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Cosmopolitan distribution

In , a cosmopolitan distribution refers to the geographic range of a —such as a , , or —that extends across most or all of the Earth's continents or oceans in suitable habitats, often spanning multiple biogeographic realms. This pattern contrasts with more restricted distributions, like , and is typically observed in organisms capable of long-distance dispersal, broad environmental tolerance, or human-mediated transport. Notable examples include the rock pigeon (Columba livia), native to Eurasia but now established on all continents except Antarctica due to human introduction, and the house mouse (Mus musculus), which has achieved worldwide presence through commensalism with human settlements. Other cosmopolitan taxa encompass microorganisms like certain diatoms (Hantzschia amphioxys) and tardigrades, as well as plants such as common reed (Phragmites australis), which thrive in diverse wetlands globally. These distributions often result from evolutionary adaptations, natural dispersal mechanisms (e.g., wind or ocean currents), or anthropogenic factors, enabling persistence in varied climates from tropical to temperate zones. However, modern genetic analyses have challenged the notion of true cosmopolitanism, revealing that many historically described cosmopolitan species are actually complexes of cryptic lineages with more localized ranges, differentiated by subtle morphological or molecular traits. For instance, in environments, apparent global uniformity may mask regional driven by isolation or adaptation. This complexity underscores the role of in understanding patterns, dispersal barriers, and responses to .

Definition and Qualification

Core Definition

In , a cosmopolitan distribution describes the geographic of a —such as a , , or higher group—that spans most or all of Earth's surface in ecologically suitable habitats, often encompassing multiple continents, oceans, and biomes while typically excluding extreme polar regions or highly isolated realms unless the taxon possesses adaptations for those environments. This pattern implies a high degree of dispersal success and ecological tolerance, allowing the taxon to maintain viable populations across vast, heterogeneous landscapes without being confined to specific biogeographic provinces. The concept of cosmopolitan distribution emerged in the 19th century amid efforts by naturalists to map global species patterns and challenge earlier notions of fixed regional boundaries. Edward Forbes, a pioneering marine biologist, contributed significantly through his 1859 monograph The Natural History of the European Seas, where he delineated zoogeographic regions in the ocean and highlighted species exhibiting apparent uniformity across wide marine expanses, laying groundwork for understanding global-scale uniformity in distributions. This historical framing underscored the idea of certain taxa achieving near-worldwide presence due to effective long-distance dispersal and broad environmental adaptability, influencing subsequent biogeographic theory. Unlike merely wide-ranging species, which may occupy large but discontinuous or regionally restricted areas (e.g., across a single or major ), cosmopolitan distributions demand a more pervasive, quasi-universal occupancy that transcends continental and oceanic barriers in habitable zones. Qualification as cosmopolitan typically requires evidence of occurrence in at least three major biogeographic realms, with genetic and ecological continuity, though precise criteria are elaborated further in related assessments and vary by and study.

Qualification Criteria

To qualify as having a cosmopolitan distribution, a must demonstrate presence across multiple major biogeographic realms, such as the traditional Wallacean divisions (Nearctic, Palearctic, Neotropical, Ethiopian, Oriental, and ), reflecting broad global occupancy beyond regional confines. More recent analyses refine this using an updated of 11 zoogeographic realms, with some studies classifying as cosmopolitan if they occur in 9 or 10 of these realms, based on distributional and phylogenetic data from over 21,000 vertebrate . This criterion emphasizes not just geographic span but also ecological viability across diverse environments; however, there is no universally standardized threshold, and assessments often consider context-specific factors. A key additional standard involves limited absence from suitable habitats to distinguish true from patchy or human-mediated introductions. Quantitative assessments often incorporate thresholds drawn from ecological modeling to ensure distributions are not relics of transient events. Exceptions apply for habitat-specific taxa, where barriers are evaluated relative to life history; truly , such as certain copepods, are deemed cosmopolitan if distributed across connected marine or freshwater systems, disregarding terrestrial divides like continents. Similarly, aerial dispersers like seabirds ignore ground-based obstacles, qualifying based on flight-enabled access to suitable realms. Modern phylogeographic studies further validate cosmopolitan status through evidence of genetic connectivity, requiring low inter-regional differentiation indicative of ongoing , as seen in analyses of widespread where haplotypes show minimal divergence across hemispheres. This genetic criterion, often derived from multi-locus sequencing, confirms that apparent reflects natural dispersal rather than cryptic or isolation.

Contrasting Distributions

represents a stark contrast to cosmopolitan distribution, characterized by the restriction of a to a single locality, region, or , where it occurs nowhere else on . This pattern emphasizes localized evolutionary and , often resulting in high hotspots like islands or mountain ranges. For instance, (Geospiza spp.), confined exclusively to the , exemplify endemism, with their 13 adapted to specific niches within this archipelago, highlighting the opposite of the broad, continuous spread seen in cosmopolitan taxa. In contrast, like the rock pigeon (Columba livia), which thrive across urban and rural environments on every continent, demonstrate the pervasive global presence that defines cosmopolitanism. Intermediate distribution patterns, such as Holarctic or , occupy a middle ground between and full , spanning large but not worldwide areas. The Holarctic distribution encompasses nontropical regions of the , including , northern , and , but excludes southern continents and equatorial zones, thus falling short of global coverage. Similarly, distributions are limited to tropical latitudes across both Eastern and Western Hemispheres, often involving weedy or adaptable species that do not extend into temperate or polar realms. These patterns illustrate regional breadth without the ubiquity required for cosmopolitan status, as per qualification criteria that demand presence in most or all suitable habitats worldwide. Disjunct distributions further differentiate from cosmopolitan patterns by featuring wide-ranging but fragmented occurrences, where populations of the same species are separated by significant geographic gaps without continuous occupation in between. This discontinuity often arises from historical vicariance or incomplete dispersal, resulting in isolated pockets rather than the seamless, near-universal coverage of s. For example, certain plant genera like exhibit disjunct ranges in southern , , and , separated by vast oceanic barriers, underscoring the gapped nature that precludes cosmopolitan classification. Such patterns highlight the importance of evaluating range continuity when distinguishing distribution types.

Biogeographic Frameworks

Biogeographic frameworks provide the foundational structures for understanding how species distributions, including those of cosmopolitan taxa, relate to Earth's major ecological divisions. Alfred Russel Wallace's seminal classification divided the terrestrial world into six primary zoogeographic realms based on patterns of animal distribution: the Palearctic (encompassing , North Asia, and ), Nearctic ( north of ), Neotropical (Central and ), Ethiopian (), Oriental (South and ), and Australian (, , and surrounding islands). These realms reflect historical barriers to dispersal and evolutionary divergence, yet cosmopolitan species are characterized by their ability to occur across multiple realms, often due to exceptional adaptability or ancient origins predating the full establishment of these boundaries. Modern refinements to Wallace's system, such as Miklos Udvardy's 1975 classification, expanded the framework to eight realms—adding the and Oceanian realms—and further subdivided them into 193 biogeographic provinces to account for finer-scale and transitional zones. Udvardy's approach emphasizes biotic provinces where species assemblages show high regional fidelity, but it also highlights how cosmopolitan distributions manifest as widespread occurrences that overlap and transcend these provincial boundaries, integrating diverse ecosystems from temperate to tropical zones. Maps of these realms often depict overlaps in transitional areas, such as the Holarctic bridge between the Palearctic and Nearctic, where cosmopolitan lineages can exploit connectivity via land bridges or climatic corridors. In marine environments, frameworks like the Marine Ecoregions of the World () refine provincial divisions to include 12 major realms and 62 provinces, such as the Indo-West Pacific and Temperate Austral, based on patterns in and distributions. These marine provinces account for oceanic currents and depth gradients, with cosmopolitan marine species demonstrating trans-provincial ranges that challenge strict delineations, as seen in overlaps between tropical and temperate zones. Diagrams of these frameworks typically illustrate realm boundaries as fluid, with cosmopolitan distributions bridging gaps like the convergence zone. The historical development of these frameworks is deeply intertwined with and geological processes, which have reshaped continental configurations and facilitated or restricted ancient lineages. During the era, tectonic movements—such as the breakup of and the closure of the Tethys Sea—created vicariance events that isolated populations, yet some lineages achieved status by predating these shifts or through subsequent long-distance dispersal across evolving landmasses. This geological underscores how frameworks like Wallace's and Udvardy's evolved to incorporate paleogeographic evidence, explaining the persistence of patterns in lineages originating from supercontinents. Contrasting distributions, such as endemic patterns, emerge as subsets within these tectonic-influenced realms.

Dispersal Mechanisms and Barriers

Modes of Dispersal

Cosmopolitan species achieve their widespread distributions through a combination of active and passive dispersal modes, which enable colonization across diverse habitats. Active dispersal relies on the organism's own locomotion, such as long-distance via flight in like the barn (Hirundo rustica), which annually traverses hemispheres to exploit seasonal resources. Similarly, swarming, as seen in species like the painted lady butterfly (Vanessa cardui), facilitates rapid spread over thousands of kilometers during favorable winds and weather patterns. These mechanisms are particularly effective in mobile taxa, allowing repeated range expansions without dependence on external carriers. Passive dispersal, in contrast, depends on abiotic or vectors to transport propagules over vast distances. currents propel lightweight structures like or in , while water flows carry aquatic larvae or floating diaspores in species, such as the cosmopolitan oligochaete Pontodrilus litoralis. also plays a key role, with organisms like epiphytic microbes or attaching to migratory hosts, including or mammals, to bridge geographic gaps. Long-distance mechanisms further underpin cosmopolitan ranges, including rafting on natural debris like or mats, which has historically enabled transoceanic crossings in and . Aerial spores in fungi and , dispersed by , exemplify passive long-range transport, with evidence from phylogeographic patterns indicating global connectivity. records, such as those of early angiosperms, provide historical corroboration, revealing dispersal events predating modern continents and supporting the role of these processes in ancient biogeographic patterns. Genetic adaptations complement these dispersal modes by enhancing survival and . High , producing vast numbers of propagules, boosts the probability of successful , as observed in invasive where elevated reproductive output correlates with rapid spread. Dormant stages, such as resilient cysts in microscopic aquatic animals or spores in microbes, allow endurance during prolonged transport, facilitating passive dispersal across inhospitable environments. Studies of , using markers like microsatellites in species such as the common reed (), reveal sustained connectivity among distant populations, preserving genetic diversity essential for persistence.

Major Obstacles

Cosmopolitan species face formidable oceanic barriers that originated from the vicariance events associated with the breakup of the supercontinent during the late era, particularly the formation of and Oceans, which isolated southern landmasses and disrupted previously continuous distributions. These expansive water bodies, including the Pacific Ocean's vast width, have historically prevented passive dispersal for many terrestrial organisms, leading to in affected lineages while requiring exceptional long-distance colonization events for species to achieve global ranges. For instance, in neobatrachian frogs, phylogenetic analyses indicate that early divergences align with the opening of these oceans around 100-80 million years ago, highlighting how such barriers shaped disjunct patterns that cosmopolitan taxa later overcame through rare transoceanic dispersals. On land, mountain ranges and deserts present significant terrestrial obstacles to the spread of species, with varying permeability influenced by and species traits. The Himalayan , rising over the past 50 million years, has acted as a directional barrier to between the and the , reducing migration for many taxa due to extreme elevations exceeding 8,000 meters and harsh altitudinal gradients that limit suitable habitats. Case studies of Himalayan diatoms reveal that while some species maintain distributions through aquatic dispersal in streams that traverse the range, others show restricted , demonstrating partial barrier permeability for microorganisms but greater impermeability for larger organisms. Similarly, hyper-arid deserts like the have long isolated from , with and phylogenetic evidence indicating low faunal exchange rates over millions of years, though insects and birds occasionally permeate these zones via wind-assisted or migratory pathways. Climatic gradients further challenge cosmopolitan distributions by exerting selective pressures that demand broad physiological tolerance, often resulting in pre-human isolation patterns within fragmented habitats. Steep transitions from tropical to temperate or arid zones create physiological barriers, where species must adapt to varying regimes, levels, and seasonal extremes to persist across continents. For example, in the reed Phragmites australis, ecophysiological studies along latitudinal gradients show that plasticity in water use efficiency and cold tolerance enables survival, but historical climatic oscillations, such as those during the Pleistocene, fragmented habitats into isolated refugia, imposing genetic bottlenecks and local adaptations that tested the limits of widespread occupancy. These natural fragmentations, driven by glacial-interglacial cycles rather than anthropogenic factors, underscore how climatic variability has historically constrained and favored resilient traits in globally distributed species.

Ecological and Population Dynamics

Ecological Boundaries

Ecological boundaries of cosmopolitan distributions are primarily shaped by the interplay between niche conservatism—the tendency for species to retain ancestral ecological tolerances—and , which allows adaptation to varying environmental conditions across biomes. Niche conservatism often constrains cosmopolitan to similar abiotic conditions, such as and ranges, limiting their effective range despite widespread dispersal. For instance, in marine copepods, populations exhibit conserved thermal tolerances but varying plasticity in salinity responses, enabling persistence in diverse coastal habitats while preventing full exploitation of extreme conditions. In contrast, high plasticity in pH tolerance facilitates broader distributions in fluctuating oceanic environments, as observed in cosmopolitan invertebrates where overlapping pH niches across habitats reduce sensitivity to acidification gradients. This balance determines whether a maintains a truly global presence or faces ecological limits in novel biomes. Habitat suitability models (HSMs) provide a framework for delineating these boundaries by integrating abiotic factors like , , , and UV radiation with interactions such as predation. HSMs, rooted in theory, predict occurrence probabilities based on environmental envelopes, revealing how UV radiation acts as a barrier for organisms by damaging DNA unless mitigated by protective mechanisms. factors, including predation avoidance, further refine suitability at local scales; for example, models incorporating predator distributions show reduced habitat occupancy in high-predation zones, even where abiotic conditions are favorable. These models emphasize that while abiotic tolerances define broad potential ranges, pressures impose finer-scale limits, often resulting in fragmented distributions within cosmopolitan patterns. The distinction between fundamental and realized niches critically delimits cosmopolitan ranges, with many species exhibiting pseudo-cosmopolitan distributions due to unfilled portions of their fundamental niche. The fundamental niche encompasses all conditions under which a species can survive absent interactions, often broader than the realized niche, which reflects actual occupancy constrained by , predation, and other factors. In pseudo-cosmopolitan cases, widespread dispersal creates an illusion of ubiquity, but ecological barriers leave potential habitats unoccupied, such as in marine species where larval transport reaches distant sites yet adult survival is limited by unsuitable conditions. This gap highlights how physical dispersal mechanisms contribute to apparent boundaries without fully realizing the species' physiological potential.

Regional and Temporal Variations

Cosmopolitan species exhibit genetic clines along environmental gradients, reflecting adaptation to varying climatic conditions across their global ranges. In the fruit fly , a classic example of a cosmopolitan , mitochondrial and nuclear DNA analyses reveal latitudinal clines in frequencies, with variants associated with resistance more prevalent in arid subtropical regions and those linked to cold tolerance increasing toward higher latitudes. These clines arise from ongoing selection pressures despite high , maintaining subtle genetic across continents. Similarly, in the cosmopolitan reed , genome-wide studies show clinal variation in traits like flowering time and stress tolerance, correlating with and gradients from tropics to temperate zones. Isolation during Pleistocene glaciations has contributed to formation in several cosmopolitan species by fragmenting populations into refugia, promoting . For the barn swallow Hirundo rustica, phylogeographic analyses of identify six across , , and , with deep divergences dated to the (approximately 0.5–1 million years ago), likely resulting from glacial isolation in southern refugia followed by northward recolonization. In the osprey Pandion haliaetus, another widely distributed raptor, mitochondrial sequencing reveals four major genetic lineages corresponding to Australasian, , Palearctic, and Nearctic groups, with splits attributed to vicariance during Pleistocene sea-level fluctuations and continental glaciations that restricted . Quaternary climatic cycles have driven temporal shifts in the distributions of cosmopolitan species, with paleontological evidence documenting range contractions during glacial periods and expansions during interglacials. These oscillations, repeated over multiple cycles, have left signatures in the , showing cyclic latitudinal migrations tied to variations of 4–6°C. In avian examples like the barn swallow, genetic data confirm population bottlenecks during glacial advances, with subsequent demographic expansions traced through coalescent models. Demographic variations in cosmopolitan species often manifest as higher densities and in tropical regions compared to polar areas, shaped by resource availability and historical stability. of Drosophila melanogaster demonstrate greater nucleotide diversity and lower coefficients in equatorial populations, reflecting larger effective population sizes (N_e > 10^6) versus reduced N_e (~10^5) at higher latitudes, where harsh conditions densities. (F_ST) metrics further quantify regional genetic differentiation in cosmopolitan species. These patterns provide baselines for understanding variations relative to ecological boundaries, such as thermal tolerances defining range s.

Notable Examples

Microorganisms and Invertebrates

Microorganisms exhibit cosmopolitan distributions due to their microscopic size, high reproductive rates, and ability to disperse passively through air, water, and biological vectors, enabling them to colonize diverse global environments. Bacteria such as are a prime example, serving as commensals in the intestines of humans and animals while persisting in external habitats like water and sediments across every continent, including . This ubiquity stems from their adaptation to varied ecological niches, from aquatic systems to soil, facilitating widespread nutrient cycling and occasionally opportunistic infections. Among zooplankton, species in the genus Daphnia, such as , demonstrate cosmopolitan patterns through their production of dormant resting eggs (ephippia) that disperse via water currents and wind, allowing establishment in freshwater bodies worldwide. These small cladocerans thrive in temperate and tropical lakes, contributing to aquatic food webs and responding to environmental cues like for predator avoidance. Their strong dispersal capabilities and large population sizes further underscore their global presence, often regarded as archetypal cosmopolitan freshwater . Fungi like those in the genus Aspergillus exemplify microbial cosmopolitanism, with species such as Aspergillus fumigatus and Aspergillus flavus distributed ubiquitously in soils, plant litter, and air worldwide, playing essential roles in decomposing organic matter and driving global nutrient cycling. As saprophytes, they break down complex carbon compounds, releasing nutrients that support ecosystem productivity across biomes, from arid deserts to humid forests. Their spore-based dispersal via atmospheric currents enhances this broad reach, making them key players in terrestrial biogeochemical processes. Invertebrates, particularly small arthropods, achieve cosmopolitan status through rapid reproduction, passive dispersal modes like wind and water, and opportunistic use. The (Musca domestica) is a notable case, with a worldwide distribution tied to its synanthropic lifestyle near human settlements, where it breeds in organic waste and spreads via flight and transport. Its high evolutionary adaptability, including quick development of resistance to environmental stresses, supports colonization of diverse climates from temperate to subtropical regions. Cockroaches, such as americana and Blattella germanica, represent resilient cosmopolitan , found in nearly all human-inhabited areas due to their tolerance for extreme conditions and omnivorous diet. These species exhibit rapid and cryptic behaviors that aid survival in urban and natural settings globally, from to temperate zones, often evolving traits like insecticide resistance to maintain their range. Their association with human activity subtly enhances dispersal without dominating their natural ecological spread.

Vertebrates and Plants

Among vertebrates, the (Passer domesticus) exemplifies a cosmopolitan species, native to and parts of but now distributed across all continents except due to introductions starting in the . Its range expansion is closely tied to human settlements, with post-glacial recolonization of around 10,000 years ago facilitating early associations with , followed by global dispersal via shipping and colonization. Similarly, rats of the genus , including the (R. norvegicus) and (R. rattus), have achieved near-global distributions, originating from —the former from northern and , the latter from the —and spreading worldwide through human-mediated transport on ships since the 1700s. These thrive in urban and rural human-altered environments, with evolutionary adaptations for enabling rapid population growth and establishment in diverse climates. In plants, the common dandelion () represents a classic cosmopolitan weed, native to (with origins traced to the approximately 30 million years ago) but now widespread across , , , and beyond as an introduced by . Its dispersal relies on wind via lightweight achenes equipped with a pappus for long-distance transport (up to 150 km or more), supplemented by animal adhesion and human activities like and air travel. Other cosmopolitan weeds, such as broadleaf cattail (), exhibit similar patterns, with wind, water, and animal-mediated dispersal contributing to their presence on every continent except . Evolutionarily, T. officinale's success stems from apomictic reproduction in triploid lineages, producing clonal seeds that maintain genetic uniformity while allowing hybridization with sexual diploids to generate diversity, alongside a short-lived that aids persistence in disturbed habitats despite lacking long-term dormancy. Regional variations in these species, such as denser urban populations, reflect local environmental adaptations without altering their overall global span.

Contemporary Influences

Anthropogenic Dispersal

Human activities, particularly since the , have dramatically accelerated the dispersal of , transforming many from regional endemics into cosmopolitan invaders by bypassing natural barriers through global trade networks. This facilitation has introduced thousands of non-native worldwide, often via unintentional vectors like shipping and , leading to homogenized biotas across continents. For instance, the volume of international shipping has increased exponentially, from about 2.6 billion tons in 1970 to over 11 billion tons by 2020, correlating with heightened invasion risks. Trade and transport mechanisms have been primary drivers of this expansion, with ship ballast water serving as a notorious conduit for aquatic species. Ballast water, used to stabilize vessels, discharges billions of liters of plankton-laden seawater daily into ports, enabling the transoceanic spread of organisms like the (Dreissena polymorpha). Native to the Ponto-Caspian region, zebra mussels were first detected in in the late 1980s, likely via ballast discharge from European freighters into the , and have since proliferated across freshwater systems in the United States and , fouling and disrupting ecosystems. By 2020, they had established populations in over 30 U.S. states, illustrating how maritime trade has rendered this species effectively cosmopolitan in temperate freshwater habitats. Similar patterns occur with air and land transport, where hitchhiking and accompany , further amplifying global distributions. Agriculture and urbanization have similarly propelled the cosmopolitanization of terrestrial species, especially weeds that thrive in disturbed habitats. Global crop trade and soil movement have disseminated "anthropochorous" plants—those dispersed by human action—forming cosmopolitan weed complexes in arable lands and cities. For example, common bermudagrass (), originally from and , was intentionally and accidentally spread via shipments and urban since the , achieving near-global coverage in warm-climate lawns, roadsides, and fields by the mid-20th century. Urban expansion exacerbates this by creating novel habitats like pavement cracks and waste grounds, where resilient weeds like , dispersed through contaminated seed and vehicle traffic, now span every continent except . These complexes not only compete with but also alter microbiomes, underscoring the intertwined role of farming intensification and city growth in species homogenization. Post-2000 data highlight the escalating impacts of these invasive cosmopolitans, with the International Union for Conservation of Nature (IUCN) and the IPBES assessment tracking over 37,000 established alien species globally, noting a surge in establishments linked to trade globalization. Economic costs have ballooned accordingly; a analysis estimated annual global damages from invasive aliens at $423 billion, with sectors like and fisheries bearing the brunt through yield losses and control efforts. In alone, invasive species inflicted $137 billion in yearly costs by 2021, including $5.4 billion from zebra mussels in infrastructure maintenance. These figures emphasize the need for enhanced , such as the IMO's ratified in 2017, to curb further anthropogenic spread.

Climate Change Impacts

Climate change is driving significant range shifts in cosmopolitan species, with observed poleward and upslope migrations documented across terrestrial, freshwater, and marine ecosystems. These shifts occur as warming temperatures alter thermal tolerances, prompting species to track suitable climates; for instance, marine ectotherms exhibit range expansions toward poles under moderate warming scenarios. Models from the project that by 2100, under high-emission pathways like SSP5-8.5, up to 50% of species could experience range contractions, while generalist cosmopolitan species may expand, leading to biotic homogenization where local decreases as similar widespread taxa dominate assemblages. Peripheral populations of species are particularly vulnerable to loss due to climate-induced at trailing edges, where warming exacerbates isolation and reduces . Studies indicate that trailing-edge populations often harbor unique adaptive alleles, but projected habitat losses could lead to and diminished evolutionary potential. Concurrently, facilitates the emergence of novel invasives by expanding suitable habitats for non-native species, such as tropical plants and microbes that previously faced barriers in temperate zones; for example, warming has enabled species like the reed Phragmites australis to adapt and invade new regions through enhanced . Recent research highlights accelerated spread of tropical pathogens as a key impact, with modeling showing that warming increases the global richness and abundance of like Vibrio species by over 20% by 2100, enabling their cosmopolitan distribution into higher latitudes and threatening human and . Conservation strategies emphasize maintaining connectivity through ecological corridors and expanding marine protected areas to safeguard cosmopolitan species like the cold-water coral Desmophyllum dianthus, where over 80% of habitats may persist under low-emission scenarios but require targeted interventions in high-risk regions such as the Patagonian Shelf. Additional approaches include genetic monitoring of peripheral populations and assisted migration to bolster adaptive diversity, prioritizing low-emission pathways to mitigate homogenization risks.

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