Low climatic niche overlap among allopatric woolly opossum species reflects phylogenetic and geographic influences in the Neotropics

Low climatic niche overlap among allopatric woolly opossum species reflects phylogenetic and geographic influences in the Neotropics | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Research Article Low climatic niche overlap among allopatric woolly opossum species reflects phylogenetic and geographic influences in the Neotropics Wellington Hannibal, Roniel Freitas-Oliveira, Ana Claudia Bernardes-Dias, and 3 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-9181042/v1 This work is licensed under a CC BY 4.0 License Status: Posted Version 1 posted You are reading this latest preprint version Abstract The study of species distribution has long intrigued naturalists, driven by ecological and evolutionary processes such as species interactions, climatic, and geological factors. Understanding these patterns is crucial for comprehending biodiversity. Despite extensive knowledge of species distributions, there is limited understanding of how climatic niches influence spatial segregation among closely related Neotropical marsupials, specifically the genus Caluromys . In this study, we investigated whether climatic niches determine the spatial segregation of Caluromys species in the Neotropical region, hypothesizing that niche overlap would be high due to geographical subdivision. We collected occurrence data for Caluromys species from the literature and the Global Biodiversity Information Facility, yielding 1,158 cleaned coordinates. We used 19 bioclimatic variables from WorldClim for historical climate data and analyzed climatic niche overlap through indices (Schoener’s D and Hellinger’s I) using Principal Component Analysis (PCA). We found low climatic niche overlap among Caluromys species pairs, with higher overlap between C. derbianus and C. lanatus than among other pairs. Contrary to our hypothesis, Caluromys species exhibited low climatic niche overlap on a broad geographic scale. The study emphasizes the need to investigate niche overlap at different geographic scales and highlights the role of historical climatic events in shaping current species distributions and niches. Caluromys derbianus Caluromys lanatus Caluromys philander Didelphidae Marsupials Figures Figure 1 Introduction Knowledge of species distribution has motivated naturalists since the 18th century to investigate the mechanisms that influence its distribution patterns (Whittaker et al. 2013). Distribution patterns are non-random and repetitive organizations resulting from complex interactions among species, climatic conditions, and geological features (Peterson et al. 2011). Congeneric species often distribute parapatrically because their environmental tolerances must differ significantly yet overlap partially, creating regions of potential sympatry where competitive exclusion could occur (Anderson et al. 2002; Gutiérrez et al. 2014). Biotic interactions can create and maintain geographic isolation at local scales and likely across larger geographic extents, for example, along extensive portions of their distributions (Gutiérrez et al. 2014). Additionally, sister species that have diverged purely as a result of geographical subdivision may show low spatial overlap but high niche overlap, with the degree of spatial overlap increasing and the degree of niche overlap decreasing among increasingly distant relatives, as a result of avoidance of competition or local adaptation (Cardillo and Warren 2016). Ecological niche models and assessments of environmental niche overlap, based on species' geographic distribution patterns, can help understand the degree of niche and spatial overlaps between species pairs (Anderson et al. 2002; Gutiérrez et al. 2014; Cardillo and Warren 2016). The Neotropical region exhibits a high diversity of mammals (Burgin et al. 2018), with a particular emphasis on marsupials. Neotropical marsupials comprise a diverse group, with approximately 124 species distributed across three orders; Didelphimorphia is the most representative, encompassing 116 species, 18 genera, and a single family – Didelphidae (Gardner 2008; Voss and Jansa 2021). The basal didelphid lineages inhabited the moist forests of South America, with most didelphids not entering North America until the Pliocene (Jansa et al. 2014). Among the didelphid marsupials, the genus Caluromys originated from a shared ancestor that also gave rise to the genus Caluromysiops and the subfamily Caluromyinae approximately 7.5 million years ago during the Miocene. The divergence between Caluromys philander and the clade comprising C. lanatus and C. derbianus likely occurred during the late Miocene (~5–6 Ma), whereas the split between C. lanatus and C. derbianus appears to be more recent, possibly in the Pliocene (~4–5 Ma) (Silva-Neto et al. 2024). Caluromys species segregate their occurrence across much of their geographical range, but occur predominantly in forested environments of Central and South America (Gardner 2008; Jansa et al. 2014; Fonseca and Astúa 2015; Ortega et al. 2021). In this study, we investigated whether the climate niche determines the spatial and phylogenetic segregation of Caluromys species in the Neotropical region. Considering that woolly opossums are sister species with high morphological and functional similarity, occurring in the canopy of forested habitats in Central and South America (Gardner 2008), we hypothesized that these species have a high niche overlap as a result of geographical subdivision (Cardillo and Warren 2016). Additionally, we expected greater niche overlap between more closely related species, such as C. derbianus and C. lanatus , than between other pairs. Materials and Methods Spatial overlap We compiled occurrence records for Caluromys species from the literature (Caramaschi 2005; Fonseca and Astúa 2015) and the Global Biodiversity Information Facility (GBIF 2023), resulting in 1,742 records. These data were cleaned using the “clean_coordinates” function from the “CoordinateClean” package (Zizka et al. 2019) to remove centroids, duplicates, and erroneous coordinates, yielding 1,158 valid occurrences (536 for C. derbianus , 412 for C. lanatus , and 210 for C. philander ; Supplementary Figure 1). To quantify spatial overlap, we generated minimum convex polygons (MCPs) for each species using the “mcp” function from the “adehabitatHR” package (Calenge and Fortmann 2023). Pairwise overlap was calculated as the proportion of intersecting area relative to the total combined area of both species (i.e., Jaccard similarity index), ranging from 0 (no overlap) to 1 (complete overlap). Niche overlap Climatic data were obtained from WorldClim version 2.1 (1970–2000) at 2.5 min resolution (Fick and Hijmans 2017). We extracted values of 19 bioclimatic variables for all occurrence points, generating the environmental presence matrix. Following Warren et al. (2008) and Broennimann et al. (2012), we used the “ecospat” package (Broennimann et al. 2022) to assess niche overlap. Background environmental conditions were defined using MCPs derived from species occurrences, from which climatic values were extracted to avoid inflating the available environmental space across the entire Neotropical region. Principal component analysis (PCA) was performed using the “dudi.pca” function from the “ade4” package (Dray and Dufour 2007), and niche overlap was quantified for all species pairs based on the first two axes. Overlap metrics included Schoener’s D and Hellinger’s I. Finally, kernel density functions were computed using “ecospat.grid.clim.dyn”, and niche overlap was estimated using “ecospat.niche.overlap”. Phylogenetic distance We used the first phylogeny of Caluromys species from Upham et al. (2019) and analyzed phylogenetic distances between species using the “cophenetic.phylo” function of the “ape” package (Paradis and Schliep 2019). In this study, we adopted the birth-death phylogeny, which provided a better representation of species within the taxonomy, for example, C. derbianus and C. lanatus within the subgenus Mallodephys Thomas, 1920, and C. philander within the subgenus Caluromys J.A. Allen, 1900 (Voss and Jansa 2009). All analyses were performed in R (R Core Team 2021). Results Spatial overlap among Caluromys species was generally low, particularly for C. derbianus , which showed minimal overlap with both C. lanatus (0.06) and C. philander (0.01). In contrast, C. lanatus and C. philander exhibited moderate spatial overlap (0.41), indicating greater geographic co-occurrence than in other species pairs. Climatic niche overlap among species pairs was low to moderate, as indicated by Schoener’s D and Hellinger’s I indices. Caluromys derbianus and C. lanatus showed the highest niche overlap (D = 0.36, I = 0.59), whereas lower values were observed for C. lanatus and C. philander (D = 0.19, I = 0.34) and C. derbianus and C. philander (D = 0.14, I = 0.30; Fig. 1 ). Phylogenetic distance followed a similar pattern, with C. derbianus and C. lanatus being more closely related (phylo.dist. = 13.52) than either C. derbianus and C. philander or C. lanatus and C. philander (phylo.dist. = 15.81), based on the birth–death phylogeny. Discussion Our results revealed a clear decoupling between spatial overlap, climatic niche similarity, and phylogenetic relatedness among Caluromys species. Spatial overlap was generally low, particularly for the Central American Woolly Opossum ( C. derbianus ), which showed minimal geographic overlap with the other species, whereas Brown-eared Woolly Opossum ( C. lanatus ) and Bare-tailed Woolly Opossum ( C. philander ) exhibited higher spatial co-occurrence. Despite this spatial segregation, phylogenetically closer species, such as C. derbianus and C. lanatus , showed higher climatic niche overlap than more distantly related pairs, as expected. This pattern is consistent with niche conservatism, whereby closely related species tend to retain similar ecological traits over evolutionary time (Wiens and Graham 2005 ; Peterson et al. 2011 ). As a general rule, two closely related species (e.g., morphologically, functionally, and phylogenetically similar) tend not to co-occur indefinitely in sympatry, resulting in Competitive exclusion (Hardin 1960 ). The low spatial overlap observed between these closely related species suggests that their geographic distributions may be structured by biotic interactions or historical biogeographic processes, which limit long-term coexistence in sympatry. Large marsupial species in South America, such as Didelphis albiventris and D. aurita , which occur allopatrically, present high niche similarity (D = 0.84), but their realized potential ranges do not overlap extensively because the presence of D. aurita acts as a biotic barrier to the realized and potential range of D. albiventris (Cáceres et al. 2016 ). On the other hand, when two congeneric species overlap in both distribution and niche similarity (e.g., Myrcia guianensis and M. splendens in South America), they are not ecologically equivalent (de Aguiar et al. 2021 ). In this context, Caluromys species, which are functionally similar (Gardner 2008 ), exhibit low to moderate climatic niche overlap and strong spatial segregation, likely reflecting both ecological and historical constraints. The genera Caluromys and Caluromysiops are included in the subfamily Caluromyinae, forming a monophyletic group supported by phylogenetic analyses based on morphological and molecular datasets (Voss and Jansa 2009 , 2021 ). The genus Caluromys has known fossil material from South America dating to the Pleistocene (Cartelle 1999 ). These animals are restricted to forest habitats and are among the American marsupials with the highest arboreal activity (Gardner 2008 ). The three species of Caluromys share similar ecological attributes, including nocturnal behavior, arboreal locomotion, and an omnivorous–frugivorous diet (Gardner 2008 ; Silva-Neto et al. 2024 ). Considering morphological attributes such as body mass, head–body length, and tail length, the three species do not appear to diverge significantly. Taxonomically, C. derbianus and C. lanatus belong to the same subgenus Mallodelphys, and both species have a well-developed marsupial pouch (Voss and Jansa 2009 ). Geographically, C. derbianus occurs in Central America and in the southern and southeastern regions of Mexico, as well as in western Colombia and western Ecuador in South America (Bucher and Hoffman 1980 ; Gardner 2008 ; Fonseca and Astúa 2015 ). Caluromys lanatus has a wide distribution in South America, occurring in the northern and central regions of Colombia, northwestern and southern Venezuela, eastern Ecuador, Peru, and Bolivia, as well as in the central-western, southeastern, and southern regions of Brazil, southern Paraguay, and Argentina (Cáceres and Carmignotto 2006 ; Gardner 2008 ; Fonseca and Astúa 2015 ). Caluromys philander is found further north in South America, occurring in Venezuela, Trinidad and Tobago, Guyana, Suriname, and French Guiana; in Brazil, it occurs in the north, central-west, southeast, and along the northeastern coast (Gardner 2008 ; Voss and Jansa 2009 ). However, C. derbianus and C. lanatus overlap in distribution in the northwest of South America, and C. lanatus and C. philander overlap in distribution in the northwest–southeast diagonal of South America (Supplementary Figure S1). At finer geographic scales (e.g., biomes and ecoregions), populations of Caluromys species may represent distinct evolutionary units. For example, C. lanatus could be divided into two morphological units: C. l. ochropus for populations of the Amazon and Cerrado, and C. l. lanatus for populations of the Atlantic Forest (Fonseca and Astúa 2015 ). Similarly, populations of C. philander from the southeastern and southern regions of Brazil (Atlantic Forest biome) may represent a distinct lineage compared to those in other regions of the country (Caramaschi 2005 ). Additionally, populations of C. philander in the Amazon and Atlantic Forests were separated approximately 1–2 million years ago due to early Pleistocene climatic oscillations. The expansion of forest areas in central Brazil played a crucial role as both current and past habitats for these forest species (Machado et al. 2024 ), and ecological niche models predict disjunct areas of high environmental suitability between Amazonian and Atlantic Forest populations. In conclusion, our results demonstrate that, at broad geographic scales, species with high morphological and functional similarity may exhibit low to moderate climatic niche overlap and strong spatial segregation. While niche similarity appears to be primarily structured by phylogenetic relatedness, spatial distribution is more strongly influenced by historical and biogeographic processes. These findings highlight the importance of considering spatial, ecological, and evolutionary dimensions when investigating patterns of species coexistence and diversification. Declarations Competing Interests The authors declare that they have no known financial or non-financial competing interests that could have appeared to influence the work reported in this paper. Funding This study was supported by the Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq), the INCT Program in Ecology, Evolution and Biodiversity Conservation and the Fundação de Amparo à Pesquisa do Estado de Goiás (FAPEG). Author Contribution W.H. conceived the study. W.H., R.F.O., and A.C.B.D. collected and organized the data. R. F. O., L.C.T. and M.S.L.R. contributed to the analytical framework. All authors contributed to writing and approved the final manuscript. Acknowledgements The authors are thankful to the three anonymous reviewers. WH, LCT, and MSLM thank the Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) for their research grant support. RFO thanks for funding as a CNPq research grant through the INCT Program in Ecology, Evolution, and Biodiversity Conservation. We thank the Fundação de Amparo à Pesquisa do Estado de Goiás - FAPEG (no. process: 202310267001404) References Anderson RP, Peterson AT, Gómez-Laverde M (2002) Using niche-based GIS modeling to test geographic predictions of competitive exclusion and competitive release in South American pocket mice. Oikos 98:3–16. https://doi.org/10.1034/j.1600-0706.2002.t01-1-980116.x Broennimann O, Fitzpatrick MC, Pearman PB et al (2012) Measuring ecological niche overlap from occurrence and spatial environmental data. Glob Ecol Biogeogr 21:481–497. https://doi.org/10.1111/j.1466-8238.2011.00698.x Bucher JE, Hoffman RS (1980) Caluromys derbianus. Mamm. Species 1 Burgin CJ, Colella JP, Kahn PL, Upham NS (2018) How many species of mammals are there? J Mammal 99:1–14. https://doi.org/10.1093/jmammal/gyx147 Cáceres NC, Carmignotto AP (2006) Caluromys lanatus. 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J Biogeogr 40:2209–2214. https://doi.org/10.1111/jbi.12235 Wiens JJ, Graham CH (2005) Niche conservatism: Integrating evolution, ecology, and conservation biology. Annu Rev Ecol Evol Syst 36:519–539. https://doi.org/10.1146/annurev.ecolsys.36.102803.095431 Zizka A, Silvestro D, Andermann T, Azevedo J, Duarte Ritter C, Edler D, Farooq H, Herdean A, Ariza M, Scharn R, Svanteson S, Wengstrom N, Zizka V, Antonelli A (2019) CoordinateCleaner: standardized cleaning of occurrence records from biological collection databases. Methods in Ecology and Evolution, -7. 10.1111/2041-210X.13152 %3Chttps: 10.1111="2041-210x.13152=" doi.org=""%3E%3C/https:%3E Additional Declarations No competing interests reported. Supplementary Files floatimage2.jpeg Supplementary Figure S1. Occurrence points based on Caramaschi (2005), Fonseca and Astúa (2015) and Global Biodiversity Information Facility (GBIF, 2023). Species distribution ranges based on the IUCN polygons (www.iucnedlist.org). 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Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-9181042","acceptedTermsAndConditions":true,"allowDirectSubmit":true,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":612717387,"identity":"4c77dd79-2537-4ee9-a524-5596c38548ef","order_by":0,"name":"Wellington Hannibal","email":"data:image/png;base64,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","orcid":"","institution":"Universidade Estadual de Goiás, UEG","correspondingAuthor":true,"prefix":"","firstName":"Wellington","middleName":"","lastName":"Hannibal","suffix":""},{"id":612717388,"identity":"e0907746-5006-4920-b21b-c6b5ea15a9fc","order_by":1,"name":"Roniel Freitas-Oliveira","email":"","orcid":"","institution":"Universidade Federal de Jataí, UFJ","correspondingAuthor":false,"prefix":"","firstName":"Roniel","middleName":"","lastName":"Freitas-Oliveira","suffix":""},{"id":612717392,"identity":"d5ac8f54-7dfc-4cdf-81c1-eff2d8f7a257","order_by":2,"name":"Ana Claudia Bernardes-Dias","email":"","orcid":"","institution":"Universidade Estadual de Goiás, UEG","correspondingAuthor":false,"prefix":"","firstName":"Ana","middleName":"Claudia","lastName":"Bernardes-Dias","suffix":""},{"id":612717405,"identity":"41478040-ef07-4a46-b7a5-bc9c28c18c16","order_by":3,"name":"Natália Albino","email":"","orcid":"","institution":"Universidade Estadual de Goiás, UEG","correspondingAuthor":false,"prefix":"","firstName":"Natália","middleName":"","lastName":"Albino","suffix":""},{"id":612717412,"identity":"8424dbf1-f06b-465e-a404-8025e3b06200","order_by":4,"name":"Levi Carina Terribile","email":"","orcid":"","institution":"Universidade Federal de Jataí, UFJ","correspondingAuthor":false,"prefix":"","firstName":"Levi","middleName":"Carina","lastName":"Terribile","suffix":""},{"id":612717428,"identity":"b963d310-abba-4011-8938-9944b1d42841","order_by":5,"name":"Matheus S. Lima-Ribeiro","email":"","orcid":"","institution":"National Institute for Science and Technology (INCT) in Ecology, Evolution and Biodiversity Conservation","correspondingAuthor":false,"prefix":"","firstName":"Matheus","middleName":"S.","lastName":"Lima-Ribeiro","suffix":""}],"badges":[],"createdAt":"2026-03-20 17:23:05","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-9181042/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-9181042/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":105603548,"identity":"c4e542e7-14ba-464c-8090-b02d6c49d6e1","added_by":"auto","created_at":"2026-03-27 21:18:17","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":418373,"visible":true,"origin":"","legend":"\u003cp\u003eClimate niche for: a) \u003cem\u003eC. derbianus \u003c/em\u003e(yellow) and \u003cem\u003eC. lanatus \u003c/em\u003e(blue), b) and \u003cem\u003eC. derbianus \u003c/em\u003e(yellow) and \u003cem\u003eC. philander \u003c/em\u003e(red), c) \u003cem\u003eC. lanatus \u003c/em\u003e(blue) and \u003cem\u003eC. philander\u003c/em\u003e (red). Purple shadows indicate the overlap between pairs of species. The lines represent the available climatic space, being 100% (solid) and 50% (dashed) line.\u003c/p\u003e","description":"","filename":"floatimage1.png","url":"https://assets-eu.researchsquare.com/files/rs-9181042/v1/a1e9014b2491ad8678228aa1.png"},{"id":108729923,"identity":"7c469ff1-7e76-462e-a1fa-da05b98f8337","added_by":"auto","created_at":"2026-05-07 18:10:11","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":566204,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-9181042/v1/dec196d1-a7a0-41d1-9415-161ae32ddf63.pdf"},{"id":105603549,"identity":"7506aef2-6bf9-47af-acae-d01f66f2a6e7","added_by":"auto","created_at":"2026-03-27 21:18:17","extension":"jpeg","order_by":1,"title":"","display":"","copyAsset":false,"role":"supplement","size":825157,"visible":true,"origin":"","legend":"\u003cp\u003eSupplementary Figure S1. Occurrence points based on Caramaschi (2005), Fonseca and Astúa (2015) and Global Biodiversity Information Facility (GBIF, 2023). Species distribution ranges based on the IUCN polygons (www.iucnedlist.org).\u003c/p\u003e","description":"","filename":"floatimage2.jpeg","url":"https://assets-eu.researchsquare.com/files/rs-9181042/v1/20a71bd47d4eab02a2332d63.jpeg"}],"financialInterests":"No competing interests reported.","formattedTitle":"Low climatic niche overlap among allopatric woolly opossum species reflects phylogenetic and geographic influences in the Neotropics","fulltext":[{"header":"Introduction","content":"\u003cp\u003eKnowledge of species distribution has motivated naturalists since the 18th century to investigate the mechanisms that influence its distribution patterns (Whittaker et al. 2013). Distribution patterns are non-random and repetitive organizations resulting from complex interactions among species, climatic conditions, and geological features (Peterson et al. 2011).\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eCongeneric species often distribute parapatrically because their environmental tolerances must differ significantly yet overlap partially, creating regions of potential sympatry where competitive exclusion could occur (Anderson et al. 2002; Gutiérrez et al. 2014). Biotic interactions can create and maintain geographic isolation at local scales and likely across larger geographic extents, for example, along extensive portions of their distributions (Gutiérrez et al. 2014). Additionally, sister species that have diverged purely as a result of geographical subdivision may show low spatial overlap but high niche overlap, with the degree of spatial overlap increasing and the degree of niche overlap decreasing among increasingly distant relatives, as a result of avoidance of competition or local adaptation (Cardillo and Warren 2016). Ecological niche models and assessments of environmental niche overlap, based on species' geographic distribution patterns, can help understand the degree of niche and spatial overlaps between species pairs (Anderson et al. 2002; Gutiérrez et al. 2014; Cardillo and Warren 2016).\u003c/p\u003e\n\u003cp\u003eThe Neotropical region exhibits a high diversity of mammals (Burgin et al. 2018), with a particular emphasis on marsupials. Neotropical marsupials comprise a diverse group, with approximately 124 species distributed across three orders; Didelphimorphia is the most representative, encompassing 116 species, 18 genera, and a single family – Didelphidae (Gardner 2008; Voss and Jansa 2021). The basal didelphid lineages inhabited the moist forests of South America, with most didelphids not entering North America until the Pliocene (Jansa et al. 2014). Among the didelphid marsupials, the genus \u003cem\u003eCaluromys\u003c/em\u003e originated from a shared ancestor that also gave rise to the genus\u003cem\u003e\u0026nbsp;Caluromysiops\u0026nbsp;\u003c/em\u003eand the subfamily Caluromyinae approximately 7.5 million years ago during the Miocene. The divergence between \u003cem\u003eCaluromys philander\u003c/em\u003e and the clade comprising \u003cem\u003eC. lanatus\u003c/em\u003e and \u003cem\u003eC. derbianus\u003c/em\u003e likely occurred during the late Miocene (~5–6 Ma), whereas the split between \u003cem\u003eC. lanatus\u003c/em\u003e and \u003cem\u003eC. derbianus\u003c/em\u003e appears to be more recent, possibly in the Pliocene (~4–5 Ma) (Silva-Neto et al. 2024). \u003cem\u003eCaluromys\u0026nbsp;\u003c/em\u003especies segregate their occurrence across much of their geographical range, but occur predominantly in forested environments of Central and South America (Gardner 2008; Jansa et al. 2014; Fonseca and Astúa 2015; Ortega et al. 2021).\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eIn this study, we investigated whether the climate niche determines the spatial and phylogenetic segregation of \u003cem\u003eCaluromys\u003c/em\u003e species in the Neotropical region. Considering that woolly opossums are sister species with high morphological and functional similarity, occurring in the canopy of forested habitats in Central and South America (Gardner 2008), we hypothesized that these species have a high niche overlap as a result of geographical subdivision (Cardillo and Warren 2016). Additionally, we expected greater niche overlap between more closely related species, such as \u003cem\u003eC. derbianus\u0026nbsp;\u003c/em\u003eand \u003cem\u003eC. lanatus\u003c/em\u003e, than between other pairs.\u003c/p\u003e"},{"header":"Materials and Methods","content":"\u003cp\u003e\u003cu\u003eSpatial overlap\u003c/u\u003e\u003c/p\u003e\n\u003cp\u003eWe compiled occurrence records for \u003cem\u003eCaluromys\u003c/em\u003e species from the literature (Caramaschi 2005; Fonseca and Astúa 2015) and the Global Biodiversity Information Facility (GBIF 2023), resulting in 1,742 records. These data were cleaned using the “clean_coordinates” function from the “CoordinateClean” package (Zizka et al. 2019) to remove centroids, duplicates, and erroneous coordinates, yielding 1,158 valid occurrences (536 for \u003cem\u003eC. derbianus\u003c/em\u003e, 412 for \u003cem\u003eC. lanatus\u003c/em\u003e, and 210 for \u003cem\u003eC. philander\u003c/em\u003e; Supplementary Figure 1).\u003c/p\u003e\n\u003cp\u003eTo quantify spatial overlap, we generated minimum convex polygons (MCPs) for each species using the “mcp” function from the “adehabitatHR” package (Calenge and Fortmann 2023). Pairwise overlap was calculated as the proportion of intersecting area relative to the total combined area of both species (i.e., Jaccard similarity index), ranging from 0 (no overlap) to 1 (complete overlap).\u003c/p\u003e\n\u003cp\u003e\u003cu\u003eNiche overlap\u003c/u\u003e\u003c/p\u003e\n\u003cp\u003eClimatic data were obtained from WorldClim version 2.1 (1970–2000) at 2.5 min resolution (Fick and Hijmans 2017). We extracted values of 19 bioclimatic variables for all occurrence points, generating the environmental presence matrix. Following Warren et al. (2008) and Broennimann et al. (2012), we used the “ecospat” package (Broennimann et al. 2022) to assess niche overlap.\u003c/p\u003e\n\u003cp\u003eBackground environmental conditions were defined using MCPs derived from species occurrences, from which climatic values were extracted to avoid inflating the available environmental space across the entire Neotropical region. Principal component analysis (PCA) was performed using the “dudi.pca” function from the “ade4” package (Dray and Dufour 2007), and niche overlap was quantified for all species pairs based on the first two axes. Overlap metrics included Schoener’s D and Hellinger’s I. Finally, kernel density functions were computed using “ecospat.grid.clim.dyn”, and niche overlap was estimated using “ecospat.niche.overlap”.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cu\u003ePhylogenetic distance\u003c/u\u003e\u003c/p\u003e\n\u003cp\u003eWe used the first phylogeny of \u003cem\u003eCaluromys\u003c/em\u003e species from Upham et al. (2019) and analyzed phylogenetic distances between species using the “cophenetic.phylo” function of the “ape” package (Paradis and Schliep 2019). In this study, we adopted the birth-death phylogeny, which provided a better representation of species within the taxonomy, for example, \u003cem\u003eC. derbianus\u0026nbsp;\u003c/em\u003eand \u003cem\u003eC. lanatus\u0026nbsp;\u003c/em\u003ewithin the subgenus \u003cem\u003eMallodephys\u0026nbsp;\u003c/em\u003eThomas, 1920, and \u003cem\u003eC. philander\u0026nbsp;\u003c/em\u003ewithin the subgenus \u003cem\u003eCaluromys\u0026nbsp;\u003c/em\u003eJ.A. Allen, 1900\u0026nbsp;(Voss and Jansa 2009). All analyses were performed in R\u0026nbsp;(R Core Team 2021).\u003c/p\u003e"},{"header":"Results","content":"\u003cp\u003eSpatial overlap among \u003cem\u003eCaluromys\u003c/em\u003e species was generally low, particularly for \u003cem\u003eC. derbianus\u003c/em\u003e, which showed minimal overlap with both \u003cem\u003eC. lanatus\u003c/em\u003e (0.06) and \u003cem\u003eC. philander\u003c/em\u003e (0.01). In contrast, \u003cem\u003eC. lanatus\u003c/em\u003e and \u003cem\u003eC. philander\u003c/em\u003e exhibited moderate spatial overlap (0.41), indicating greater geographic co-occurrence than in other species pairs.\u003c/p\u003e \u003cp\u003eClimatic niche overlap among species pairs was low to moderate, as indicated by Schoener\u0026rsquo;s D and Hellinger\u0026rsquo;s I indices. \u003cem\u003eCaluromys derbianus\u003c/em\u003e and \u003cem\u003eC. lanatus\u003c/em\u003e showed the highest niche overlap (D\u0026thinsp;=\u0026thinsp;0.36, I\u0026thinsp;=\u0026thinsp;0.59), whereas lower values were observed for \u003cem\u003eC. lanatus\u003c/em\u003e and \u003cem\u003eC. philander\u003c/em\u003e (D\u0026thinsp;=\u0026thinsp;0.19, I\u0026thinsp;=\u0026thinsp;0.34) and \u003cem\u003eC. derbianus\u003c/em\u003e and \u003cem\u003eC. philander\u003c/em\u003e (D\u0026thinsp;=\u0026thinsp;0.14, I\u0026thinsp;=\u0026thinsp;0.30; Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003ePhylogenetic distance followed a similar pattern, with \u003cem\u003eC. derbianus\u003c/em\u003e and \u003cem\u003eC. lanatus\u003c/em\u003e being more closely related (phylo.dist. = 13.52) than either \u003cem\u003eC. derbianus\u003c/em\u003e and \u003cem\u003eC. philander\u003c/em\u003e or \u003cem\u003eC. lanatus\u003c/em\u003e and \u003cem\u003eC. philander\u003c/em\u003e (phylo.dist. = 15.81), based on the birth\u0026ndash;death phylogeny.\u003c/p\u003e"},{"header":"Discussion","content":"\u003cp\u003eOur results revealed a clear decoupling between spatial overlap, climatic niche similarity, and phylogenetic relatedness among \u003cem\u003eCaluromys\u003c/em\u003e species. Spatial overlap was generally low, particularly for the Central American Woolly Opossum (\u003cem\u003eC. derbianus\u003c/em\u003e), which showed minimal geographic overlap with the other species, whereas Brown-eared Woolly Opossum (\u003cem\u003eC. lanatus\u003c/em\u003e) and Bare-tailed Woolly Opossum (\u003cem\u003eC. philander\u003c/em\u003e) exhibited higher spatial co-occurrence.\u003c/p\u003e \u003cp\u003eDespite this spatial segregation, phylogenetically closer species, such as \u003cem\u003eC. derbianus\u003c/em\u003e and \u003cem\u003eC. lanatus\u003c/em\u003e, showed higher climatic niche overlap than more distantly related pairs, as expected. This pattern is consistent with niche conservatism, whereby closely related species tend to retain similar ecological traits over evolutionary time (Wiens and Graham \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2005\u003c/span\u003e; Peterson et al. \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2011\u003c/span\u003e). As a general rule, two closely related species (e.g., morphologically, functionally, and phylogenetically similar) tend not to co-occur indefinitely in sympatry, resulting in Competitive exclusion (Hardin \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e1960\u003c/span\u003e). The low spatial overlap observed between these closely related species suggests that their geographic distributions may be structured by biotic interactions or historical biogeographic processes, which limit long-term coexistence in sympatry.\u003c/p\u003e \u003cp\u003eLarge marsupial species in South America, such as \u003cem\u003eDidelphis albiventris\u003c/em\u003e and \u003cem\u003eD. aurita\u003c/em\u003e, which occur allopatrically, present high niche similarity (D\u0026thinsp;=\u0026thinsp;0.84), but their realized potential ranges do not overlap extensively because the presence of \u003cem\u003eD. aurita\u003c/em\u003e acts as a biotic barrier to the realized and potential range of \u003cem\u003eD. albiventris\u003c/em\u003e (C\u0026aacute;ceres et al. \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). On the other hand, when two congeneric species overlap in both distribution and niche similarity (e.g., \u003cem\u003eMyrcia guianensis\u003c/em\u003e and \u003cem\u003eM. splendens\u003c/em\u003e in South America), they are not ecologically equivalent (de Aguiar et al. \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). In this context, \u003cem\u003eCaluromys\u003c/em\u003e species, which are functionally similar (Gardner \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2008\u003c/span\u003e), exhibit low to moderate climatic niche overlap and strong spatial segregation, likely reflecting both ecological and historical constraints.\u003c/p\u003e \u003cp\u003eThe genera \u003cem\u003eCaluromys\u003c/em\u003e and \u003cem\u003eCaluromysiops\u003c/em\u003e are included in the subfamily Caluromyinae, forming a monophyletic group supported by phylogenetic analyses based on morphological and molecular datasets (Voss and Jansa \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2009\u003c/span\u003e, \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). The genus \u003cem\u003eCaluromys\u003c/em\u003e has known fossil material from South America dating to the Pleistocene (Cartelle \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e1999\u003c/span\u003e). These animals are restricted to forest habitats and are among the American marsupials with the highest arboreal activity (Gardner \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2008\u003c/span\u003e). The three species of \u003cem\u003eCaluromys\u003c/em\u003e share similar ecological attributes, including nocturnal behavior, arboreal locomotion, and an omnivorous\u0026ndash;frugivorous diet (Gardner \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Silva-Neto et al. \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). Considering morphological attributes such as body mass, head\u0026ndash;body length, and tail length, the three species do not appear to diverge significantly. Taxonomically, \u003cem\u003eC. derbianus\u003c/em\u003e and \u003cem\u003eC. lanatus\u003c/em\u003e belong to the same subgenus Mallodelphys, and both species have a well-developed marsupial pouch (Voss and Jansa \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2009\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eGeographically, \u003cem\u003eC. derbianus\u003c/em\u003e occurs in Central America and in the southern and southeastern regions of Mexico, as well as in western Colombia and western Ecuador in South America (Bucher and Hoffman \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e1980\u003c/span\u003e; Gardner \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Fonseca and Ast\u0026uacute;a \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). \u003cem\u003eCaluromys lanatus\u003c/em\u003e has a wide distribution in South America, occurring in the northern and central regions of Colombia, northwestern and southern Venezuela, eastern Ecuador, Peru, and Bolivia, as well as in the central-western, southeastern, and southern regions of Brazil, southern Paraguay, and Argentina (C\u0026aacute;ceres and Carmignotto \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Gardner \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Fonseca and Ast\u0026uacute;a \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). \u003cem\u003eCaluromys philander\u003c/em\u003e is found further north in South America, occurring in Venezuela, Trinidad and Tobago, Guyana, Suriname, and French Guiana; in Brazil, it occurs in the north, central-west, southeast, and along the northeastern coast (Gardner \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Voss and Jansa \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2009\u003c/span\u003e). However, \u003cem\u003eC. derbianus\u003c/em\u003e and \u003cem\u003eC. lanatus\u003c/em\u003e overlap in distribution in the northwest of South America, and \u003cem\u003eC. lanatus\u003c/em\u003e and \u003cem\u003eC. philander\u003c/em\u003e overlap in distribution in the northwest\u0026ndash;southeast diagonal of South America (Supplementary Figure S1).\u003c/p\u003e \u003cp\u003eAt finer geographic scales (e.g., biomes and ecoregions), populations of \u003cem\u003eCaluromys\u003c/em\u003e species may represent distinct evolutionary units. For example, \u003cem\u003eC. lanatus\u003c/em\u003e could be divided into two morphological units: \u003cem\u003eC. l. ochropus\u003c/em\u003e for populations of the Amazon and Cerrado, and \u003cem\u003eC. l. lanatus\u003c/em\u003e for populations of the Atlantic Forest (Fonseca and Ast\u0026uacute;a \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). Similarly, populations of \u003cem\u003eC. philander\u003c/em\u003e from the southeastern and southern regions of Brazil (Atlantic Forest biome) may represent a distinct lineage compared to those in other regions of the country (Caramaschi \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2005\u003c/span\u003e). Additionally, populations of \u003cem\u003eC. philander\u003c/em\u003e in the Amazon and Atlantic Forests were separated approximately 1\u0026ndash;2\u0026nbsp;million years ago due to early Pleistocene climatic oscillations. The expansion of forest areas in central Brazil played a crucial role as both current and past habitats for these forest species (Machado et al. \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2024\u003c/span\u003e), and ecological niche models predict disjunct areas of high environmental suitability between Amazonian and Atlantic Forest populations.\u003c/p\u003e \u003cp\u003eIn conclusion, our results demonstrate that, at broad geographic scales, species with high morphological and functional similarity may exhibit low to moderate climatic niche overlap and strong spatial segregation. While niche similarity appears to be primarily structured by phylogenetic relatedness, spatial distribution is more strongly influenced by historical and biogeographic processes. These findings highlight the importance of considering spatial, ecological, and evolutionary dimensions when investigating patterns of species coexistence and diversification.\u003c/p\u003e"},{"header":"Declarations","content":" \u003cp\u003e \u003cstrong\u003eCompeting Interests\u003c/strong\u003e \u003cp\u003eThe authors declare that they have no known financial or non-financial competing interests that could have appeared to influence the work reported in this paper.\u003c/p\u003e \u003c/p\u003e\u003ch2\u003eFunding\u003c/h2\u003e \u003cp\u003eThis study was supported by the Conselho Nacional de Desenvolvimento Cient\u0026iacute;fico e Tecnol\u0026oacute;gico (CNPq), the INCT Program in Ecology, Evolution and Biodiversity Conservation and the Funda\u0026ccedil;\u0026atilde;o de Amparo \u0026agrave; Pesquisa do Estado de Goi\u0026aacute;s (FAPEG).\u003c/p\u003e\u003ch2\u003eAuthor Contribution\u003c/h2\u003e\u003cp\u003eW.H. conceived the study. W.H., R.F.O., and A.C.B.D. collected and organized the data. R. F. O., L.C.T. and M.S.L.R. contributed to the analytical framework. All authors contributed to writing and approved the final manuscript.\u003c/p\u003e\u003ch2\u003eAcknowledgements\u003c/h2\u003e \u003cp\u003eThe authors are thankful to the three anonymous reviewers. WH, LCT, and MSLM thank the Conselho Nacional de Desenvolvimento Cient\u0026iacute;fico e Tecnol\u0026oacute;gico (CNPq) for their research grant support. RFO thanks for funding as a CNPq research grant through the INCT Program in Ecology, Evolution, and Biodiversity Conservation. We thank the Funda\u0026ccedil;\u0026atilde;o de Amparo \u0026agrave; Pesquisa do Estado de Goi\u0026aacute;s - FAPEG (no. process: 202310267001404)\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\u003cli\u003e\u003cspan\u003eAnderson RP, Peterson AT, G\u0026oacute;mez-Laverde M (2002) Using niche-based GIS modeling to test geographic predictions of competitive exclusion and competitive release in South American pocket mice. Oikos 98:3\u0026ndash;16. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1034/j.1600-0706.2002.t01-1-980116.x\u003c/span\u003e\u003cspan address=\"10.1034/j.1600-0706.2002.t01-1-980116.x\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eBroennimann O, Fitzpatrick MC, Pearman PB et al (2012) Measuring ecological niche overlap from occurrence and spatial environmental data. 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Methods in Ecology and Evolution, -7. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003e10.1111/2041-210X.13152 %3Chttps: 10.1111=\"2041-210x.13152=\" doi.org=\"\"%3E%3C/https:%3E\u003c/span\u003e\u003cspan address=\"10.1111/2041-210X.13152 %3Chttps: 10.1111=\u0026quot;2041-210x.13152=\u0026quot; doi.org=\u0026quot;\u0026quot;%3E%3C/https:%3E\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":true,"highlight":"","institution":"","isAcceptedByJournal":false,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true},"keywords":"Caluromys derbianus, Caluromys lanatus, Caluromys philander, Didelphidae, Marsupials","lastPublishedDoi":"10.21203/rs.3.rs-9181042/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-9181042/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eThe study of species distribution has long intrigued naturalists, driven by ecological and evolutionary processes such as species interactions, climatic, and geological factors. Understanding these patterns is crucial for comprehending biodiversity. Despite extensive knowledge of species distributions, there is limited understanding of how climatic niches influence spatial segregation among closely related Neotropical marsupials, specifically the genus \u003cem\u003eCaluromys\u003c/em\u003e. In this study, we investigated whether climatic niches determine the spatial segregation of \u003cem\u003eCaluromys\u003c/em\u003e species in the Neotropical region, hypothesizing that niche overlap would be high due to geographical subdivision. We collected occurrence data for \u003cem\u003eCaluromys\u003c/em\u003e species from the literature and the Global Biodiversity Information Facility, yielding 1,158 cleaned coordinates. We used 19 bioclimatic variables from WorldClim for historical climate data and analyzed climatic niche overlap through indices (Schoener\u0026rsquo;s D and Hellinger\u0026rsquo;s I) using Principal Component Analysis (PCA). We found low climatic niche overlap among \u003cem\u003eCaluromys\u003c/em\u003e species pairs, with higher overlap between \u003cem\u003eC. derbianus\u003c/em\u003e and \u003cem\u003eC. lanatus\u003c/em\u003e than among other pairs. Contrary to our hypothesis, \u003cem\u003eCaluromys\u003c/em\u003e species exhibited low climatic niche overlap on a broad geographic scale. The study emphasizes the need to investigate niche overlap at different geographic scales and highlights the role of historical climatic events in shaping current species distributions and niches.\u003c/p\u003e","manuscriptTitle":"Low climatic niche overlap among allopatric woolly opossum species reflects phylogenetic and geographic influences in the Neotropics","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2026-03-27 21:18:12","doi":"10.21203/rs.3.rs-9181042/v1","editorialEvents":[{"type":"communityComments","content":0}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true}}],"origin":"","ownerIdentity":"538d88d3-f290-4ff8-9324-c903577f10b9","owner":[],"postedDate":"March 27th, 2026","published":true,"recentEditorialEvents":[{"type":"decision","content":"Rejected","date":"2026-05-07T18:02:48+00:00","index":"","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2026-05-07T10:57:44+00:00","index":32,"fulltext":""},{"type":"reviewerAgreed","content":"191553433796195016105573168809717119968","date":"2026-05-07T10:48:51+00:00","index":31,"fulltext":""}],"rejectedJournal":[],"revision":"","amendment":"","status":"posted","subjectAreas":[],"tags":[],"updatedAt":"2026-05-07T18:09:30+00:00","versionOfRecord":[],"versionCreatedAt":"2026-03-27 21:18:12","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-9181042","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-9181042","identity":"rs-9181042","version":["v1"]},"buildId":"XKTyCvWXoU3ODBz1xrDgd","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}