming demic patterns in non-European contexts. Tao et al. in 2023 reported genome-wide data from 11 ancient individuals in southwest China dating 4,500–3,000 years ago, showing that millet and rice mixed-farming populations derived approximately 90% of their ancestry from Neolithic Yellow River farmers. This substantial genetic contribution underscores demic diffusion as the primary mechanism for the spread of millet farming from the Yellow River Basin into southern regions.^[33]

Applications and Examples

Neolithic Expansion in Europe

The spread of Neolithic farming practices from the Near East to Europe exemplifies demic diffusion, where migrating agricultural populations played a primary role in disseminating farming technologies and lifestyles. This process originated around 9000 BCE in the Fertile Crescent, where early domestication of plants and animals enabled population growth and expansion.^[34] By approximately 6500 BCE, farming communities had reached the Balkans through initial migrations across Anatolia, marking the entry point into continental Europe.^[4] The expansion continued westward and northward, arriving in Britain by around 4000 BCE, facilitated by a steady "wave of advance" at rates of about 1 km per year.^[17] The primary migration routes included the Danube corridor, which channeled farmers northward into central Europe, and the Mediterranean arcs, allowing coastal dispersal along southern Europe.^[7] Models integrating archaeological and genetic data indicate that demic diffusion accounted for approximately 60% of the spread, with farmers' migrations outpacing cultural transmission through local adoption.^[17] These routes supported rapid population influxes, as evidenced by higher dispersal speeds—up to five times faster along the Danube-Rhine pathway and ten times along the Mediterranean—compared to purely cultural models.^[4] Cultural markers of this demic expansion prominently featured the introduction of domesticated wheat species, such as einkorn and emmer, alongside cattle herding, which transformed subsistence economies from foraging to agriculture.^[4] Sedentism emerged as a key outcome, with permanent settlements arising from the reliable food surplus provided by these innovations, enabling larger communities and social complexity in regions like the Balkans and central Europe.^[4] The demic waves contributed to the formation of megalithic cultures in western and northern Europe around 5000–3000 BCE, where farming descendants constructed monumental tombs reflecting organized societies tied to agricultural prosperity.^[35] Additionally, these migrations potentially influenced language shifts, as agricultural languages dispersed alongside the farming populations, shaping early European linguistic diversity.^[36]

Spread in Other Regions

In East Asia, demic diffusion is evidenced by the spread of millet farming originating from the Yellow River Basin around 8000 BCE, where ancient genomic data from sites in southwest China indicate that local populations derived approximately 90% of their ancestry from Neolithic Yellow River farmers, signifying substantial population replacement and migration-driven agricultural dissemination.^[37] This process involved successive waves of farmer migrations into regions previously dominated by hunter-gatherers, facilitating the integration of millet cultivation across northern and central areas.^[38] In South Asia, genetic analyses reveal Iranian farmer-related admixture in the Indus Valley region dating to approximately 5400–3700 BCE, marking an early instance of demic diffusion as herding and farming populations from the Iranian plateau migrated eastward, contributing significantly to the ancestry of later Indus Valley inhabitants without Steppe pastoralist input at that stage.^[39] This admixture, estimated to have occurred after 7000–6000 BCE, underscores a demographic expansion that blended incoming farmer lineages with local Ancient Ancestral South Indian hunter-gatherer groups, laying the genetic foundation for subsequent Bronze Age developments.^[40] Beyond continental Asia, demic diffusion manifests in oceanic contexts through Austronesian migrations starting around 3000 BCE, where voyaging populations from Taiwan spread rice farming to Pacific islands, replacing or admixing with Paleolithic settlers in Island Southeast Asia and beyond via total or near-total demographic shifts.^[41] Analogous to the Bantu expansion in Africa—where genetic evidence shows a demic spread of farming and ironworking from West Africa around 3000 BCE, resulting in cline-like diversity gradients and substantial ancestry contributions in eastern and southern regions—this pattern highlights how maritime and terrestrial migrations propelled agricultural frontiers in the Americas and Oceania, though at varying scales influenced by environmental barriers.^[42] Across these global cases, expansion speeds consistently range from 1 to 3 km per year in terrestrial or coastal suitable environments, mirroring the benchmark observed in the European Neolithic.^[43]

Criticisms and Modern Perspectives

Limitations of the Model

The pure demic diffusion model, which posits the spread of farming primarily through population migration and admixture with local groups, has been critiqued for overemphasizing demographic expansion at the expense of local adaptations and cultural transmission mechanisms. In regions such as parts of Central Europe, archaeological evidence indicates that indigenous hunter-gatherers adopted agricultural practices without complete population turnover, suggesting that cultural diffusion played a significant role alongside migration. For instance, studies integrating radiocarbon dates estimate that cultural diffusion accounted for approximately 40% of the Neolithic transition's pace, highlighting the model's failure to fully capture hybrid processes of adoption.^[17] A key theoretical weakness lies in the model's predicted expansion speed, which assumes a uniform rate of about 1 km per year based on population growth and dispersal parameters. However, empirical observations reveal discrepancies, with faster spreads observed in fertile river valleys and coastal areas, where rates exceeded 2 km per year due to favorable conditions accelerating settlement. This uniform speed assumption oversimplifies regional variations, leading to mismatches between modeled and actual timelines in heterogeneous landscapes.^[44] Early formulations of the demic diffusion model, developed before the widespread availability of ancient DNA data in the 1990s and 2000s, relied heavily on modern genetic patterns and archaeological proxies, resulting in assumptions of substantial farmer genetic contributions that have since been revised. Ancient DNA analyses from sites across Europe have demonstrated substantial genetic continuity with pre-Neolithic populations, indicating higher levels of admixture than initially proposed, thus underscoring data limitations in pre-genomic era models.^[45] Furthermore, the model often oversimplifies environmental influences by treating landscapes as homogeneous, neglecting how climate barriers and ecological constraints could halt or redirect migration waves. For example, in northern and mountainous regions of Europe, cooler climates and terrain features impeded the predicted steady advance, as evidenced by delayed Neolithic arrivals correlated with environmental suitability rather than solely demographic pressures. This environmental oversight limits the model's applicability to diverse prehistoric settings.^[46]

Integration with Cultural Diffusion

Hybrid models of demic and cultural diffusion integrate population migration with the transmission of ideas and practices, providing a more nuanced explanation for the spread of innovations like agriculture. In a seminal 2012 study, researchers developed a unified framework using reaction-diffusion equations to combine these processes, estimating that cultural diffusion accounted for approximately 40% of the Neolithic transition's spread rate in Europe, with demic diffusion contributing the remaining 60%.^[17] This mixed approach reconciles archaeological evidence of variable regional rates, where demic movement filled gaps left by uneven cultural adoption.^[17] Recent simulations have advanced these hybrid frameworks by incorporating complex geographic and demographic factors. A 2025 agent-based modeling study of the European Neolithic expansion revealed an initial demic core of high early farmer ancestry through migration, followed by a limited cultural halo where hunter-gatherers adopted farming practices at low rates (about 0.1% per year).^[4] Similarly, a 2024 analysis using dispersal kernels estimated cultural contributions at 13-25% in Europe, emphasizing demic dominance but highlighting interplay in slower spreads.^[47] These models demonstrate how combined mechanisms better account for patchy archaeological distributions than pure demic scenarios, as cultural transmission allows for localized accelerations without uniform population replacement.^[17] The benefits of such integrations include improved fits to empirical data on ancestry clines and radiocarbon dates, revealing non-uniform expansion routes like northern and southern paths in Europe.^[4] Future directions involve expanding agent-based simulations to quantify proportions across diverse regions, incorporating ancient DNA and environmental variables for more precise delineations of demic versus cultural roles.^[4]

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We argue that the historical processes behind the Neolithization of Europe were influenced by environmental factors predisposing occupation of regions.
[42]
Cultural versus demic diffusion in agricultural expansions according ...
Feb 19, 2024 · Mathematical models of agricultural spread use distances between birthplaces of parents and their children (often called “birthplace ...