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361 Geographical patterns in the distribution of naturalized plants in Central America Eduardo Chacón-Madrigal1,2 , Julissa Rojas-Sandoval3,4 , Lilian Ferrufino-Acosta5, Rodolfo Flores6,7 , Pablo Galán8, AnaLu MacVean9, Dagoberto Rodríguez-Delcid10 , Iris Saldivar-Gómez11 , Yader Ruiz12 , Omar López13 1 Herbario Luis Fournier Origgi, Centro de Investigación en Biodiversidad y Ecología Tropical (CIBET), Universidad de Costa Rica, San José, Costa Rica 2 Herbario Nacional, Departamento de Historia Natural, Museo Nacional de Costa Rica, San José, Costa Rica 3 Institute of the Environment, University of Connecticut, Storrs, CT, USA 4 Department of Geography, Sustainability, Community, and Urban Studies, University of Connecticut, Storrs, CT, USA 5 Escuela de Biología, Facultad de Ciencias, Universidad Nacional Autónoma de Honduras, Tegucigalpa, Honduras 6 Los Naturalistas, P.O. Box 0426-01459, David, Chiriquí, Panama 7 Departamento de Botánica, Universidad de Panamá, Estafeta Universitaria, Panama City, Panama 8 San Salvador, El Salvador 9 Environmental Horticulture Department, York College of Pennsylvania, York, PA, USA 10 Herbario LAGU, Jardín Botánico La Laguna, La Libertad, El Salvador 11 Centro de Investigación Capacitación y Formación Ambiental. CICFA, Managua, Nicaragua 12 Departamento de Ciencias Naturales, Facultad Multidisciplinaria Oriental, Universidad de El Salvador, San Salvador, El Salvador 13 Instituto Interamericano para la Investigación del Cambio Global, Panama City, Panama Corresponding author: Eduardo Chacón-Madrigal ([email protected]) Copyright: © Eduardo Chacón-Madrigal et al. This is an open access article distributed under terms of the Creative Commons Attribution License (Attribution 4.0 International – CC BY 4.0). Research Article Abstract Non-native plant species are on the rise globally, yet the distribution patterns and environmental drivers in biodiversity-rich regions such as Central America remain poorly understood. These species are affecting biodiversity, ecosystem integrity, and conservation efforts, especially when they become invasive. We analyzed the spatial distribution of 751 naturalized plant species using more than 42,000 records collected through the Global Biodiversity Information Facility (GBIF) across the seven countries in Central America. We evaluated the influence of environmental variables, human population density, protected areas, and life zones on both occurrence and species richness. Human population density appeared as the strongest predictor of naturalized species occurrence and richness, highlighting the role of human activity in promoting invasions. The annual mean temperature and biodiversity integrity were negatively associated with the occurrence and species richness. Tropical rainforests and other humid life zones have more naturalized species than expected by chance. Protected areas had fewer naturalized species overall, but a higher species: observation ratio, showing both conservation value and vulnerability. Naturalized rare species, in terms of the number of records, were found outside protected zones, particularly in disturbed and urbanized areas. Our findings highlight the need for early detection, targeted management, and strengthened protection strategies, especially in mid-elevation zones and densely populated areas. By identifying key environmental and anthropogenic drivers and the most affected regions, this study offers actionable insights for conservation planning and invasive species management in one of the world’s most biodiverse and socio-environmentally vulnerable regions. Key words: Invasion patterns, invasive species, life zones, population density, protected areas Academic editor: Anibal Pauchard Received: 29 April 2025 Accepted: 23 November 2025 Published: 19 December 2025 NeoBiota 104: 361–379 (2025) DOI: 10.3897/neobiota.104.157379 Advancing research on alien species and biological invasions A peer-reviewed open-access journal NeoBiota This article is part of: Developing lists of alien taxa in the Global South: workflows, protocols, processes, and experiences Edited by John Wilson, Michele Dechoum, Katelyn Faulkner, Barbara Langdon, Shyama Pagad, Aníbal Pauchard, Hanno Seebens, Tsungai Zengeya, Silvía Ziller
362 NeoBiota 104: 361–379 (2025), DOI: 10.3897/neobiota.104.157379 Eduardo Chacón-Madrigal et al.: Distribution of naturalized plants in Central America Introduction Naturalized plant species, those that establish self-sustaining populations outside their native range, play a complex and significant role in global change and biodiversity conservation (IPBES 2023). When naturalized species become invasive, they can severely affect ecosystem processes, economic stability, food and water security, human health, and even cultural identities (Pimentel et al. 2001; Clements et al. 2021; Heringer et al. 2021; Bacher et al. 2023). Climate, land use, economic activity, and human activity determine the diversity and abundance of naturalized plants in the geographical context (Carboni et al. 2010; Pyšek et al. 2010; Wohlwend et al. 2021; Dong et al. 2025). Understanding the drivers and distribution patterns of naturalized species is essential for developing effective management strategies to conserve biodiversity and promote ecological resilience. Although global studies have advanced invasion ecology (Carboni et al. 2010; Pysek et al. 2010), a critical gap remains in characterizing these patterns and relationships within regions that harbor exceptionally diverse native biotas, where the impacts of invasive species may be particularly significant. Central America, with its high biodiversity and unique ecological contexts, provides an ideal setting to examine how environmental factors and human activities interact to shape naturalization patterns. With approximately 20,000 plant species and 23% endemism (Govaerts 2024), Central America is a biodiversity hotspot (Coates 1999). However, the region also faces a high degree of ecological and socioeconomic vulnerability due to a long history of colonialism, political instability, inequality, poverty, inadequate use of its territory, and rapid unplanned urbanization (Morales-Marroquín et al. 2022). These pressures and the limited investment in scientific research result in weak conservation policies and poor regulation of activities that affect native ecosystems (Harvey et al. 2005). For instance, managing non-native invasive and naturalized species is rarely addressed in environmental laws and policies across Central American countries (Chacón-Madrigal et al. 2022). Thus, empirical research on plant invasions in the region is still scarce compared to other regions. Recent studies have begun to address long-standing knowledge gaps in our understanding of plant invasions in Central America (see Avalos et al. 2021; Chacón-Madrigal et al. 2022; Rojas-Sandoval et al. 2023; MacVean and Zinn 2023). These efforts have provided valuable baseline data, including species inventories, introduction pathways, and invasion histories, and highlight the pressing need for more coordinated, region-wide research on plant invasions. A persistent challenge found across these studies is the lack of updated herbarium records and systematic botanical surveys focused specifically on non-native species. In an earlier work, we documented 1,228 species in the non-native flora of Central America, including 835 naturalized species and 393 casuals (Rojas-Sandoval et al. 2023). That study also showed major introduction pathways, economic uses, and country-specific patterns. However, this region still lacks fine-scale analyses of species distributions and the environmental and human factors influencing their spread. Addressing these gaps is crucial, as they limit the region’s ability to develop predictive tools and proactive strategies to manage invasive species and mitigate their ecological and socioeconomic impacts. In this study, we evaluate the distribution patterns of naturalized species across Central America with key environmental and anthropogenic factors. Specifically, we assess the distribution of these species across ecosystems classified as life zones, Citation: Chacón-Madrigal E, Rojas-Sandoval J, FerrufinoAcosta L, Flores R, Galán P, MacVean A, Rodríguez-Delcid D, Saldivar-Gómez I, Ruiz Y, López O (2025) Geographical patterns in the distribution of naturalized plants in Central America. NeoBiota 104: 361– 379. https://doi.org/10.3897/ neobiota.104.157379
363 NeoBiota 104: 361–379 (2025), DOI: 10.3897/neobiota.104.157379 Eduardo Chacón-Madrigal et al.: Distribution of naturalized plants in Central America protected areas versus unprotected areas, biodiversity integrity, gradients of human population density, mean annual temperature (MAT), and annual precipitation (AP). We selected these variables because they represent well-established drivers of naturalization or invasibility, supported by both theoretical and empirical evidence (Pyšek et al. 2010; Wohlwend et al. 2021). Climatic variables (MAT and AP) influence naturalized species by determining environmental suitability; we predict higher richness in warmer, wetter areas (Pyšek et al. 2010; Wohlwend et al. 2021). Human population density reflects propagule pressure and disturbance, and it is expected to be positively associated with naturalization (Lockwood et al. 2005). Biodiversity integrity, in contrast, is expected to show a negative association, consistent with the biotic resistance hypothesis (Levine et al. 2004). Finally, protection status may reduce naturalization due to lower disturbance (Bhatta et al. 2025), though mixed evidence suggests that protected areas can also be susceptible to invasions (Hiley et al. 2023). By identifying the ecological and socioeconomic drivers of naturalized species distributions in Central America, we aim to conduct future research, guide policy development, and support evidence-based management strategies. Our findings offer crucial insight into the role of protected areas in buffering against biological invasions and keeping biodiversity, as well as the vulnerability of disturbed landscapes and the potential for climatic and demographic trends to shape future invasions. This research enhances the scientific understanding of biological invasions in the tropics. It offers practical guidance for preserving the ecological integrity of diverse landscapes in a region of global conservation significance. Methods Study site We analyzed all seven countries comprising Central America: Belize, Costa Rica, El Salvador, Guatemala, Honduras, Nicaragua, and Panama. These countries cover a total area of 525,300 km2 (World Bank 2022) and encompass a complex and diverse geography, including mountain ranges, volcanoes, plains, and significant bodies of water. Elevation ranges from sea level to 4,220 m a.s.l. at Tajumulco Volcano in Guatemala. The mean annual temperature is 24 °C, with average minimum temperatures ranging from 0 °C during the coldest month in the highlands to maximum temperatures of around 30 °C during the hottest month in the lowlands (Taylor and Alfaro 2005). Annual rainfall varies widely due to topographic and elevational gradients. For instance, El Salvador and parts of Guatemala, Honduras, and Nicaragua receive less than 1,000 mm of rainfall annually. In contrast, other regions in Guatemala, Panama, and Costa Rica receive more than 2,500 mm, reaching 5,000 mm/year in some areas of the Caribbean slope of Costa Rica and Nicaragua, as well as in the southern part of Costa Rica. The region has distinct wet and dry seasons, with the wet season typically extending from May to November (coinciding with increased hurricanes and tropical storm activity), while the dry season extends from December to April. The Caribbean coast is generally more humid and receives more rainfall than the Pacific coast (Taylor and Alfaro 2005). Central America is a narrow land bridge connecting North America and South America, separating the Pacific Ocean from the Caribbean Sea. Its formation began 60 Ma ago and the land bridge closed the interoceanic canal between Middle and Late Miocene, ca. 13-3 million years ago (Montes et al. 2015), allowing the
364 NeoBiota 104: 361–379 (2025), DOI: 10.3897/neobiota.104.157379 Eduardo Chacón-Madrigal et al.: Distribution of naturalized plants in Central America movement of previously isolated biota of North and South America in an extraordinary biogeographical event (Bagley and Johnson 2014; Leigh et al. 2014). As a corridor between two major biogeographical regions, it harbors ca. 7% of the world’s plant and animal species (Greenheck 2002). Despite its rich biodiversity and abundant natural resources, the region faces significant environmental challenges. It is considered a threatened biological hotspot with high conservation priority due to habitat degradation, soil erosion, water pollution, and climate change (Harvey et al. 2005). Deforestation, primarily driven by agriculture, is a significant concern. Substantial portions of the forest have been cleared for cattle ranching, palm oil plantations, and other agricultural activities. Estimates suggest that less than 20% of the region keeps dense forest cover, and much of the remaining forest is highly fragmented or at risk of conversion (Hoang and Kanemoto 2021). The population of Central America is ca. 50 million people. The country with the highest population is Guatemala, and the smallest is Belize (Table 1). The region has a high population growth rate, along with accelerating urbanization and migration. Economically, Central America relies heavily on agriculture, with coffee, bananas, sugarcane, and other crops as key exports (Grau and Aide 2008). However, in recent years, the economy has shifted towards manufacturing and service industries, particularly in Costa Rica and Panama (Ulku and Zaourak 2021). According to the World Bank, nearly 30% of the population lives below the poverty line, with rural areas and Indigenous communities being especially vulnerable (ECLAC 2024). The region is highly vulnerable to climate change due to its geographic location, socioeconomic inequalities, agricultural dependence, low educational levels, and weak infrastructure (Castellanos et al. 2022). Expected impacts of climate change include more frequent and intense hurricanes, floods, landslides, and droughts (Hannah et al. 2017; Donatti et al. 2019), which can lead to potential loss of life, displacement, and economic disruption. Rising sea levels may threaten coastal communities and infrastructure, and growing water scarcity could further worsen food insecurity (Hagen et al. 2022). Due to its strategic geographic position, Central America is one of the most important global trade routes (e.g., the Panama Canal). As a result, the region is undergoing rapid economic development and increased international trade (Kerf 2021), with ecological consequences including pollution, resource degradation, and overexploitation. Infrastructure expansion, such as roads, ports, and urban areas, further contributes to environmental disturbances (ECLAC 2015) and may ease the introduction and spread of non-native species. Table 1. Geographical and socioeconomic data of the Central American countries. Land area (km2), human population (people), Gross Domestic Product 2022 (G C2019-24 DP), number of occurrence records, and naturalized plant species. Country Land area (km2) Population GDP (Billions US$) Occurrences Species Belize 22299.42 405272 2.8305 909 138 Costa Rica 51144.33 5180829 69.244 19179 545 El Salvador 20539.22 6336392 31.989 3261 364 Guatemala 108811.24 17357886 95.003 2904 273 Honduras 112236.73 10432860 31.426 3416 283 Nicaragua 128691.43 6948392 15.65 5373 287 Panama 74530.31 4408581 76.523 7616 286
365 NeoBiota 104: 361–379 (2025), DOI: 10.3897/neobiota.104.157379 Eduardo Chacón-Madrigal et al.: Distribution of naturalized plants in Central America Data collection and analysis This study builds on an open-access dataset of 1,228 non-native plant species occurring across Central America compiled by Rojas-Sandoval et al. (2023) as part of the FINCA Project, which includes a detailed account of the taxonomic composition of alien flora in this region (Flora Introduced and Naturalized in Central America: https://finca.collaboration.uconn.edu/). The non-native species list for each country has been generated and reviewed by botanical experts from each country who work in local herbaria or botanical gardens, using information from herbaria and local and regional treatments, including Flora Mesoamericana (https://legacy.tropicos.org/Project/FM), Manual de Plantas de Costa Rica (https://legacy.tropicos.org/Project/Costa Rica), Flora de Nicaragua (https://legacy.tropicos.org/Project/FN), and Panama Checklist (https://legacy.tropicos.org/ Project/PAC). Also, local experts added species based on their experience. However, all species included were confirmed by records containing at least one voucher specimen in a herbarium. We standardized species names using the taxonomic name resolution service website (https://tnrs.biendata.org/) (Boyle et al. 2013) according to World Flora Online (http://www.worldfloraonline.org/). We used that list to compile records for each country, sourced from the Global Biodiversity Information Facility (GBIF, http://www.gbif.org) and local herbarium collections (CR, USJ) (Chacón and Saborío-R 2006; Lopez 2012; MacVean and Zinn 2023). All observations are based entirely on secondary sources. As GBIF’s data providers include iNaturalist (www.inaturalist.org), it contains data collected by citizen science. However, GBIF only includes observations classified as “research grade” by iNaturalist, i.e., those with taxonomic identification confirmed by two or three users and with coordinates and date in the record. In the FINCA dataset, we classified species as either casual or naturalized following Richardson et al. (2000) and Blackburn et al. (2011). We classified species as naturalized based on any of the following criteria: expert opinion for each country, information in local treatments (as mentioned above), or the number of records for each species in each country, and considered a species naturalized if the number exceeded 10 records. We selected from the original GBIF dataset only the occurrences of the species classified as naturalized. For all occurrences, we applied the workflow by Seebens and Kaplan (2022), which includes data-cleaning procedures to eliminate errors in geographic coordinates and duplicate records. Because an exotic species may be native in another country within the same region, we restricted searches to records from countries where it was listed as exotic. The species list used for each country is available in the repository https://zenodo.org/records/16748538. Central America comprises only seven countries, limiting the statistical power of analyses that treat countries as the sampling units. To address this, we analyzed patterns using a grid-based approach when variables were available at comparable spatial resolutions. Additionally, we performed country-level correlation analyses to capture general patterns. Specifically, we examined correlations between the number of occurrences and species per country, as well as among three national-level variables: land area (km2), population size, and Gross Domestic Product (GDP in billions of US dollars) for the year 2022, which is the latest year with data available for all countries. We obtained country-level data from the World Bank Data (World Bank 2022, 2025). We calculate Pearson correlation coefficients using R software (R Development Core Team 2022).
366 NeoBiota 104: 361–379 (2025), DOI: 10.3897/neobiota.104.157379 Eduardo Chacón-Madrigal et al.: Distribution of naturalized plants in Central America With a grid-based approach, we use regression analyses. For independent variables, we use annual mean temperature (AMT), annual precipitation (AP), human population density (human population), and a biodiversity integrity index. Climate data (AMT and AP) at 5-minute resolution were obtained from WorldClim (Fick and Hijmans 2017). Human population density data were obtained from WorldPop Hub at 1 km resolution (https://hub.worldpop.org/). We obtained biodiversity integrity index data from Gassert et al. (2022) at a 1 km resolution. We resampled all datasets in a grid with a 5-minute resolution using bilinear interpolation with the terra package (Hijmans 2023), ensuring a uniform resolution. To construct our dependent variables, we rasterized the occurrence data of naturalized species in a grid with a 5-minute resolution, estimating both the number of naturalized species records and the number of species per cell (Both raster datasets are available in https://zenodo.org/records/16748538). We used a mixed-effect regression model to assess the influence of independent variables on the number of records. Latitude and longitude were included as random effects using a Matérn correlation matrix to control spatial autocorrelation. The country was included as a fixed effect. We used the fitme function from the spaMM package (Rousset and Ferdy 2014). Species richness was analyzed using a similar model, incorporating the number of records as a fixed effect. As expected, we found that the number of species per cell was dependent on the number of records. This was evaluated and verified through a regression analysis between the number of occurrences and the number of species. For both models, we used a Poisson distribution for the response variable. We generated maps of predicted records and predicted richness for each model. To evaluate whether life zones influence the number of occurrences of naturalized species, we used a contemporary map of life zones from Elsen et al. (2021) with a resolution of 30 seconds. We calculated the number of cells with naturalized species within each life zone. We compared the number of cells expected for each life zone with a normal distribution of values estimated by randomizing 1000 times the overall number of cells with naturalized species (n = 14,125) across all geographical space. We counted the number of cells within each life zone each time. We plotted the density of the distribution of expected values by random and contrasted it with the observed value per life zone. We also use the climenv package (Tsakalos et al. 2023) to visualize species occurrences within a climatic space diagram, comparing them to the overall availability in Central America. We obtained and joined vector shapefiles of protected areas for each Central American country (Suppl. material 3) and then rasterized at a 30-second resolution. We obtained protected area boundaries from national datasets rather than the UNEP-WCMC database, as the latter omits several protected areas known to exist in the region; national datasets provided more complete and up-to-date coverage for all countries. We calculated the number of observations and species inside and outside protected areas and determined the number of cells with naturalized species for each category (i.e., with and without protection). We compared proportions using a Chi-square test of independence with Yates’s correction. We also used rarefaction analysis with Hill numbers of order 0, 1, and 2 to compare species richness between protected and unprotected cells using the iNEXT package (Chao et al. 2014). We chose this analysis because it is not biased by sampling effort. We conducted all the analyses in R (R Development Core Team 2022).
367 NeoBiota 104: 361–379 (2025), DOI: 10.3897/neobiota.104.157379 Eduardo Chacón-Madrigal et al.: Distribution of naturalized plants in Central America Results We compiled 42,658 occurrence records for 751 naturalized plant species across Central America. The number of species was positively associated with the number of records (Suppl. material 1). However, records were unevenly distributed among countries; Costa Rica accounted for the highest number of species (19,179), while Belize had the fewest (909; Table 1). Neither the number of occurrence records nor the number of species per country was correlated with land area, population size, or GDP (Suppl. material 2). At the spatial cell level, population density, AMT, and biodiversity integrity significantly explained the number of occurrence records (Table 2, Fig. 1). Cells with higher population density had more occurrences of naturalized species. In contrast, AMT and the biodiversity integrity index negatively affect the number of records (Table 2, Fig. 1). The species richness of naturalized plants is significantly influenced by population density, AMT, and the biodiversity integrity index (Table 2). Specifically, species richness increased with population density, while it decreased in areas with higher AMT and higher biodiversity integrity values (Table 2). When analyzing life zones, we found no significant deviation from random expectations in the cells observed with naturalized species for tropical moist, tropical dry, and premontane moist forests (Figs 2, 3). In contrast, all other life zones showed a higher-than-expected number of cells with naturalized species. Figure 1. Maps of the number of occurrence records (A) and species richness (B) of naturalized species. Predicted number of occurrences (C) is based on population density, MAT, and biodiversity integrity, while predicted species richness (D) is based on population density, temperature, and biodiversity integrity, standardized to 300 occurrences.
368 NeoBiota 104: 361–379 (2025), DOI: 10.3897/neobiota.104.157379 Eduardo Chacón-Madrigal et al.: Distribution of naturalized plants in Central America The Tropical premontane wet forest had the largest positive deviation from random expectations (Figs 2, 3), with nearly 4% more occupied cells than expected. In contrast, Tropical moist forest and Tropical dry forest had fewer cells with naturalized species than expected (Fig. 3). On protected areas, we found 33,211 occurrences of naturalized species outside protected areas, compared to 8755 observations inside. In total, 729 naturalized species were recorded outside protected areas, while 540 were found within protected areas (Fig. 4). Most species occurring exclusively outside protected areas were seen at low frequencies (Fig. 4). No significant differences were observed in the number of common or frequent species between protected and unprotected Table 2. Coefficients of the mixed-effect regression model predicting the number of occurrences and the species richness of naturalized species in Central America using as predictors human population, annual mean temperature, annual precipitation, and biodiversity integrity index (see methods). Variable Estimate SE df Chi sq. value Pr (>F) Number of occurrences (Intercept) 5.365 0.569 1 88.93 <0.001** Population density 0.0006 0.00007 1 77.74 <0.001*** Annual mean temperature -0.072 0.0137 1 27.882 <0.001*** Annual precipitation 0.000003 0.00009 1 0.002 0.96 Biodiversity integrity -2.90 0.402 1 52.167 <0.001*** Species richness (Intercept) 4.848 0.499 1 94.28 <0.001*** Population density 0.0005 0.00006 1 66.23 <0.001*** Annual mean temperature -0.0609 0.0121 1 25.21 <0.001*** Annual precipitation 0.000023 0.00008 1 0.085 0.77 Biodiversity integrity -2.827 0.354 1 63.88 <0.001*** Figure 2. Holdridge life zones scheme based on biotemperature, annual precipitation, and potential evapotranspiration ratio (A). Environmental space is available within the Central American Region with (B) in the dimensions of the Holdridge life zone variables. Environmental space where naturalized species have been recorded within the dimensions of the Holdridge life zones (C). The blue points indicate low-density areas, and yellow points indicate high-density areas of species records.
369 NeoBiota 104: 361–379 (2025), DOI: 10.3897/neobiota.104.157379 Eduardo Chacón-Madrigal et al.: Distribution of naturalized plants in Central America areas (Fig. 4). The proportion of cells with naturalized species was significantly different between protected and unprotected areas (χ2 = 296, df = 2, p < 0.001), with higher numbers of cells with naturalized species occurring outside protected areas (0.029 cells with naturalized species in unprotected areas/overall number of cells in unprotected areas) than inside (0.019 cells with naturalized species in protected areas/ overall number of cells in protected areas). However, the observation density was similar, with approximately 6.9 occurrences of naturalized species per 100 km2 outside protected areas versus approximately 6.5 occurrences of naturalized species per 100 km2 inside. Interestingly, the ratio of species to observations was higher within protected areas, averaging approximately six species per 100 records, compared to approximately two species per 100 records outside protected areas. Figure 3. Observed versus expected cells with naturalized species occurrences across Holdridge life zones in Central America. Dots indicate the observed number of occupied cells, while density areas represent the expected distribution obtained by randomizing the same number of occupied cells (n = 14,125) 1,000 times across all life zones. Red dots indicate values higher than expected, blue dots lower than expected, and black dots values consistent with expectations. Figure 4. Rarefaction curves of the naturalized plant species in Central America occurring within and outside protected areas based on Hill’s numbers 0 (species richness), 1 (common species), and 2 (frequent species). Curves represent the estimated diversity for each category, allowing a comparison of species accumulation and community structure between protected and unprotected areas. The shadow areas represent standard errors.
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379 NeoBiota 104: 361–379 (2025), DOI: 10.3897/neobiota.104.157379 Eduardo Chacón-Madrigal et al.: Distribution of naturalized plants in Central America Supplementary material 3 Sources of layers of Protected Areas for each Central American country used in the analysis Authors: Eduardo Chacón-Madrigal, Julissa Rojas-Sandoval, Lilian Ferrufino-Acosta, Rodolfo Flores, Pablo Galán, AnaLu MacVean, Dagoberto Rodríguez-Delcid, Iris Saldivar-Gómez, Yader Ruiz, Omar López Data type: pdf Copyright notice: This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited. Link: https://doi.org/10.3897/neobiota.104.157379.suppl3