Testing Darwin´s conundrum in a threatened biome: invasive and non-invasive exotic species have the same phylogenetic field

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Despite those efforts, we are yet to definitively solve the question of what makes an exotic species successful in a new environment. Darwin’s conundrum represents that quest, with two contrasting hypotheses regarding exotic species establishment and success. We tested those hypotheses in a threatened biome: Mexican cloud forest (MCF). To do that, we estimated the phylogenetic field of non-invasive and invasive exotic angiosperm species, which allows us to see if they co-occur with more species closely related or distant to them. We also assessed if the phylogenetic field of exotic species is different in species with different growth forms and family representation in the MCF flora. We found that there is no difference between non-invasive and invasive exotic angiosperms’ phylogenetic fields, but there is a tendency for all exotic species to co-occur in assemblages with more closely related species, suggesting the importance for environmental filtering and support for the pre-adaptation hypothesis. Additionally, we did not find conclusive evidence that species with different growth forms had different phylogenetic fields. Finally, exotic species from angiosperm families that have more species on the MCF did not tend to co-occur in clustered assemblages, showing no effect of family representation on the MCF in the exotic species phylogenetic fields. Darwin’s conundrum pre-adaptation naturalization phylogenetic field exotic species cloud forest Figures Figure 1 Figure 2 Figure 3 Figure 4 Introduction The impact of exotic and invasive species on native species and their ecosystems has been a growing concern for the past few decades (Morales and Traveset 2009 ; Pyšek et al. 2017 ; Seebens et al. 2017 ; Roy et al. 2024). Especially in the context of global change, invasive species can add to the threat and increase the damage caused by abiotic changes (Brook et al. 2008 ; Daru et al. 2021 ). For example, by leading to local competitive exclusion and even to the global extinction of some species (Savidge 1987 ; Bellard et al. 2016 ). Biological invasions are context-dependent, and several factors can act both separately and in a synergistic manner to modulate their effect, such as the abiotic and biotic traits of invaded areas, stage of invasion, and human influence (González-Moreno et al. 2014 ; Wrzesień and Denisow 2017 ; Orbán et al. 2021 ; Falcão et al. 2022 ; Guo et al. 2024 ). Several studies have tried to find general patterns of invasion and their drivers, as well as the determinants of the susceptibility to invasion (Fuentes-Lillo et al. 2021 , Mounger et al. 2021 , Gioria et al. 2023 ). However, given the context-dependent nature of biological invasions, we are far from reaching a consensus about mechanisms and variables determining if a species will become invasive and the susceptibility of a particular environment. One of the open questions about the mechanisms driving biological invasions is the so-called Darwin’s conundrum (Diez et al. 2008 ), composed by two contrasting hypotheses considering the interaction between the exotic species and the native community as modulating invasions. These two hypotheses are: the (1) ‘pre-adaptation’ hypothesis, which predicts that exotic species that are more closely related to native species would have an advantage in colonizing their environment due to similar ecological requirements and mutual opportunities (Richardson and Pyšek 2006 ); and the (2) ‘naturalization’ hypothesis predicting that exotic species that are distantly related to native species would have such advantage due to mechanisms like enemy release or empty niche space, and would be less affected by competition with native species (Richardson and Pyšek 2006 ). Studies trying to solve Darwin’s conundrum have reached different conclusions, with support for both hypotheses (i.e., Park and Potter 2013 ; Qian and Sandel 2017 ; Wang et al. 2023 ). Although there is no consensus on the contexts that favor either hypothesis, some generalities have emerged such as the effect of spatial scale, invasion stage, and habitat disturbance (Diez et al. 2008 ; Bezeng et al. 2015 ; Wang et al. 2023 ; Almeida et al. 2024; Guo et al. 2024 ). Generally, there is more evidence to support the naturalization hypothesis for invasive exotic species at small spatial scales, whereas the pre-adaptation hypothesis seems to be favored for non-invasive exotic species at large spatial scales (Procheş et al. 2008 ; Schaefer et al. 2011 ), but there is conflicting evidence (i.e., Park and Potter 2013 ; Almeida et al. 2024). Initial studies on Darwin’s conundrum were based on taxonomic classifications – genera or families – to represent species relatedness (Duncan and Williams 2002 ; Ricciardi and Atkinson 2004 ). Then, as phylogenetic information became more available, they were incorporated into biological invasion studies to evaluate Darwin’s conundrum more robustly by directly considering species evolutionary relationships (Strauss et al. 2006 ). A recent approach leveraging phylogenetic information along with coexistence patterns to test Darwin’s conundrum is the phylogenetic field (Villalobos et al. 2013 ; Kusumoto et al. 2019 ). The phylogenetic field is the property of a focal species that is determined by the phylogenetic structure of the species that co-occur with it (Villalobos et al. 2013 ) and can be described at the level of its complete geographic distribution (e.g., Villalobos et al. 2013 ; Kusumoto et al. 2019 ) or within its occupied local assemblages (Pinto-Ledezma et al. 2020 ). In the context of biological invasions, we can use the phylogenetic field to evaluate if the phylogenetic structure of co-occurrence between an exotic species and the native recipients (e.g., co-occurrence among closely-related or distantly-related species; clustering or over-dispersion, respectively) are related to different components of biological invasions (Kusumoto et al. 2019 ; Pinto-Ledezma et al. 2020 ). Since it has been proposed, the phylogenetic field has been used in different taxa, spatial and temporal scales (Barnagaud et al. 2014 ; Villalobos et al. 2016 ; Ceccarelli et al. 2020 ). Specifically, regarding the study of biological invasions, three studies explicitly used the phylogenetic field approach to test Darwin’s conundrum, all of them with plants (Kusumoto et al. 2019 ; Pinto-Ledezma et al. 2020 ; Almeida et al. 2024). Two of these studies focused on local assemblages of native species occupied by exotic species and found more support for the pre-adaptation hypothesis (Pinto-Ledezma et al. 2020 ; Almeida et al. 2024), but Pinto-Ledezma et al. ( 2020 ) also found that environmental disturbances, such as fire, can change that relationship. Alternatively, Kusumoto et al. ( 2019 ) considered the complete distributions of exotics (instead of local assemblages) in the Japanese archipelago and found evidence to support both hypotheses. They also evaluated the importance of spatial scale and species attributes, such as their geographic extent, in influencing whether an exotic species tends to be more clustered or over-dispersed in relation to the native recipient community. With such few studies using the phylogenetic field approach, we are only starting to elucidate the relationship between different variables (from the species and the environment) and the two hypotheses of Darwin’s conundrum. It is important to highlight that the quest to solve Darwin´s conundrum is not only a theoretical one, because if we can understand the variables that can make an assemblage and potential invasive species support either side of the conundrum, we can better predict which species would potentially be more detrimental to certain communities and help targeting conservation efforts. Here, we evaluated Darwin’s conundrum in angiosperms of the Mexican Cloud Forest (MCF), a biome with the smallest remaining area in Mexico, currently covering less than one percent of the country and, at the same time, great biodiversity, with many species being endemic to this biome (Villaseñor and Gual-Díaz 2014 ). The peculiarities of the MCF make it a biome with high priority for conservation (CONABIO 2021 ) and give us a rare opportunity to conserve a high number of endangered species and ecosystem services within small geographical areas. At the same time, these characteristics highlight the need to study different components of MCF, including exotic species and MCF invasibility, in order to better inform future management strategies. Despite MCF importance, there are only a few studies about biological invasions on this biome in Mexico. We found a list of exotic plant species reported for the MCF (Villaseñor 2010 ), as well as some studies evaluating the effect of one or a few exotic plant species on different ecological processes of native species and management of exotic ones (e.g., higher tolerance of an invasive species: González de León et al. 2021 ; restoration of invaded MCF: Toledo-Aceves et al. 2022 ; fruit consumption of an exotic tree: Carpinteiro-Díaz et al. 2024 ). Given this context and to the best of our knowledge, our study is the first to comprehensively use phylogenetic data of a whole taxonomic group to elucidate biological invasion patterns in the MCF, and one of the few that use the phylogenetic field approach to test Darwin´s conundrum. We used the phylogenetic field approach to answer the following questions about MCF angiosperms: (1) Invasive and non-invasive exotic angiosperms support different sides of Darwin´s conundrum? (2) How is the phylogenetic structure of exotic species compared to the recipient angiosperm community? (3) Exotic species with different growth forms have differences in the structure of their phylogenetic fields? Methods Study area and species lists First, we geographically defined our study area by obtaining a polygon of the geographic distribution of MCF in Mexico. For this, we used a ‘land-use and vegetation’ polygon (INEGI 2021 ) and selected the polygons that were classified as ‘bosque de niebla o montaña’, as this represents the MCF biome in the country. Then, we obtained an angiosperm species list from the floristic-taxonomic catalogue of the Mexican cloud forest (MCF) compiled by Villaseñor ( 2010 ). This species list was based on an exhaustive literature revision and consists of the most complete list of the MCF floristic composition, including 6212 native and exotic species. We resolved the taxonomy of species in this list using the Leipzig catalogue of vascular plants (Freiberg et al. 2020 ) and removed species without georeferenced occurrence records inside the MCF polygon. The occurrence records were obtained from the Global Biodiversity Information Facility using the rgbif package (Chamberlain et al. 2025; see full data citation in Supplementary material 1). For a few species that had an unresolved status on the Leipzig catalogue, we used The World Flora Online (WFO 2024 ) to obtain an accepted name. The list ended up with 5128 native cloud forest angiosperm species in Mexico. To get the most up-to-date list of exotic angiosperm species with occurrence records in the MCF, we performed a search on the National Biodiversity Information System of Mexico (CONABIO 2024) which resulted in 281 exotic angiosperm species with at least one record in the MCF. The CONABIO database of occurrence records provides information on whether the exotic species is invasive or not (based on a rapid risk assessment, see Golubov et al. 2014). We used this label to separate our focal species into exotic and exotic-invasive (195 and 86 species, respectively). Geographic distribution data We obtained geographic distribution polygons (range maps) for 4364 of the MCF native species of our list from the Botanical Information and Ecology Network database (Enquist et al. 2016 ), using the R-package BIEN (Maitner et al. 2018 ). For the remaining native species (764), we used the same parameters reported by BIEN to generate their range map polygons (Maitner et al. 2018 ). We generated all the ranges for the exotic species as we were interested in their introduced range in the MCF instead of their native ranges. For the species for which we generated their geographic ranges, we downloaded occurrence data from the Global Biodiversity Information Facility using the rgbif package (GBIF.org 2024 ; Owens et al. 2025). Based on the geographic ranges of all considered species, natives and exotic/exotic-invasive, we built a presence-absence matrix using the letsR package (Vilela and Villalobos 2015 ). This matrix was then used to generate separate matrices for each of the 281 focal exotic and exotic-invasive species considering only the native species with which they co-occur. Both the presence-absence and co-occurrence matrices were constructed based on a grid-cell system of ~ 10km resolution, to coincide with the resolution of the environmental variables and models we used from BIEN (Maitner et al. 2018 ). To evaluate if exotic species with different growth forms could have differences in their phylogenetic fields. We downloaded growth form data from The Botanical Information and Ecology Network (BIEN), which is a standardized botanical database with data for 28 traits and around 93000 species (Maitner et al. 2018 ; see full data citation at Online resources 1). After joining trait names that represented the same growth form (e.g., vine and climbers), we ended up with five categories: vines, herbs, trees, shrubs and grasses. Phylogenetic information and metrics To obtain the phylogenetic relationships among our selected species, we used the phylogenetic plant megatree based on the LCVP taxonomic database (Freiberg et al., 2020 ) available in the V.PhyloMaker2 package (‘GBOTB.extended.LCVP.tre’; Jin and Qian 2022 ; available at https://github.com/megatrees/ ). This megatree was based on the megatree of seed plants reconstructed by Smith and Brown ( 2018 ). We used the U.PhyloMaker package to reconstruct the phylogenetic tree for our species, under the scenario 3 to add missing species at a midpoint of the genus or family branch (Jin and Qian 2023 ) . To evaluate the effect of the introduction of exotic species on the phylogenetic diversity (PD) of the MCF, we calculated Faith’s phylogenetic diversity for three sets of species: all species (exotic and natives), only native, and only exotic species. Then, to control for the effect of species richness (SR) on the phylogenetic diversity metric, we adjusted a loess model for each group (PD ~ SR) and used their residuals to test if the component of PD that is not explained by species richness (hereafter residual PD, resPD) between the native and the native + exotic species sets across the grid-cells of the MCF, using a Kolmogorov-Smirnov test. The phylogenetic fields for our 281 focal exotic species were constructed using a standardized effect size of the mean pairwise phylogenetic distance (sesMPD) of the native species that co-occurred within the distribution of each exotic species across the MCF. We chose the “taxa.labels” null model to calculate the standardized effect size, which randomly shuffle species from the phylogeny before calculating the mean phylogenetic distance between them, with 1000 iterations. All phylogenetic metrics were calculated using the R package picante (Kembel et al. 2010 ). Considering the complex and unique floristic composition of MCF and the different biogeographic affiliations of its species, we wanted to see if the potential patterns of exotics’ phylogenetic field over-dispersion or clustering could be explained by the under or over representation of certain angiosperm families in the MCF. For this, we used a supplementary table provided by Luna-Vega and Magallón ( 2010 ). On this table, they reported the number of species from all angiosperm families present in the global and MCF flora. We used these values to calculate the proportional representation of each family in our focal species list, classifying them as under or over-represented. Statistical analysis To test if there were differences between the phylogenetic fields of non-invasive and invasive exotic species and among their different growth forms, we used Kruskal-Wallis non-parametric tests. When a significant difference was found by this omnibus test, we used a Dunn’s test with a Bonferroni p correction to test the difference between groups. We also fitted a generalized linear model to test the influence of the values of proportional family representation on the phylogenetic field of our focal exotic species (sesMPD ~ proportional family representation, gaussian). We made all analyses and figures using the R-software (R Core Team 2024 ). Results Considering the 10km resolution, the maximum richness (measured as the overlapping of species ranges) was 3445 native species, 200 exotic species, 138 non-invasive exotics, and 65 invasive exotics (Fig. 1 ; the total species for these groups were, respectively: 5128, 281, 195, 86). The phylogenetic diversity, as described by resPD, represented by the MCF native angiosperms did not change when adding the phylogenetic diversity of exotic species (Kolmogorov-Smirnov, p > 0.1, Fig. 2 ). Regarding the phylogenetic fields of exotic species, there was no difference between invasive and non-invasive exotic MCF angiosperms (Kruskal-Wallis, p > 0.05; Fig. 3 ). From the 281 exotic species, only 16 species had a significant phylogenetic field when compared to the null model, with all of these species showing a clustered phylogenetic field (Fig. 3 ). In our omnibus test, we found initial evidence that species growth form could influence their phylogenetic field (Kruskal-Wallis, p 0.05, Table 1 ). Additionally, the fact that a family is more or less represented in the MCF than in the global flora did not explain clustered or over-dispersed phylogenetic field patterns (GLM, p > 0.05; Fig. 4 ). Table 1 Results (p-value with Bonferroni’s correction) of the p ost-hoc pairwise Dunn’s test comparing the Standardized effect of mean pairwise phylogenetic distance for different growth forms of exotic angiosperm species in the Mexican montane cloud forest vines trees herbs grasses trees 1.00 - - - herbs 0.78 0.12 - - grasses 1.00 0.63 1.00 - shrubs 0.30 0.07 1.00 1.00 Discussion Our findings reveal that the addition of 281 exotic species did not significantly increase phylogenetic diversity of the MCF native assemblage and that the majority of invasive and non-invasive exotic species in the Mexican cloud forest (MCF) do not co-occur with a particular, either closely or distantly related, set of native species; a result that does supports neither hypothesis of Darwin´s conundrum. There is some evidence to support that invasive species tend to be less relate to natives (Strauss et al. 2006 ), and therefore would align more with the naturalization hypothesis and thus show over-dispersed phylogenetic fields. However, this was not the case in our study, suggesting that other factors could play a more important role in the assembly of exotics within the MCF such as environmental filtering. Environmental filtering, known to foster clustered assemblages (Aldana et al. 2017 ; Shivaprakash et al. 2018 ), could explain the pattern we observed, as exotic species with significant phylogenetic fields were those co-occurring with clustered assemblages of natives. This finding instead supports the pre-adaptation hypothesis, which posits that exotic species that are closely related to the native recipients are more likely to succeed given their shared preferences with those natives as a result of their shared ancestry. In addition, species from over-represented families in the MCF did not tend to be more clustered than those from underrepresented families (Fig. 4 ). Finally, we didn’t find evidence supporting a difference between the phylogenetic fields of exotic species with different growth forms, contrary to what has been suggested in previous studies (e.g., Liu et al. 2024) . To test the role of invasion stages in shaping phylogenetic patterns, it is essential to clarify how these stages are defined. Different studies use varying criteria to define invasive species, and sometimes these criteria are not clearly stated. In our study, species were defined as invasive based on a national risk assessment (Golubov et al. 2014), and, despite the “being invasive elsewhere” criteria have been used in several studies before (Bradley et al. 2023 ), it may have poor precision. Such a lack of a precise definition of truly invasive species could explain our finding of no difference between the phylogenetic fields of invasive and non-invasive exotics. For example, Strauss et al. ( 2006 ) found that pest species—also classified based on government authorities—were less related to native taxa than non-pest exotics. Another dimension frequently linked to patterns of invasive and non-invasive species is the traits of exotic species ((Pyšek and Richardson 2008 , Palma et al. 2021 ). A lot of effort has been put to compile information about plant species traits (i.e., the BIEN database we used on our study, Maitner et al. 2018 ), but several studies evaluating how species traits are related to different patterns of biological invasions had to eliminate a considerable amount of the species pool from their analysis due to lack of data (i.e. Almeida et al. 2024). Here, we evaluated growth form because it was the only trait available for all of our 5409 considered species, and we did not want to eliminate species in order to evaluate the most comprehensive phylogenetic structure possible. Despite the indication of difference between the phylogenetic fields of exotic angiosperms with different growth forms, we could not find a difference between the groups in the pairwise comparison. This divergence between the omnibus and the post-hoc tests is not totally unexpected in hierarchical testing and could be explained by the more conservative approach of the multiple comparison correction in order to avoid false positives (Chen et al. 2018). Regardless, we cannot discard the possibility that there is a subtle difference that could be detected if the sample sizes of each group were more equal or we if we had more refined data available, like some quantitative traits. The biogeographical origins of the MCF flora, with both temperate and tropical affiliations (Morrone 2020 ), have contributed to its high phylogenetic diversity. The fact that most exotic species displayed random phylogenetic fields relative to the native assemblage may reflect this preexisting diversity. Exotic species entering such a diverse native assemblage in the MCF might neither cluster nor diverge significantly from the native pool, as they enter an already phylogenetically diverse assemblage. This situation likely explains why the introduction of exotic angiosperm species did not significantly increase the phylogenetic diversity of the MCF assemblage. As other studies have shown, the Mexican MCF biome is highly fragmented and have suffered several anthropogenic threats (CONABIO 2010 ), which leads us to think that our results align with the well-known intermediate disturbance hypothesis, which posits that some level of disturbance can facilitate the introduction of exotic species (Catford et al. 2012 ). In disturbed environments, competition is reduced, and environmental filtering plays a more prominent role, favoring species with pre-adaptations to local conditions (Wang et al. 2023 ). This could be one factor explaining why some exotic species in the highly disturbed MCF exhibited clustered phylogenetic fields, since human activities such as agriculture, urbanization, and deforestation have already drastically reduced the MCF's coverage (CONABIO 2010 ), further exacerbating its vulnerability to invasion. Despite the substantial body of literature on the MCF, with some studies considering phylogenetic information (i.e., for 15 species of plants, birds and rodents, Ornelas et al. 2013 ), our study is the first attempt to evaluate the phylogenetic structure of all angiosperm exotic species, in relation to the native species. Future studies could refine these insights by investigating how non-phylogenetic, ecological factors such as propagule pressure, competitive dynamics, and species traits, to better understand the processes shaping the phylogenetic structure of invaded assemblages. Given the diversity of pteridophytes in the MCF (Gual-Díaz and Rendón-Correa 2014 ), would be interesting that future studies include these taxa in order to better elucidate invasion patterns. Studies considering those additional subjects could help untangle Darwin’s conundrum and improve our ability to predict invasion dynamics, leading to more effective management strategies for this threatened ecosystem. Finally, replicating this study across other Mexican biomes could reveal whether the predominance of non-significant phylogenetic field patterns holds true in regions with different species pools and disturbance regimes. Declarations Funding This works was made with the support of the Secretaría de Ciencia, Humanidades, Tecnología e Innovación (Secihti) and the Instituto de Ecología, A.C. (INECOL), Mexico. Competing Interests The authors have no relevant financial or non-financial interests to disclose. Author Contributions All authors participated in the conception and design of the study. JF compiled the data, performed analyses, and wrote the first draft of the manuscript. FV advised JF on all the steps of the study and edited the manuscript. All authors approved the final manuscript. Data availability The data used in this study is available online on the databases cited in the main text. 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Diversity and Distributions 23:1323–1333. https://doi.org/10.1111/ddi.12620 R Core Team (2024) R: A Language and Environment for Statistical Computing. R Foundation for Statistical Computing, Vienna, Austria. https://www.R-project.org/ Ricciardi A, Atkinson SK (2004) Distinctiveness magnifies the impact of biological invaders in aquatic ecosystems. Ecology Letters 7:781–784. https://doi.org/10.1111/j.1461-0248.2004.00642.x Richardson DM, Pyšek P (2006) Plant invasions: Merging the concepts of species invasiveness and community invasibility. Progress in Physical Geography 30:409–431. https://doi.org/10.1191/0309133306pp490pr Rzedowski J (1996) Análisis preliminar de la flora vascular de los bosques mesófilos de montaña de México | Acta Botanica Mexicana. Acta Botanica Mexicana 35:25–44. https://doi.org/10.21829/abm35.1996.955 Savidge JA (1987) Extinction of an Island Forest Avifauna by an Introduced Snake. Ecology 68:660–668. https://doi.org/10.2307/1938471 Schaefer H, Hardy OJ, Silva L, et al (2011) Testing Darwin’s naturalization hypothesis in the Azores. Ecology Letters 14:389–396. https://doi.org/10.1111/j.1461- 0248.2011.01600.x Seebens H, Blackburn TM, Dyer EE, et al (2017) No saturation in the accumulation of alien species worldwide. Nature Communications 8:14435. https://doi.org/10.1038/ncomms14435 Shivaprakash KN, Ramesh BR, Umashaanker R, Dayanandan S (2018) Functional trait and community phylogenetic analyses reveal environmental filtering as the major determinant of assembly of tropical forest tree communities in the Western Ghats biodiversity hotspot in India. Forest Ecosystems 5:25. https://doi.org/10.1186/s40663-018-0144-0 Smith SA, Brown JW (2018) Constructing a broadly inclusive seed plant phylogeny. American Journal of Botany 105:302–314. https://doi.org/10.1002/ajb2.1019 Strauss SY, Webb CO, Salamin N (2006) Exotic taxa less related to native species are more invasive. Proceedings of the National Academy of Sciences of the United States of America 103:5841–5845. https://doi.org/10.1073/pnas.0508073103 Toledo-Aceves T, López-Barrera F, Vásquez-Reyes V, Günter S (2022) Restoration of tropical montane cloud forest in bracken dominated pastures: The role of nurse shrubs. Forest Ecology and Management 508:120055. https://doi.org/10.1016/j.foreco.2022.120055 Vilela B, Villalobos F (2015) letsR: a new R package for data handling and analysis in macroecology. Methods in Ecology and Evolution 6:1229–1234. https://doi.org/10.1111/2041-210X.12401 Villalobos F, Olalla‐Tárraga MÁ, Cianciaruso MV, et al (2016) Global patterns of mammalian co‐occurrence: phylogenetic and body size structure within species ranges. Journal of Biogeography 44:136–146. https://doi.org/10.1111/jbi.12826 Villalobos F, Rangel TF, Diniz-Filho JAF (2013) Phylogenetic fields of species: cross- species patterns of phylogenetic structure and geographical coexistence. Proceedings of the Royal Society B: Biological Sciences. https://doi.org/10.1098/rspb.2012.2570 Villaseñor JL (2010) El bosque húmedo de montaña en México y sus plantas vasculares: catálogo florístico-taxonómico. Comisión Nacional para el Conocimiento y Uso de la Biodiversidad - Universidad Nacional Autónoma de México, México, D.F. Villaseñor JL, Gual-Díaz M (2014) El bosque mesófilo de montaña en méxico y sus plantas con flores. In: Gual-Díaz M, Rendón-Correa A (eds) Bosques mesófilos de montaña de México: diversidad, ecología y manejo. Comisión Nacional para el Conocimiento y Uso de la Biodiversidad, Mexico, pp 221-236 Wang J, Li S-P, Ge Y, et al (2023) Darwin’s naturalization conundrum reconciled by changes of species interactions. Ecology 104:e3850. https://doi.org/10.1002/ecy.3850 WFO (2024) World Flora Online. Published on the Internet; http://www.worldfloraonline.org. Accessed 26 July 2024 Wrzesień M, Denisow B (2017) Factors responsible for the distribution of invasive plant species in the surroundings of railway areas. A case study from SE Poland. Biologia 72:1275–1284. https://doi.org/10.1515/biolog-2017-0146 Supplementary Files ESM1.docx Cite Share Download PDF Status: Under Review Version 1 posted Reviewers agreed at journal 30 Apr, 2025 Reviewers invited by journal 01 Apr, 2025 Editor invited by journal 21 Mar, 2025 Editor assigned by journal 20 Mar, 2025 First submitted to journal 20 Mar, 2025 You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. 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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-6271970","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":436640813,"identity":"550e43de-2aee-4750-ae30-9ba3338887a6","order_by":0,"name":"Jessica Falcao","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAA6klEQVRIie2PsQrCMBCGI4G4BFwPFPsKlkIp6MN06pTuDiKCYJfSuZuvoItzpBiXumeVgoM4FARxElMddEo7CuYbfv7hPu4OIYPhB2nNVIyRpxJzFdBrpuQIVBK/UmizVW+FDqper+Bofiz5BKxOFF/PcuJR1M52K+1hsXCAC7DT/LAZMqEOo0EgtUrKULck4CMZbhxGlALU1SvLC77zB/iWZCeHPZooKSWwXYA/kAwX4aKJEgeut03AXufCxWEClNT9YkdZIfltZPX38+LKbtN+p50JvTL7dAKv1I1XWF8dl3XTBoPB8J88ATlDSM0TKZDVAAAAAElFTkSuQmCC","orcid":"https://orcid.org/0000-0002-3748-0939","institution":"INECOL: Instituto de Ecologia","correspondingAuthor":true,"prefix":"","firstName":"Jessica","middleName":"","lastName":"Falcao","suffix":""},{"id":436640814,"identity":"a43fec08-218b-455a-83e5-3a5d5d558a95","order_by":1,"name":"Fabricio Villalobos","email":"","orcid":"","institution":"INECOL: Instituto de Ecologia","correspondingAuthor":false,"prefix":"","firstName":"Fabricio","middleName":"","lastName":"Villalobos","suffix":""}],"badges":[],"createdAt":"2025-03-20 18:39:20","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-6271970/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-6271970/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":81030382,"identity":"69ddb740-2139-4ba7-9676-cdb4d6c2b10a","added_by":"auto","created_at":"2025-04-21 11:15:10","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":170975,"visible":true,"origin":"","legend":"\u003cp\u003eGeographic pattern of species richness of angiosperms on the Mexican cloud forest. (A) Native species; (B) All exotic species; (C) Non-invasive exotic species; (D) Invasive exotic species\u003c/p\u003e","description":"","filename":"floatimage1.png","url":"https://assets-eu.researchsquare.com/files/rs-6271970/v1/01934cd2b3578257cfe1b035.png"},{"id":81028781,"identity":"720ebf4d-b6bc-4d56-84fb-da18a623574c","added_by":"auto","created_at":"2025-04-21 11:07:10","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":8177,"visible":true,"origin":"","legend":"\u003cp\u003eDensity distribution of residual phylogenetic diversity (resPD) of angiosperms from the Mexican montane cloud forest (Kolmogorov-Smirnov test, p\u0026gt;0.1). Red distribution corresponds to the set of native species, whereas the blue distribution corresponds to the set of species composed by both natives and exotics (invasive and non-invasive ones)\u003c/p\u003e","description":"","filename":"floatimage2.png","url":"https://assets-eu.researchsquare.com/files/rs-6271970/v1/9c2e731d907177883179cfa0.png"},{"id":81028778,"identity":"b0574d0d-2f6c-4d49-87e7-af2becf5b6ef","added_by":"auto","created_at":"2025-04-21 11:07:10","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":13333,"visible":true,"origin":"","legend":"\u003cp\u003ePhylogenetic fields of exotic angiosperms on the Mexican montane cloud forest, measured by the standardized effect size of the mean pairwise phylogenetic distance (sesMPD). The curve represents the density of sesMPD values for all exotic species, while points and asterisks represent the 281 individual species. Blue symbols are for non-invasive exotic species, while red symbols are for invasive exotic species. Exotic species with significant phylogenetic fields are represented by asterisks, while the ones with non-significant phylogenetic fields are depicted with circles\u003c/p\u003e","description":"","filename":"floatimage3.png","url":"https://assets-eu.researchsquare.com/files/rs-6271970/v1/53d7765cba0e03bedc5a772b.png"},{"id":81031420,"identity":"d0a20742-ddc7-48d6-9623-74b059c14141","added_by":"auto","created_at":"2025-04-21 11:23:10","extension":"png","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":164914,"visible":true,"origin":"","legend":"\u003cp\u003eRelationship between exotic angiosperm families representation in the Mexican cloud forest (MCF; difference between the percentage of species from each family at MCF and globally, from Luna-Vega and Magallón 2010) and the standardized effect size of mean pairwise phylogenetic distance of the assemblages where those species occur\u003c/p\u003e","description":"","filename":"floatimage4.png","url":"https://assets-eu.researchsquare.com/files/rs-6271970/v1/ba825e5306a8db89f85f93e4.png"},{"id":81032331,"identity":"a19a87d2-dc5d-4fc8-8dea-f8f00a80a4dd","added_by":"auto","created_at":"2025-04-21 11:31:10","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":810224,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-6271970/v1/3f424080-e19e-406f-9479-ef44a1257570.pdf"},{"id":81031422,"identity":"c3aa3b52-9d9b-4c57-aeb2-836a513ca21d","added_by":"auto","created_at":"2025-04-21 11:23:10","extension":"docx","order_by":5,"title":"","display":"","copyAsset":false,"role":"supplement","size":29773,"visible":true,"origin":"","legend":"","description":"","filename":"ESM1.docx","url":"https://assets-eu.researchsquare.com/files/rs-6271970/v1/b1949612f710acd7133989d6.docx"}],"financialInterests":"","formattedTitle":"Testing Darwin´s conundrum in a threatened biome: invasive and non-invasive exotic species have the same phylogenetic field","fulltext":[{"header":"Introduction","content":"\u003cp\u003eThe impact of exotic and invasive species on native species and their ecosystems has been a growing concern for the past few decades (Morales and Traveset \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e2009\u003c/span\u003e; Pyšek et al. \u003cspan citationid=\"CR53\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Seebens et al. \u003cspan citationid=\"CR64\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Roy et al. 2024). Especially in the context of global change, invasive species can add to the threat and increase the damage caused by abiotic changes (Brook et al. \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Daru et al. \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). For example, by leading to local competitive exclusion and even to the global extinction of some species (Savidge \u003cspan citationid=\"CR62\" class=\"CitationRef\"\u003e1987\u003c/span\u003e; Bellard et al. \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). Biological invasions are context-dependent, and several factors can act both separately and in a synergistic manner to modulate their effect, such as the abiotic and biotic traits of invaded areas, stage of invasion, and human influence (Gonz\u0026aacute;lez-Moreno et al. \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Wrzesień and Denisow \u003cspan citationid=\"CR77\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Orb\u0026aacute;n et al. \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Falc\u0026atilde;o et al. \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Guo et al. \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). Several studies have tried to find general patterns of invasion and their drivers, as well as the determinants of the susceptibility to invasion (Fuentes-Lillo et al. \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2021\u003c/span\u003e, Mounger et al. \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e2021\u003c/span\u003e, Gioria et al. \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). However, given the context-dependent nature of biological invasions, we are far from reaching a consensus about mechanisms and variables determining if a species will become invasive and the susceptibility of a particular environment.\u003c/p\u003e \u003cp\u003eOne of the open questions about the mechanisms driving biological invasions is the so-called Darwin\u0026rsquo;s conundrum (Diez et al. \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2008\u003c/span\u003e), composed by two contrasting hypotheses considering the interaction between the exotic species and the native community as modulating invasions. These two hypotheses are: the (1) \u0026lsquo;pre-adaptation\u0026rsquo; hypothesis, which predicts that exotic species that are more closely related to native species would have an advantage in colonizing their environment due to similar ecological requirements and mutual opportunities (Richardson and Pyšek \u003cspan citationid=\"CR60\" class=\"CitationRef\"\u003e2006\u003c/span\u003e); and the (2) \u0026lsquo;naturalization\u0026rsquo; hypothesis predicting that exotic species that are distantly related to native species would have such advantage due to mechanisms like enemy release or empty niche space, and would be less affected by competition with native species (Richardson and Pyšek \u003cspan citationid=\"CR60\" class=\"CitationRef\"\u003e2006\u003c/span\u003e). Studies trying to solve Darwin\u0026rsquo;s conundrum have reached different conclusions, with support for both hypotheses (i.e., Park and Potter \u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Qian and Sandel \u003cspan citationid=\"CR55\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Wang et al. \u003cspan citationid=\"CR74\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). Although there is no consensus on the contexts that favor either hypothesis, some generalities have emerged such as the effect of spatial scale, invasion stage, and habitat disturbance (Diez et al. \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Bezeng et al. \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Wang et al. \u003cspan citationid=\"CR74\" class=\"CitationRef\"\u003e2023\u003c/span\u003e; Almeida et al. 2024; Guo et al. \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). Generally, there is more evidence to support the naturalization hypothesis for invasive exotic species at small spatial scales, whereas the pre-adaptation hypothesis seems to be favored for non-invasive exotic species at large spatial scales (Procheş et al. \u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Schaefer et al. \u003cspan citationid=\"CR63\" class=\"CitationRef\"\u003e2011\u003c/span\u003e), but there is conflicting evidence (i.e., Park and Potter \u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Almeida et al. 2024). Initial studies on Darwin\u0026rsquo;s conundrum were based on taxonomic classifications \u0026ndash; genera or families \u0026ndash; to represent species relatedness (Duncan and Williams \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2002\u003c/span\u003e; Ricciardi and Atkinson \u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e2004\u003c/span\u003e). Then, as phylogenetic information became more available, they were incorporated into biological invasion studies to evaluate Darwin\u0026rsquo;s conundrum more robustly by directly considering species evolutionary relationships (Strauss et al. \u003cspan citationid=\"CR67\" class=\"CitationRef\"\u003e2006\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eA recent approach leveraging phylogenetic information along with coexistence patterns to test Darwin\u0026rsquo;s conundrum is the phylogenetic field (Villalobos et al. \u003cspan citationid=\"CR71\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Kusumoto et al. \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). The phylogenetic field is the property of a focal species that is determined by the phylogenetic structure of the species that co-occur with it (Villalobos et al. \u003cspan citationid=\"CR71\" class=\"CitationRef\"\u003e2013\u003c/span\u003e) and can be described at the level of its complete geographic distribution (e.g., Villalobos et al. \u003cspan citationid=\"CR71\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Kusumoto et al. \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2019\u003c/span\u003e) or within its occupied local assemblages (Pinto-Ledezma et al. \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). In the context of biological invasions, we can use the phylogenetic field to evaluate if the phylogenetic structure of co-occurrence between an exotic species and the native recipients (e.g., co-occurrence among closely-related or distantly-related species; clustering or over-dispersion, respectively) are related to different components of biological invasions (Kusumoto et al. \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Pinto-Ledezma et al. \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Since it has been proposed, the phylogenetic field has been used in different taxa, spatial and temporal scales (Barnagaud et al. \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Villalobos et al. \u003cspan citationid=\"CR70\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Ceccarelli et al. \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2020\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eSpecifically, regarding the study of biological invasions, three studies explicitly used the phylogenetic field approach to test Darwin\u0026rsquo;s conundrum, all of them with plants (Kusumoto et al. \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Pinto-Ledezma et al. \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Almeida et al. 2024). Two of these studies focused on local assemblages of native species occupied by exotic species and found more support for the pre-adaptation hypothesis (Pinto-Ledezma et al. \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Almeida et al. 2024), but Pinto-Ledezma et al. (\u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e2020\u003c/span\u003e) also found that environmental disturbances, such as fire, can change that relationship. Alternatively, Kusumoto et al. (\u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2019\u003c/span\u003e) considered the complete distributions of exotics (instead of local assemblages) in the Japanese archipelago and found evidence to support both hypotheses. They also evaluated the importance of spatial scale and species attributes, such as their geographic extent, in influencing whether an exotic species tends to be more clustered or over-dispersed in relation to the native recipient community. With such few studies using the phylogenetic field approach, we are only starting to elucidate the relationship between different variables (from the species and the environment) and the two hypotheses of Darwin\u0026rsquo;s conundrum. It is important to highlight that the quest to solve Darwin\u0026acute;s conundrum is not only a theoretical one, because if we can understand the variables that can make an assemblage and potential invasive species support either side of the conundrum, we can better predict which species would potentially be more detrimental to certain communities and help targeting conservation efforts.\u003c/p\u003e \u003cp\u003eHere, we evaluated Darwin\u0026rsquo;s conundrum in angiosperms of the Mexican Cloud Forest (MCF), a biome with the smallest remaining area in Mexico, currently covering less than one percent of the country and, at the same time, great biodiversity, with many species being endemic to this biome (Villase\u0026ntilde;or and Gual-D\u0026iacute;az \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). The peculiarities of the MCF make it a biome with high priority for conservation (CONABIO \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2021\u003c/span\u003e) and give us a rare opportunity to conserve a high number of endangered species and ecosystem services within small geographical areas. At the same time, these characteristics highlight the need to study different components of MCF, including exotic species and MCF invasibility, in order to better inform future management strategies. Despite MCF importance, there are only a few studies about biological invasions on this biome in Mexico. We found a list of exotic plant species reported for the MCF (Villase\u0026ntilde;or \u003cspan citationid=\"CR72\" class=\"CitationRef\"\u003e2010\u003c/span\u003e), as well as some studies evaluating the effect of one or a few exotic plant species on different ecological processes of native species and management of exotic ones (e.g., higher tolerance of an invasive species: Gonz\u0026aacute;lez de Le\u0026oacute;n et al. \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; restoration of invaded MCF: Toledo-Aceves et al. \u003cspan citationid=\"CR68\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; fruit consumption of an exotic tree: Carpinteiro-D\u0026iacute;az et al. \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2024\u003c/span\u003e ). Given this context and to the best of our knowledge, our study is the first to comprehensively use phylogenetic data of a whole taxonomic group to elucidate biological invasion patterns in the MCF, and one of the few that use the phylogenetic field approach to test Darwin\u0026acute;s conundrum. We used the phylogenetic field approach to answer the following questions about MCF angiosperms: (1) Invasive and non-invasive exotic angiosperms support different sides of Darwin\u0026acute;s conundrum? (2) How is the phylogenetic structure of exotic species compared to the recipient angiosperm community? (3) Exotic species with different growth forms have differences in the structure of their phylogenetic fields?\u003c/p\u003e"},{"header":"Methods","content":"\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e \u003ch2\u003eStudy area and species lists\u003c/h2\u003e \u003cp\u003eFirst, we geographically defined our study area by obtaining a polygon of the geographic distribution of MCF in Mexico. For this, we used a \u0026lsquo;land-use and vegetation\u0026rsquo; polygon (INEGI \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2021\u003c/span\u003e) and selected the polygons that were classified as \u0026lsquo;bosque de niebla o monta\u0026ntilde;a\u0026rsquo;, as this represents the MCF biome in the country. Then, we obtained an angiosperm species list from the floristic-taxonomic catalogue of the Mexican cloud forest (MCF) compiled by Villase\u0026ntilde;or (\u003cspan citationid=\"CR72\" class=\"CitationRef\"\u003e2010\u003c/span\u003e). This species list was based on an exhaustive literature revision and consists of the most complete list of the MCF floristic composition, including 6212 native and exotic species. We resolved the taxonomy of species in this list using the Leipzig catalogue of vascular plants (Freiberg et al. \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2020\u003c/span\u003e) and removed species without georeferenced occurrence records inside the MCF polygon. The occurrence records were obtained from the Global Biodiversity Information Facility using the rgbif package (Chamberlain et al. 2025; see full data citation in Supplementary material 1). For a few species that had an unresolved status on the Leipzig catalogue, we used The World Flora Online (WFO \u003cspan citationid=\"CR75\" class=\"CitationRef\"\u003e2024\u003c/span\u003e) to obtain an accepted name. The list ended up with 5128 native cloud forest angiosperm species in Mexico.\u003c/p\u003e \u003cp\u003eTo get the most up-to-date list of exotic angiosperm species with occurrence records in the MCF, we performed a search on the National Biodiversity Information System of Mexico (CONABIO 2024) which resulted in 281 exotic angiosperm species with at least one record in the MCF. The CONABIO database of occurrence records provides information on whether the exotic species is invasive or not (based on a rapid risk assessment, see Golubov et al. 2014). We used this label to separate our focal species into exotic and exotic-invasive (195 and 86 species, respectively).\u003c/p\u003e \u003c/div\u003e\n\u003ch3\u003eGeographic distribution data\u003c/h3\u003e\n\u003cp\u003eWe obtained geographic distribution polygons (range maps) for 4364 of the MCF native species of our list from the Botanical Information and Ecology Network database (Enquist et al. \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2016\u003c/span\u003e), using the R-package BIEN (Maitner et al. \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). For the remaining native species (764), we used the same parameters reported by BIEN to generate their range map polygons (Maitner et al. \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). We generated all the ranges for the exotic species as we were interested in their introduced range in the MCF instead of their native ranges. For the species for which we generated their geographic ranges, we downloaded occurrence data from the Global Biodiversity Information Facility using the rgbif package (GBIF.org \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2024\u003c/span\u003e; Owens et al. 2025). Based on the geographic ranges of all considered species, natives and exotic/exotic-invasive, we built a presence-absence matrix using the letsR package (Vilela and Villalobos \u003cspan citationid=\"CR69\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). This matrix was then used to generate separate matrices for each of the 281 focal exotic and exotic-invasive species considering only the native species with which they co-occur. Both the presence-absence and co-occurrence matrices were constructed based on a grid-cell system of ~\u0026thinsp;10km resolution, to coincide with the resolution of the environmental variables and models we used from BIEN (Maitner et al. \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). To evaluate if exotic species with different growth forms could have differences in their phylogenetic fields. We downloaded growth form data from The Botanical Information and Ecology Network (BIEN), which is a standardized botanical database with data for 28 traits and around 93000 species (Maitner et al. \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; see full data citation at Online resources 1). After joining trait names that represented the same growth form (e.g., vine and climbers), we ended up with five categories: vines, herbs, trees, shrubs and grasses.\u003c/p\u003e\n\u003ch3\u003ePhylogenetic information and metrics\u003c/h3\u003e\n\u003cp\u003eTo obtain the phylogenetic relationships among our selected species, we used the phylogenetic plant megatree based on the LCVP taxonomic database (Freiberg et al., \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2020\u003c/span\u003e) available in the V.PhyloMaker2 package (\u0026lsquo;GBOTB.extended.LCVP.tre\u0026rsquo;; Jin and Qian \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; available at \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://github.com/megatrees/\u003c/span\u003e\u003cspan address=\"https://github.com/megatrees/\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e). This megatree was based on the megatree of seed plants reconstructed by Smith and Brown (\u003cspan citationid=\"CR66\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). We used the U.PhyloMaker package to reconstruct the phylogenetic tree for our species, under the scenario 3 to add missing species at a midpoint of the genus or family branch (Jin and Qian \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2023\u003c/span\u003e) .\u003c/p\u003e \u003cp\u003eTo evaluate the effect of the introduction of exotic species on the phylogenetic diversity (PD) of the MCF, we calculated Faith\u0026rsquo;s phylogenetic diversity for three sets of species: all species (exotic and natives), only native, and only exotic species. Then, to control for the effect of species richness (SR) on the phylogenetic diversity metric, we adjusted a loess model for each group (PD\u0026thinsp;~\u0026thinsp;SR) and used their residuals to test if the component of PD that is not explained by species richness (hereafter residual PD, resPD) between the native and the native\u0026thinsp;+\u0026thinsp;exotic species sets across the grid-cells of the MCF, using a Kolmogorov-Smirnov test.\u003c/p\u003e \u003cp\u003eThe phylogenetic fields for our 281 focal exotic species were constructed using a standardized effect size of the mean pairwise phylogenetic distance (sesMPD) of the native species that co-occurred within the distribution of each exotic species across the MCF. We chose the \u0026ldquo;taxa.labels\u0026rdquo; null model to calculate the standardized effect size, which randomly shuffle species from the phylogeny before calculating the mean phylogenetic distance between them, with 1000 iterations. All phylogenetic metrics were calculated using the R package picante (Kembel et al. \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2010\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eConsidering the complex and unique floristic composition of MCF and the different biogeographic affiliations of its species, we wanted to see if the potential patterns of exotics\u0026rsquo; phylogenetic field over-dispersion or clustering could be explained by the under or over representation of certain angiosperm families in the MCF. For this, we used a supplementary table provided by Luna-Vega and Magall\u0026oacute;n (\u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e2010\u003c/span\u003e). On this table, they reported the number of species from all angiosperm families present in the global and MCF flora. We used these values to calculate the proportional representation of each family in our focal species list, classifying them as under or over-represented.\u003c/p\u003e \u003cdiv id=\"Sec6\" class=\"Section2\"\u003e \u003ch2\u003eStatistical analysis\u003c/h2\u003e \u003cp\u003eTo test if there were differences between the phylogenetic fields of non-invasive and invasive exotic species and among their different growth forms, we used Kruskal-Wallis non-parametric tests. When a significant difference was found by this \u003cem\u003eomnibus\u003c/em\u003e test, we used a Dunn\u0026rsquo;s test with a Bonferroni p correction to test the difference between groups. We also fitted a generalized linear model to test the influence of the values of proportional family representation on the phylogenetic field of our focal exotic species (sesMPD\u0026thinsp;~\u0026thinsp;proportional family representation, gaussian). We made all analyses and figures using the R-software (R Core Team \u003cspan citationid=\"CR56\" class=\"CitationRef\"\u003e2024\u003c/span\u003e).\u003c/p\u003e \u003c/div\u003e"},{"header":"Results","content":"\u003cp\u003eConsidering the 10km resolution, the maximum richness (measured as the overlapping of species ranges) was 3445 native species, 200 exotic species, 138 non-invasive exotics, and 65 invasive exotics (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e; the total species for these groups were, respectively: 5128, 281, 195, 86). The phylogenetic diversity, as described by resPD, represented by the MCF native angiosperms did not change when adding the phylogenetic diversity of exotic species (Kolmogorov-Smirnov, p\u0026thinsp;\u0026gt;\u0026thinsp;0.1, Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). Regarding the phylogenetic fields of exotic species, there was no difference between invasive and non-invasive exotic MCF angiosperms (Kruskal-Wallis, p\u0026thinsp;\u0026gt;\u0026thinsp;0.05; Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e). From the 281 exotic species, only 16 species had a significant phylogenetic field when compared to the null model, with all of these species showing a clustered phylogenetic field (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eIn our \u003cem\u003eomnibus\u003c/em\u003e test, we found initial evidence that species growth form could influence their phylogenetic field (Kruskal-Wallis, p\u0026thinsp;\u0026lt;\u0026thinsp;0.05). Despite that the \u003cem\u003epost-hoc\u003c/em\u003e test did not find a significant difference between any pair of groups after the correction for multiple comparisons (Dunn\u0026rsquo;s test, all p\u0026thinsp;\u0026gt;\u0026thinsp;0.05, Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). Additionally, the fact that a family is more or less represented in the MCF than in the global flora did not explain clustered or over-dispersed phylogenetic field patterns (GLM, p\u0026thinsp;\u0026gt;\u0026thinsp;0.05; Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab1\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 1\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eResults (p-value with Bonferroni\u0026rsquo;s correction) of the p\u003cem\u003eost-hoc\u003c/em\u003e pairwise Dunn\u0026rsquo;s test comparing the Standardized effect of mean pairwise phylogenetic distance for different growth forms of exotic angiosperm species in the Mexican montane cloud forest\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"5\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003evines\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003etrees\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eherbs\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003egrasses\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003etrees\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e1.00\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e-\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e-\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e-\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eherbs\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e0.78\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.12\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e-\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e-\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003egrasses\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e1.00\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.63\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e1.00\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e-\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eshrubs\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e0.30\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.07\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e1.00\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e1.00\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003e \u003c/p\u003e"},{"header":"Discussion","content":"\u003cp\u003eOur findings reveal that the addition of 281 exotic species did not significantly increase phylogenetic diversity of the MCF native assemblage and that the majority of invasive and non-invasive exotic species in the Mexican cloud forest (MCF) do not co-occur with a particular, either closely or distantly related, set of native species; a result that does supports neither hypothesis of Darwin\u0026acute;s conundrum. There is some evidence to support that invasive species tend to be less relate to natives (Strauss et al. \u003cspan citationid=\"CR67\" class=\"CitationRef\"\u003e2006\u003c/span\u003e), and therefore would align more with the naturalization hypothesis and thus show over-dispersed phylogenetic fields. However, this was not the case in our study, suggesting that other factors could play a more important role in the assembly of exotics within the MCF such as environmental filtering. Environmental filtering, known to foster clustered assemblages (Aldana et al. \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Shivaprakash et al. \u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e2018\u003c/span\u003e), could explain the pattern we observed, as exotic species with significant phylogenetic fields were those co-occurring with clustered assemblages of natives. This finding instead supports the pre-adaptation hypothesis, which posits that exotic species that are closely related to the native recipients are more likely to succeed given their shared preferences with those natives as a result of their shared ancestry. In addition, species from over-represented families in the MCF did not tend to be more clustered than those from underrepresented families (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e). Finally, we didn\u0026rsquo;t find evidence supporting a difference between the phylogenetic fields of exotic species with different growth forms, contrary to what has been suggested in previous studies (e.g., Liu et al. 2024) .\u003c/p\u003e \u003cp\u003eTo test the role of invasion stages in shaping phylogenetic patterns, it is essential to clarify how these stages are defined. Different studies use varying criteria to define invasive species, and sometimes these criteria are not clearly stated. In our study, species were defined as invasive based on a national risk assessment (Golubov et al. 2014), and, despite the \u0026ldquo;being invasive elsewhere\u0026rdquo; criteria have been used in several studies before (Bradley et al. \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2023\u003c/span\u003e), it may have poor precision. Such a lack of a precise definition of truly invasive species could explain our finding of no difference between the phylogenetic fields of invasive and non-invasive exotics. For example, Strauss et al. (\u003cspan citationid=\"CR67\" class=\"CitationRef\"\u003e2006\u003c/span\u003e) found that pest species\u0026mdash;also classified based on government authorities\u0026mdash;were less related to native taxa than non-pest exotics. Another dimension frequently linked to patterns of invasive and non-invasive species is the traits of exotic species ((Pyšek and Richardson \u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e2008\u003c/span\u003e, Palma et al. \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). A lot of effort has been put to compile information about plant species traits (i.e., the BIEN database we used on our study, Maitner et al. \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2018\u003c/span\u003e), but several studies evaluating how species traits are related to different patterns of biological invasions had to eliminate a considerable amount of the species pool from their analysis due to lack of data (i.e. Almeida et al. 2024). Here, we evaluated growth form because it was the only trait available for all of our 5409 considered species, and we did not want to eliminate species in order to evaluate the most comprehensive phylogenetic structure possible. Despite the indication of difference between the phylogenetic fields of exotic angiosperms with different growth forms, we could not find a difference between the groups in the pairwise comparison. This divergence between the \u003cem\u003eomnibus\u003c/em\u003e and the \u003cem\u003epost-hoc\u003c/em\u003e tests is not totally unexpected in hierarchical testing and could be explained by the more conservative approach of the multiple comparison correction in order to avoid false positives (Chen et al. 2018). Regardless, we cannot discard the possibility that there is a subtle difference that could be detected if the sample sizes of each group were more equal or we if we had more refined data available, like some quantitative traits.\u003c/p\u003e \u003cp\u003eThe biogeographical origins of the MCF flora, with both temperate and tropical affiliations (Morrone \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e2020\u003c/span\u003e), have contributed to its high phylogenetic diversity. The fact that most exotic species displayed random phylogenetic fields relative to the native assemblage may reflect this preexisting diversity. Exotic species entering such a diverse native assemblage in the MCF might neither cluster nor diverge significantly from the native pool, as they enter an already phylogenetically diverse assemblage. This situation likely explains why the introduction of exotic angiosperm species did not significantly increase the phylogenetic diversity of the MCF assemblage. As other studies have shown, the Mexican MCF biome is highly fragmented and have suffered several anthropogenic threats (CONABIO \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2010\u003c/span\u003e), which leads us to think that our results align with the well-known intermediate disturbance hypothesis, which posits that some level of disturbance can facilitate the introduction of exotic species (Catford et al. \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2012\u003c/span\u003e). In disturbed environments, competition is reduced, and environmental filtering plays a more prominent role, favoring species with pre-adaptations to local conditions (Wang et al. \u003cspan citationid=\"CR74\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). This could be one factor explaining why some exotic species in the highly disturbed MCF exhibited clustered phylogenetic fields, since human activities such as agriculture, urbanization, and deforestation have already drastically reduced the MCF's coverage (CONABIO \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2010\u003c/span\u003e), further exacerbating its vulnerability to invasion.\u003c/p\u003e \u003cp\u003eDespite the substantial body of literature on the MCF, with some studies considering phylogenetic information (i.e., for 15 species of plants, birds and rodents, Ornelas et al. \u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e2013\u003c/span\u003e), our study is the first attempt to evaluate the phylogenetic structure of all angiosperm exotic species, in relation to the native species. Future studies could refine these insights by investigating how non-phylogenetic, ecological factors such as propagule pressure, competitive dynamics, and species traits, to better understand the processes shaping the phylogenetic structure of invaded assemblages. Given the diversity of pteridophytes in the MCF (Gual-D\u0026iacute;az and Rend\u0026oacute;n-Correa \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2014\u003c/span\u003e), would be interesting that future studies include these taxa in order to better elucidate invasion patterns. Studies considering those additional subjects could help untangle Darwin\u0026rsquo;s conundrum and improve our ability to predict invasion dynamics, leading to more effective management strategies for this threatened ecosystem. Finally, replicating this study across other Mexican biomes could reveal whether the predominance of non-significant phylogenetic field patterns holds true in regions with different species pools and disturbance regimes.\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e\u003cstrong\u003eFunding\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThis works was made with the support of the Secretar\u0026iacute;a de Ciencia, Humanidades, Tecnolog\u0026iacute;a e Innovaci\u0026oacute;n (Secihti) and the Instituto de Ecolog\u0026iacute;a, A.C. (INECOL), Mexico.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eCompeting Interests\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe authors have no relevant financial or non-financial interests to disclose.\u003c/p\u003e\n\n\u003cp\u003eAuthor Contributions\u003c/p\u003e\n\u003cp\u003eAll authors participated in the conception and design of the study. JF compiled the data, performed analyses, and wrote the first draft of the manuscript. FV advised JF on all the steps of the study and edited the manuscript. All authors approved the final manuscript.\u003c/p\u003e\n\n\u003cp\u003eData availability \u003c/p\u003e\n\u003cp\u003eThe data used in this study is available online on the databases cited in the main text.\u003c/p\u003e\n"},{"header":"References","content":"\u003col\u003e\n\u003cli\u003eAldana AM, Carlucci MB, Fine PVA, Stevenson PR (2017) Environmental filtering of eudicot lineages underlies phylogenetic clustering in tropical South American flooded forests. Oecologia 183:327\u0026ndash;335. https://doi.org/10.1007/s00442- 016-3734-y\u003c/li\u003e\n\u003cli\u003eAlmeida, Thieres Santos, Oliveira, Eduardo Vin\u0026iacute;cius da Silva, Gouveia SF (2024) Exotic-to-native affinities and plant invasibility in a tropical dry forest. 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In: Gual-D\u0026iacute;az M, Rend\u0026oacute;n-Correa A (eds) Bosques mes\u0026oacute;filos de monta\u0026ntilde;a de M\u0026eacute;xico: diversidad, ecolog\u0026iacute;a y manejo. Comisi\u0026oacute;n Nacional para el Conocimiento y Uso de la Biodiversidad, Mexico, pp 221-236\u003c/li\u003e\n\u003cli\u003eWang J, Li S-P, Ge Y, et al (2023) Darwin\u0026rsquo;s naturalization conundrum reconciled by changes of species interactions. Ecology 104:e3850. https://doi.org/10.1002/ecy.3850\u003c/li\u003e\n\u003cli\u003eWFO (2024) World Flora Online. Published on the Internet; http://www.worldfloraonline.org. Accessed 26 July 2024\u003c/li\u003e\n\u003cli\u003eWrzesień M, Denisow B (2017) Factors responsible for the distribution of invasive plant species in the surroundings of railway areas. A case study from SE Poland. Biologia 72:1275\u0026ndash;1284. https://doi.org/10.1515/biolog-2017-0146\u003c/li\u003e\n\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":false,"highlight":"","institution":"","isAcceptedByJournal":false,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":true,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"biological-invasions","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"binv","sideBox":"Learn more about [Biological Invasions](https://www.springer.com/journal/10530)","snPcode":"10530","submissionUrl":"https://submission.nature.com/new-submission/10530/3","title":"Biological Invasions","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false},"keywords":"Darwin’s conundrum, pre-adaptation, naturalization, phylogenetic field, exotic species, cloud forest","lastPublishedDoi":"10.21203/rs.3.rs-6271970/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-6271970/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eExotic and invasive species have been the subject of several studies in the last few decades, due to their negative impact on native species and ecosystems. Despite those efforts, we are yet to definitively solve the question of what makes an exotic species successful in a new environment. Darwin\u0026rsquo;s conundrum represents that quest, with two contrasting hypotheses regarding exotic species establishment and success. We tested those hypotheses in a threatened biome: Mexican cloud forest (MCF). To do that, we estimated the phylogenetic field of non-invasive and invasive exotic angiosperm species, which allows us to see if they co-occur with more species closely related or distant to them. We also assessed if the phylogenetic field of exotic species is different in species with different growth forms and family representation in the MCF flora. We found that there is no difference between non-invasive and invasive exotic angiosperms\u0026rsquo; phylogenetic fields, but there is a tendency for all exotic species to co-occur in assemblages with more closely related species, suggesting the importance for environmental filtering and support for the pre-adaptation hypothesis. Additionally, we did not find conclusive evidence that species with different growth forms had different phylogenetic fields. Finally, exotic species from angiosperm families that have more species on the MCF did not tend to co-occur in clustered assemblages, showing no effect of family representation on the MCF in the exotic species phylogenetic fields.\u003c/p\u003e","manuscriptTitle":"Testing Darwin´s conundrum in a threatened biome: invasive and non-invasive exotic species have the same phylogenetic field","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2025-04-21 11:07:05","doi":"10.21203/rs.3.rs-6271970/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"reviewerAgreed","content":"","date":"2025-04-30T07:45:09+00:00","index":0,"fulltext":""},{"type":"reviewersInvited","content":"","date":"2025-04-01T04:52:39+00:00","index":"","fulltext":""},{"type":"editorInvited","content":"Biological Invasions","date":"2025-03-21T11:49:45+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2025-03-21T02:33:13+00:00","index":"","fulltext":""},{"type":"submitted","content":"Biological Invasions","date":"2025-03-20T14:37:38+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"biological-invasions","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"binv","sideBox":"Learn more about [Biological Invasions](https://www.springer.com/journal/10530)","snPcode":"10530","submissionUrl":"https://submission.nature.com/new-submission/10530/3","title":"Biological Invasions","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false}}],"origin":"","ownerIdentity":"72c6a40c-0921-4e93-821f-5581d49578ad","owner":[],"postedDate":"April 21st, 2025","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"under-review","subjectAreas":[],"tags":[],"updatedAt":"2026-05-15T23:34:09+00:00","versionOfRecord":[],"versionCreatedAt":"2025-04-21 11:07:05","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-6271970","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-6271970","identity":"rs-6271970","version":["v1"]},"buildId":"8U1c8b4HqxoKbykW_rLl7","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}