1 Introduction

The genus Gossypium, commonly known as cotton, comprises approximately 50 species distributed across tropical and subtropical regions worldwide, excluding Europe (Viot and Wendel, 2023). The evolutionary history of Gossypium is marked by multiple episodes of transoceanic dispersal, which have played a crucial role in the diversification and distribution of the genus. Notably, the genus originated around 5 to 10 million years ago and underwent rapid diversification into major genome groups shortly thereafter. The transoceanic dispersal events facilitated the spread of Gossypium species to new ecological niches, leading to the evolution of both diploid and allopolyploid species. For instance, the allopolyploid cottons, which include the commercially significant species G. hirsutum and G. barbadense, arose from the transoceanic dispersal of an A-genome taxon to the New World, followed by hybridization with an indigenous D-genome diploid.

Transoceanic dispersal has had profound implications for the evolution and biogeography of Gossypium . This process has not only facilitated the geographic expansion of the genus but also contributed to its genetic and morphological diversity. The long-distance dispersal events have led to the establishment of Gossypium species in diverse habitats, promoting speciation and adaptation to different environmental conditions (Grover et al., 2017; Grover et al., 2018).

Additionally, the repeated domestication of different wild progenitors in Africa-Asia and the Americas represents a remarkable case of human-driven parallel evolution (Wendel and Cronn, 2003; Wendel and Grover, 2015). The genomic consequences of these dispersal events include genome downsizing, interspecific hybridization, and the accumulation of deletions, which have further shaped the evolutionary trajectory of the genus.

The primary objective of this study is to elucidate the mechanisms and evolutionary significance of transoceanic dispersal in the genus Gossypium . Through integrating data from taxonomy, biogeography, molecular genetics, and phylogenetic analysis, this research aims to provide a comprehensive understanding of the role of transoceanic dispersal in the diversification and adaptation of cotton species.

2 Biological Characteristics of Gossypium

2.1 Plant morphology and ecological adaptability

Gossypium, commonly known as the cotton genus, exhibits remarkable morphological diversity, ranging from herbaceous perennials to trees as tall as 15 meters. This genus is characterized by a wide array of reproductive and vegetative traits, which have evolved to adapt to various ecological niches in arid to semi-arid regions of the tropics and subtropics (Wendel and Cronn, 2003; Wendel and Grover, 2015). The morphological variation within Gossypium includes differences in growth forms, leaf shapes, flower structures, and seed capsule sizes, which contribute to its adaptability and ecological success. The genus has also shown a high degree of cytogenetic and genomic diversity, which has facilitated its adaptation to different environments and contributed to its widespread distribution (Wendel et al., 2009).

2.2 Diversity and global distribution of Gossypium

Gossypium comprises over 50 recognized species distributed across tropical and subtropical regions worldwide, except Europe (Viot and Wendel, 2023). The genus is divided into eight major genome groups (A through G, and K), each representing a distinct lineage that has adapted to specific ecological conditions. The global distribution of Gossypium species is a result of multiple transoceanic dispersal events and subsequent diversification. Notably, four species have been independently domesticated for their fiber: two diploids from Africa-Asia and two allopolyploids from the Americas. This domestication has led to the development of commercially important cotton species such as G. hirsutum and G. barbadense, which are widely cultivated for their high-quality fibers (Grover et al., 2018).

2.3 Biological traits related to transoceanic dispersal

The evolutionary history of Gossypium is marked by several episodes of transoceanic dispersal, which have played a crucial role in the genus's diversification and global distribution. These long-distance dispersal events have facilitated the colonization of new ecological niches and the formation of new species through hybridization and polyploidization. For instance, the allopolyploid cottons, which include G. hirsutum and G. barbadense, originated from the hybridization of an A-genome taxon that dispersed to the New World with an indigenous D-genome diploid. This hybridization event, followed by genome doubling, has led to the emergence of new genetic combinations and traits that have enhanced the ecological adaptability and agronomic potential of these species (Huang et al., 2020). Additionally, the ability of Gossypium species to undergo natural interspecific hybridization, despite geographical isolation, has further contributed to their genetic diversity and evolutionary success.

3 Evolutionary History of Gossypium

3.1 Origin and evolutionary pathways of Gossypium

The genus Gossypium, commonly known as cotton, comprises approximately 50 species distributed across arid and semi-arid regions of the tropics and subtropics. The origin of Gossypium is estimated to be around 5 to 10 million years ago (mya), with a rapid diversification into eight major genome groups (A through G, and K) shortly thereafter (Wendel and Cronn, 2003; Wendel et al., 2009; Wendel and Grover, 2015). This diversification was driven by multiple episodes of transoceanic dispersal and natural hybridization among geographically isolated lineages. Notably, four species of Gossypium were independently domesticated for their fiber, two in Africa-Asia and two in the Americas, representing a remarkable case of parallel evolution driven by human selection (Wendel and Grover, 2015).

3.2 Insights from molecular phylogenetics on species relationships

Molecular phylogenetic studies have provided significant insights into the evolutionary relationships within Gossypium . Whole genome resequencing and comparative genomics have revealed that the subgenus Houzingenia, which includes New World diploid cottons, likely originated from a transoceanic dispersal event from Africa around 6.6 million years ago. Subsequent rapid diversification in the mid-Pleistocene (0.5~2.0 mya) led to the current biodiversity observed in this subgenus (Grover et al., 2018). Additionally, phylogenetic analyses have clarified the relationships among the eight genome groups and the origins of allopolyploid cottons, which arose from hybridization events between A-genome and D-genome diploids following transoceanic dispersal (Wendel et al., 2012).

3.3 Possible dispersal events in the course of evolution

The evolutionary history of Gossypium is marked by several key dispersal events. One of the most significant events was the transoceanic dispersal of an A-genome taxon to the New World, which occurred within the last 1 to 2 million years. This event led to the formation of allopolyploid cottons through hybridization with indigenous D-genome diploids (Wendel et al., 2012). These allopolyploids subsequently diversified into three modern lineages, including the commercially important species G. hirsutum and G. barbadense. Other notable dispersal events include the spread of New World diploid cottons to regions such as Arizona, the Galapagos Islands, and Peru, which required multiple long-distance dispersals (Grover et al., 2018). The biogeographic disjunction observed in the genera Kokia and Gossypioides, restricted to the Hawaiian Islands and Madagascar/East Africa respectively, also highlights the role of long-distance insular dispersal in the evolution of the cotton tribe (Grover et al., 2017).

4 Evolutionary Perspectives on Gossypium Dispersal

4.1 Genetic adaptations supporting dispersal and colonization

Gossypium species have exhibited remarkable genetic adaptations that facilitate their dispersal and colonization across diverse geographical regions. The genus has undergone significant genome downsizing, particularly in the genera Kokia and Gossypioides, which are closely related to Gossypium . This downsizing is characterized by a loss of approximately 30% of gene content and a genome-wide bias toward deletions over insertions, which may have contributed to their ability to colonize new environments (Grover et al., 2017). Additionally, the New World diploid cottons (Gossypium, subg. Houzingenia) have shown a bias toward deletions over small insertions, which helps maintain relatively small genome sizes despite the presence of repetitive DNA elements. These genetic adaptations likely play a crucial role in the ability of Gossypium species to disperse over long distances and establish populations in new habitats.

4.2 Phylogenetic evidence of transoceanic dispersal events in Gossypium

Phylogenetic analyses provide compelling evidence for multiple transoceanic dispersal events in the evolutionary history of Gossypium. The cotton tribe (Gossypieae) has experienced several long-distance dispersals, resulting in a wide geographic range that includes much of the tropics and subtropics (Grover et al., 2018). For instance, the New World diploid cottons likely originated from a transoceanic dispersal event from Africa approximately 6.6 million years ago, followed by rapid diversification in the mid-Pleistocene (Álvarez et al., 2005; Zhu et al., 2020). Furthermore, the allopolyploid cottons, which include the agronomically important species G. hirsutum and G. barbadense, emerged within the last 1 to 2 million years due to the transoceanic dispersal of an A genome taxon to the New World and subsequent hybridization with an indigenous D genome diploid. These phylogenetic insights underscore the significance of transoceanic dispersal in shaping the evolutionary trajectory of Gossypium.

4.3 Role of environmental factors in shaping genetic diversity across geographical regions

Environmental factors have played a pivotal role in shaping the genetic diversity of Gossypium species across different geographical regions. The distribution and genetic structure of Gossypium are influenced by interactions between long-distance dispersal mechanisms and geographical barriers. For example, the global distribution of mangroves in the genus Rhizophora, which shares similar dispersal mechanisms with Gossypium, is attributed to long-distance propagule dispersal and geographical barriers, leading to distinct genetic structuring between Pacific and Atlantic populations (Takayama et al., 2013). Similarly, the genetic diversity observed in Gossypium species is a result of both historical biogeographic events and contemporary environmental conditions. The repeated domestication of different wild progenitors in geographically isolated regions, such as Africa-Asia and the Americas, highlights the role of environmental factors in driving parallel evolution and genetic diversification (Zaccara et al., 2020; Viot and Wendel, 2023). These findings illustrate how environmental factors, combined with genetic adaptations, have shaped the evolutionary history and genetic diversity of Gossypium across its global range.

5 Ecological Factors Influencing Dispersal and Colonization

5.1 Role of ocean currents, wind patterns, and other natural vectors in Gossypium dispersal

The dispersal of Gossypium species across vast oceanic distances is significantly influenced by natural vectors such as ocean currents and wind patterns. Ocean currents play a crucial role in the transoceanic dispersal of seeds, as evidenced by the buoyancy and saltwater tolerance of Gossypium seeds, which allow them to survive prolonged periods in seawater and facilitate their movement across shorelines and between islands. Historical biogeographic studies have shown that ocean currents, such as the Antarctic Circumpolar Current and the West Wind Drift, have aided in the long-distance dispersal of various species, including Gossypium, by providing a means for seeds to raft across oceans (Figure 1) (Ceccarelli et al., 2016). Additionally, wind patterns can also contribute to the dispersal process by carrying seeds over shorter distances, complementing the role of ocean currents (Givnish and Renner, 2004).

5.2 Habitat adaptability and ecological adaptation of Gossypium in new regions

Gossypium species exhibit remarkable adaptability to diverse habitats, which is a key factor in their successful colonization of new regions. The ability of Gossypium seeds to remain viable after long periods of immersion in seawater allows them to establish populations in coastal and insular habitats. Once dispersed, these species can adapt to a variety of ecological conditions, including different soil types and climatic conditions. For instance, the cotton tribe (Gossypieae) has undergone multiple transoceanic dispersals, leading to the establishment of species in various tropical and subtropical regions worldwide (Grover et al., 2017). This adaptability is further supported by genetic studies, which reveal that Gossypium species have undergone significant genomic changes, such as genome downsizing and the accumulation of deletions, to better suit their new environments (Teixeira et al., 2017).

5.3 Effects of climate and soil conditions on the survival and reproduction of dispersed populations

The survival and reproduction of dispersed Gossypium populations are heavily influenced by the climate and soil conditions of the new regions they colonize. Climate plays a pivotal role in determining the suitability of a habitat for Gossypium species, as these plants typically thrive in warm, tropical, and subtropical climates (Bronnenhuber et al., 2011; Grover et al., 2017). Soil conditions, including nutrient availability and pH levels, also affect the growth and reproductive success of Gossypium populations. Studies have shown that Gossypium species can adapt to a range of soil types, which enhances their ability to establish and persist in new environments. Additionally, the genetic diversity within Gossypium populations, resulting from historical dispersal events, contributes to their resilience and ability to cope with varying climatic and soil conditions (Grover et al., 2018).

6 Dispersal and Colonization of Gossypium Species

6.1 Criteria for case study selection

The selection of case studies for examining the dispersal and colonization of Gossypium species is based on several critical criteria. These criteria ensure that the chosen examples provide comprehensive insights into the mechanisms and evolutionary consequences of transoceanic dispersal in this genus. The geographical distribution and historical biogeography of the species are paramount. Studies that elucidate the origins and spread of Gossypium species across different continents and islands are prioritized. For instance, the research on New World diploid cottons (Gossypium, subg. Houzingenia) highlights the transoceanic dispersal from Africa approximately 6.6 million years ago, followed by rapid diversification in the mid-Pleistocene, which led to the species' current distribution in regions such as Arizona, the Galapagos Islands, and Peru (Grover et al., 2018).

The genetic and genomic data available for the species play a crucial role. Comprehensive genomic analyses provide insights into the evolutionary processes, such as interspecific introgression and genome downsizing, which are essential for understanding the adaptive strategies of Gossypium species. For example, the comparative genomics study of the cotton tribe (Gossypieae) reveals significant genome downsizing in the genera Kokia and Gossypioides, which are closely related to Gossypium and exhibit remarkable geographic disjunction between the Hawaiian Islands and Madagascar/East Africa (Grover et al., 2017).

The ecological and evolutionary implications of dispersal mechanisms are considered. Studies that investigate the ecological adaptations and evolutionary pressures faced by Gossypium species during their dispersal and colonization are included. The evolutionary history of Gossypium, including the domestication of American allopolyploid species, provides a detailed account of the genetic and ecological factors that have shaped the current distribution and diversity of the genus (Derycke et al., 2008; Takayama et al., 2013).

6.2 Dispersal of Gossypium barbadense to the Americas

The dispersal of Gossypium barbadense to the Americas is a fascinating example of transoceanic seed movement and subsequent domestication. Gossypium barbadense, one of the two allopolyploid cotton species domesticated in the Americas, likely reached the New World through a combination of natural and anthropogenic mechanisms. The seeds of G. barbadense exhibit traits such as buoyancy and saltwater tolerance, which would have facilitated their dispersal across oceanic distances. This is particularly evident in the case of G. barbadense var. darwinii, whose seeds are capable of surviving prolonged immersion in seawater, enabling them to reach the Galapagos Islands from mainland South America (Cui et al., 2021).

The evolutionary history of Gossypium suggests that the genus originated around 5 to 10 million years ago, with allopolyploid cottons like G. barbadense appearing within the last 1 to 2 million years. This emergence was a result of the improbable transoceanic dispersal of an A genome taxon to the New World, followed by hybridization with an indigenous D genome diploid (Grover et al., 2018). The subsequent domestication of G. barbadense in South America, alongside G. hirsutum in Mesoamerica, represents a remarkable case of parallel evolution driven by human agricultural practices (Viot and Wendel, 2023).

The biogeographic history of Gossypium is marked by multiple long-distance dispersal events, which have significantly contributed to the genus's extensive geographic range. For instance, the New World diploid cottons, which include G. barbadense, likely originated from a transoceanic dispersal event from Africa approximately 6.6 million years ago. This was followed by rapid diversification during the mid-Pleistocene, involving further long-distance dispersals to regions such as Arizona, the Galapagos Islands, and Peru (Krosch et al., 2011).

6.3 Dispersal of Gossypium hirsutum to Pacific islands

The dispersal of Gossypium hirsutum, the most widely cultivated species of cotton, to the Pacific Islands exemplifies an important case of transoceanic dispersal within the Gossypium genus. The origins of G. hirsutum trace back to the Mesoamerican region, specifically Mexico and Central America, where its domestication likely began around 5 000 years ago. The presence of G. hirsutum across Pacific islands far from its origin in the Americas suggests a remarkable dispersal route shaped by natural and potentially anthropogenic factors (Figure 2) (Moura et al., 2019; Alavez et al., 2021).

Historical and botanical records reveal G. hirsutum among the flora of multiple Pacific islands by the time European explorers documented these regions in the 16th century. Several hypotheses exist regarding how this transoceanic dispersal may have occurred. Early studies propose that the buoyant seeds of G. hirsutum, encased in hardy, water-resistant fibers, might have floated across the Pacific Ocean, traveling with ocean currents that connect the western coast of Central America with islands in the Pacific. This natural mechanism of water-borne seed transport has been well-documented in other coastal plant species and offers a plausible scenario for G. hirsutum's dispersal across vast ocean distances (Wendel and Cronn, 2003).

Another prominent hypothesis involves early human influence. Evidence suggests that Polynesian navigators might have carried G. hirsutum seeds during their long-distance voyages between islands, as the plant's fibers provided valuable material for making textiles and other goods. Archaeological findings indicate the presence of Gossypium fibers in pre-contact Polynesian settlements, supporting the theory that early humans played a role in the dispersal and establishment of G. hirsutum across the Pacific (Alavez et al., 2021).

Genomic studies provide insights into the genetic diversity and structure of G. hirsutum populations across different Pacific islands, reinforcing both natural and human-mediated dispersal theories. Genetic analysis reveals minimal genetic divergence between G. hirsutum populations on islands and those in its native Mesoamerican range, suggesting recent dispersal events. Further, the identification of specific genetic markers associated with domesticated G. hirsutum supports the notion of human-mediated transport, as domesticated varieties tend to exhibit distinct genetic profiles that differentiate them from wild types. Molecular data thus reveal a dual dispersal scenario where natural seed flotation and early human navigation jointly facilitated the spread of G. hirsutum across the Pacific (Ulloa et al., 2013; Page et al., 2013).

The adaptation of G. hirsutum to various island ecosystems illustrates its ecological flexibility and resilience. On the Pacific islands, this species has successfully established itself in diverse environments, from coastal sandy soils to volcanic slopes, showcasing adaptability in reproductive strategies and water-use efficiency. These traits may have been pivotal for its successful establishment in the sometimes harsh and variable island climates. Studies highlight how the species' genetic plasticity and adaptability to environmental stresses enabled its persistence and proliferation across isolated Pacific ecosystems (Wendel et al., 2010).

7. Impact of Transoceanic Dispersal on Gossypium Evolution

7.1 Effects of dispersal on genetic diversity and speciation in Gossypium

Transoceanic dispersal has played a significant role in shaping the genetic diversity and speciation within the Gossypium genus. The long-distance dispersal events have led to the establishment of geographically isolated populations, which, through founder effects and genetic drift, have diverged significantly from their ancestral populations. For instance, the divergence of the genera Kokia and Gossypioides from Gossypium, which occurred approximately 5 million years ago, highlights the impact of such dispersal events on genetic diversity and speciation (Grover et al., 2021). Additionally, the New World diploid cottons (Gossypium, subgenus Houzingenia) originated from a transoceanic dispersal event from Africa around 6.6 million years ago, followed by rapid diversification during the mid-Pleistocene, further illustrating the role of dispersal in speciation (Liu et al., 2009; Grover et al., 2018). These events have resulted in a complex phylogenetic structure within the genus, with multiple instances of interspecific introgression and hybridization contributing to the genetic mosaic observed in modern Gossypium species (Cronn and Wendel, 2003).

7.2 Role in the domestication and intercontinental spread of cotton

The domestication and intercontinental spread of cotton are deeply intertwined with its history of transoceanic dispersal. The genus Gossypium includes four species that were independently domesticated for their seed fiber: two diploids from Africa-Asia and two allopolyploids from the Americas. The transoceanic dispersal of an A genome taxon to the New World, followed by hybridization with a native D genome diploid, led to the formation of allopolyploid cottons approximately 1 to 2 million years ago. This event was crucial for the development of the agronomically important species G. hirsutum and G. barbadense, which have since spread globally due to human cultivation (Cumming et al., 2014). The repeated polyploidization events and subsequent domestication have resulted in significant genetic and phenotypic innovations, such as the development of spinnable cotton fibers, which have been pivotal for the agricultural success of these species (Paterson et al., 2016).

7.3 Impact on ecosystem dynamics and biodiversity

The transoceanic dispersal of Gossypium species has had profound effects on ecosystem dynamics and biodiversity. The establishment of Gossypium populations in new environments has led to the development of unique ecological interactions and adaptations. For example, the polyploidization events in Gossypium, which were facilitated by transoceanic dispersal, have resulted in emergent properties such as higher fiber productivity and quality in tetraploid cottons compared to their diploid counterparts (Paterson et al., 2012). These adaptations have allowed Gossypium species to thrive in diverse ecological niches, contributing to the overall biodiversity of the regions they inhabit. Moreover, the genetic exchange between isolated populations through historical hybridization events has further enriched the genetic diversity within the genus, enhancing its ecological resilience (Hu et al., 2016).

8 Molecular Biology and Genomic Evidence

8.1 Analysis of gene flow in Gossypium using molecular markers

Gene flow in Gossypium species has been extensively studied using various molecular markers, revealing significant insights into the genetic structure and evolutionary history of these plants. For instance, the study on Gossypium hirsutum at its center of origin in Mexico demonstrated high genetic variation among metapopulations, with evidence of both historical and recent gene flow, including the presence of transgenes in wild populations. This suggests that gene flow has played a crucial role in shaping the genetic diversity of Gossypium species. Additionally, the use of chloroplast microsatellites and nuclear markers has helped uncover the complex patterns of gene flow and genetic differentiation in these species, highlighting the importance of both natural and anthropogenic factors in their evolution (Wegier et al., 2011).

8.2 Phylogenetic relationships and genetic diversity revealed by genomic data

Genomic data have provided a deeper understanding of the phylogenetic relationships and genetic diversity within the Gossypium genus. Whole genome resequencing of New World diploid cottons (Gossypium, subg. Houzingenia) has revealed a complex phylogenetic history, with evidence of multiple long-distance dispersals and interspecific introgression events (Grover et al., 2021). Comparative genomic analyses have shown that repetitive DNA elements, particularly transposable elements, play a significant role in genome size variation and evolution within the genus (Hawkins et al., 2006). Furthermore, the assembly of the Gossypium herbaceum genome and updates to the Gossypium arboreum and Gossypium hirsutum genomes have provided valuable insights into the phylogenetic relationships and evolutionary history of cotton A-genomes, resolving controversial concepts surrounding their origins (Huang et al., 2020).

8.3 Support and challenges from molecular biology studies on dispersal mechanisms

Molecular biology studies have both supported and challenged existing theories on the dispersal mechanisms of Gossypium species. Phylogenomic methods have indicated that the New World diploid cottons likely originated from a transoceanic dispersal event from Africa, followed by rapid diversification and multiple long-distance dispersals (Palmer et al., 2012). This supports the hypothesis that oceanic currents and wind patterns could facilitate the long-distance dispersal of cotton seeds. However, the differential amplification of transposable elements and the resulting genome size variation among Gossypium species present challenges in understanding the precise mechanisms of dispersal and adaptation. Additionally, the genomic stability observed in some species over thousands of years contrasts with the rapid genomic reorganization seen in others, suggesting that both gradual and punctuated evolutionary processes may be at play (Cheng et al., 2019). These findings underscore the complexity of dispersal mechanisms and the need for further research to fully elucidate the evolutionary dynamics of Gossypium species.

9 Conclusion and Future Perspectives

The study of transoceanic dispersal mechanisms in Gossypium has provided significant insights into the evolutionary and ecological dynamics of this genus. The ability of Gossypium seeds to remain viable after prolonged immersion in seawater and their buoyancy are critical factors that facilitate their dispersal across vast oceanic distances. For instance, the seeds of Gossypium barbadense var. darwinii have demonstrated sufficient buoyancy and saltwater tolerance to reach the Galapagos Islands from mainland South America, highlighting the role of ocean currents and possibly human activity in their distribution. Additionally, the genomic analyses of Gossypium and its relatives, such as Kokia and Gossypioides, have revealed significant genome downsizing and a bias towards deletions over insertions, which are likely consequences of their demographic and biogeographic history.

The evolutionary significance of transoceanic dispersal in Gossypium is underscored by the divergence and speciation events that have occurred as a result of these long-distance movements. The phylogenetic history of New World diploid cottons, for example, indicates that their diversification was driven by multiple transoceanic dispersals and subsequent rapid diversification during the mid-Pleistocene. This has led to a wide geographic distribution and significant genetic diversity within the genus. The ecological implications are equally profound, as the ability to colonize new and isolated habitats has allowed Gossypium species to exploit a variety of ecological niches, contributing to their survival and adaptation in diverse environments.

Understanding the mechanisms and consequences of transoceanic dispersal in Gossypium has practical applications for both conservation and resource utilization. Conservation efforts can benefit from this knowledge by identifying and protecting key habitats that facilitate seed dispersal and genetic exchange among populations. Additionally, the genetic diversity resulting from these dispersal events can be harnessed for crop improvement and resilience against environmental stresses. For instance, the saltwater tolerance observed in certain Gossypium species could be a valuable trait for developing cotton varieties that are more resilient to saline conditions. Furthermore, the insights gained from the genomic studies of Gossypium and its relatives can inform breeding programs aimed at enhancing desirable traits such as disease resistance and fiber quality.

Acknowledgments

We extend our sincere thanks to two anonymous peer reviewers for their invaluable feedback on the initial draft of this paper, whose critical evaluations and constructive suggestions have greatly contributed to the improvement of our manuscript.

Conflict of Interest Disclosure

The authors affirm that this research was conducted without any commercial or financial relationships that could be construed as a potential conflict of interest.

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