Urban forests as habitats for vascular epiphytes and allied terrestrial plants

Urban forests as habitats for vascular epiphytes and allied terrestrial plants | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Research Article Urban forests as habitats for vascular epiphytes and allied terrestrial plants Maria Berenice Jarquin-Pacheco, Samaria Armenta-Montero, Jazmín Contreras-López, and 1 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-6116120/v1 This work is licensed under a CC BY 4.0 License Status: Published Journal Publication published 26 Jun, 2025 Read the published version in Urban Ecosystems → Version 1 posted 10 You are reading this latest preprint version Abstract Epiphytic plants and allied terrestrial groups, which are sensitive to changes in humidity and temperature, play a crucial role in understanding the dynamics of ecosystem disruption caused by human activity impact. In the conurbation of Xalapa-Banderilla, Veracruz, Mexico, urban and peri-urban forests have different levels of disturbance conditions. This study aimed to analyze the patterns of vascular epiphytes and related terrestrial plants in urban forests with varying transformation histories in Xalapa, located in the central region of Veracruz Mexico. Five sampling plots were established in each forest (three urban and two peri-urban) where the richness of these groups was recorded. In a sample of 1 ha, 103 species distributed among 58 genera and 22 families were recorded, with ferns being the most represented. The peri-urban forest “Clavijero” exhibited the highest species richness compared to the others. Overall, heterogeneity in species composition was observed between sites, being lower when comparing only urban forests. In peri-urban forests, species considered indicator species were recorded, while in urban forests, mostly generalist species adapted to stressful conditions were recorded. The urban forests studied are reservoirs of diversity, highlighting the importance of forests in the periphery that shelter rare and conservation indicator species, suggesting that the original environmental conditions are still being maintained for the benefit of diversity in general. bromeliads ecological indicators ferns peri-urban forests urban green areas Figures Figure 1 Figure 2 Figure 3 Figure 4 Introduction Epiphytic plants are an important component of plant communities in the tropics, especially in humid mountain forests. These host a great diversity, accounting for up to 50% of the leaf biomass, and, in some forests, can represent up to 25% of the total vascular species (Cach-Pérez et al. 2014 ; Gotsh et al. 2015). Vascular epiphytes are mostly represented by species of ferns, peperomias, bromeliads, aroids and orchids (Zotz 2013 ; Krömer et al. 2014 ). These main groups can occasionally grow terrestrially in ecosystems, mainly ferns, and to a lesser extent, Araceae, orchids, and Peperomia. These plants provide habitat for organisms such as insects, birds, and amphibians, offering resources such as food and shelter; they also participate in canopy interactions (Zotz 2013 ; González and Ceballos 2021 ; Mendoza et al. 2023 ). However, these types of plants depend on atmospheric conditions like humidity and temperature; that is, they are sensitive to changes in their habitat, which can lead to shifts in the composition of their communities and a reduction in their richness. Therefore, they can serve as bioindicators of climate change and ecological damage (Köster et al. 2009 ; Larrea and Werner 2010 ; Köster et al. 2011 ; Reina-Rodríguez et al. 2017 ; Carvajal-Hernández et al. 2017 ). Vascular epiphytes are one of the most important elements in humid mountain forests (HMF) (Gotsh et al. 2015). In Mexico, this type of vegetation supports a significant diversity of plants due to the transition zone between two biogeographic regions that converge in the country (Rzedowski 1991 ; 2006 ). One of the most vital ecosystem services provided by this type of vegetation is the availability of water resources, derived from precipitation and fog, which is essential for the establishment of groups dependent on humid conditions, such as vascular epiphytes (Krömer et al. 2014 ; Muñoz-Villers et al. 2015 ). Despite its ecological importance, this forest is one of those currently presents strong anthropogenic pressure, primarily due to changes in land use for creating agricultural land, livestock, and urban development (Challenger and Caballero 1998 ; Muñoz-Villers et al. 2015 ; Rosas et al. 2019 ). Recent studies have revealed that over a period of 17 years (1999–2016), a 47.2% reduction in this type of vegetation has been observed, highlighting the current issues faced by the HMF (Espinoza-Guzmán et al. 2019 ). The loss, reduction, and/or fragmentation of this forest type leads to the decline of species in general and vascular epiphytes in particular (Paredes et al. 2021 ). One of the areas in Mexico that still maintains remnants of the HMF within the city and in its surroundings is the conurbation of Xalapa, located in the central part of the State of Veracruz, along the slope of the Atlantic Ocean (Williams-Linera et al. 2002 ). However, urban growth in Xalapa has occurred at the expense of rural areas and natural ecosystems (Von Thaden et al. 2021), modifying 90% of the original HMF (Williams-Linera et al. 2002 ). These transformations have altered the structure and functioning of ecosystems, creating habitat fragmentation, causing a lack of connectivity between them (Von Thaden et al. 2021), and inducing the loss of plant species. However, the presence of forest fragments within urban areas can still sustain some biodiversity in small islands of vegetation (Rosas et al. 2019 ). Given the threat posed by biodiversity loss and its combination with overall urban growth, priority has been given to preserving areas with some type of vegetation, such as urban forests (UF), which are defined as areas with native and secondary vegetation, existing within transformed ecosystems in cities (Zewdie and Tegegne 2019 ). In that sense, the city of Xalapa is still recognized as a “green city" because of the presence of areas with remnant forests of native vegetation and/or protected natural areas that exist in a fragmented manner and at various stages of ecological succession (Williams-Linera et al. 2002 ; Von Thaden et al. 2021), which, on a smaller scale, also include parks and gardens medians. UF provides a variety of benefits to cities, including the conservation of environmental services (such as carbon sequestration, microclimate regulation, and mitigation of weather events), protection of flora and fauna, and space for recreation (Heidt and Neef 2008 ; WHO 2017; Lemoine-Rodriguez et al. 2019). Urban forests can support high levels of biodiversity, including endangered species (Alvey 2006 ). This richness is highly dependent on the conservation status of the UF (Pesola et al. 2017 ) and the existence of remnants of native vegetation (Kowarik 2011 ). For instance, forests adjacent to cities tend to support a greater diversity of medium-sized mammals than more urbanized areas (Tee et al. 2018 ). Similarly, a 10–30% increase in native vegetation cover has been found to support a rise in the presence of bats, birds, and insects (Threlfall et al. 2017 ) by providing shelter and food (Mohamad et al 2013 ). However, different taxa respond distinctly to urban spaces (MacGregor-Fors et al. 2016 ), as evident in the case of epiphytes (Santana et al. 2017 ), whose presence is associated with trees of larger diameter and height (Izuddin and Webb 2015 ). Considering that the conurbation of Xalapa contains forest fragments surrounded by urban development on its periphery, there is an opportunity to understand the patterns of vascular epiphyte diversity under various anthropogenic pressures typical of urban dynamics, including isolation, modifications to tree structure, biodiversity extraction, microclimatic changes, soil degradation, and the introduction of exotic flora. Therefore, the aim is to examine the patterns of vascular epiphytes in urban forests with varying transformation histories in Xalapa, situated in central Veracruz, Mexico. This study contributes to recognizing the essential role of vegetation remnants in the city and their significance as a refuge for the diversity of vascular epiphytes and associated terrestrial plants. Materials and Methods Study area The study was carried out in the conurbation of the municipalities of Xalapa and Banderilla, located in the central region of the State of Veracruz in southeastern Mexico (Fig. 1 ). The area contains remnants of humid mountain forest, consisting of patches of vegetation of various sizes. The surface area of the urban region of Xalapa encompasses approximately 60 km², where a mixture of native and exotic species has been recently observed, similar to what occurs in various cities (Falfán and MacGregor-Fors 2016 ). The altitude ranges from 1,120 to 1,700 meters, with the predominant climates being semi-warm humid, characterized by abundant summer rainfall, and semi-warm humid with year-round rainfall. The temperature range oscillates between 18 and 24°C, while precipitation levels vary from 1,100 to 1,600 mm annually (SIEGVER 2020; Jara-Toto et al. 2023 ). The study was conducted in five urban forests, three of which are located within the city of Xalapa: 1) Protected Natural Area “El Tejar Garnica” (TEJ) (133 ha, altitude of 1310 m), corresponding to the polygon known as Parque Natura; 2) Protected Natural Area “Macuiltépetl Ecological Park” (MAC) (31 ha; 1600 masl); 3) “Campus para la Cultura, las Artes y el Deporte” (CAD) primarily in its section known as Agrobosque Kaná (28 ha; 1366 masl); 4) “ Francisco Xavier Clavijero Ecological Reserve” (CLA) (59.85 ha; 1362 masl), hereafter referred to as Clavijero Park; and 5) “La Martinica” Natural Protected Area (MAR) (52.30 ha; 1599 masl). The last two sites are situated on the periphery of the urban areas of the municipalities of Xalapa and Banderilla. Of all the study sites, only the CAD Campus is not officially classified by the government as a protected natural area; however, it is under the protection of the Universidad Veracruzana as an environmental management unit. The remaining sites are protected natural areas administered by different levels of government; the vegetation they harbor are remnants of humid mountain forest with different degrees of anthropogenic disturbance and different transformation histories (Jara-Toto et al. 2023 ). In the forest fragments within the city (El Tejar Garnica, Campus CAD, and Parque Macuiltépetl), the vegetation structure is mainly composed of tree species from secondary vegetation derived from the HMF, such as Dendropanax arboreus , Piper amalago , and Heliocarpus appendiculatus (Jara-Toto et al. et al. 2023), where the presence of native and introduced plant species is also recorded. In the peri-urban area of Clavijero Park, typical HMF species exhibit a higher state of maturity, including Clethra macrophylla, Liquidambar styraciflua, Quercus lancifolia , and Turpinia insignis (Williams-Linera et al. 2013 ). Meanwhile, in La Martinica, there is a mix of primary and secondary HMF species at various levels of conservation. Field work Sampling was conducted by establishing five plots measuring 20 x 20 m (400 m²) at each of the five sites mentioned earlier, following the methodology proposed by Kessler and Bach ( 1999 ). As a result, a total of 25 plots (1 ha) were sampled across the five areas. Each plot was selected to maintain a distance from water bodies and prevent the establishment of a well-defined tree and shrub layer. In each plot, data on the incidence of understory species were recorded, including both holoepiphytes and related groups found on the ground or on fallen trunks and branches (e.g., ferns, Araceae, peperomias, and terrestrial orchids). This approach was adopted to ensure adequate representation of specific groups seen as indicators of mountain ecosystem quality, such as certain ferns found in terrestrial habitats. However, some species are almost exclusively epiphytes, like the genus Tillandsia . This strategy aimed to increase the number of species recorded, providing a more comprehensive view of the plant community within the studied ecosystem. Furthermore, certain plants, such as specific ferns, are recognized as indicators of habitat quality (Della and Falkenberg, 2019 ). Including both epiphytes and terrestrial species allowed for better identification of the groups that reflect the condition and status of the ecosystems. The observed species were collected and herborized according to the methodology of Lot and Chiang ( 1986 ) and Lorea and Riba ( 1990 ). Subsequently, dichotomous keys were used for their identification, and the specimens were deposited in the herbarium of the Universidad Veracruzana in Xalapa (XALU), with duplicates stored in the National Herbarium (MEXU) of the Institute of Biology at the Universidad Autónoma de México. The resulting floristic list was organized at the family level following APG IV ( 2016 ) for angiosperms and PPGI ( 2016 ) for ferns. The scientific names of each species were verified using the Tropics.org database ( https://www.tropicos.org/home ). The IUCN Red List was consulted to compile species in any risk category. At the national level, the Official Mexican Standard (NOM-059-SEMARNAT-2010) was reviewed to identify species that fall into a risk category within Mexican territory. Data analysis To assess alpha diversity, the data were analyzed using the online version of iNEXT software to generate rarefaction curves that evaluate sampling effort and calculate Hill numbers to measure diversity at each study site (Chao et al. 2014 ; 2016 ). The Kruskal-Wallis test was conducted to confirm differences in total species richness across all sites, followed by Dunn's post hoc test to assess the differences between each site. For beta diversity, Whittaker's species turnover index was calculated. These analyses were conducted using Past 4.04 (Hammer et al. 2001 ; Koleff et al. 2003 ; Tuomisto 2010 ). A similarity dendrogram was also created to group the various urban forests based on their floristic composition, utilizing the Bray-Curtis distance matrix through the Vegan package in RStudio. Additionally, a one-way similarity analysis (ANOSIM) was conducted using the Bray-Curtis similarity index to confirm statistical differences in species composition among the sampled communities. The similarity percentage (SIMPER) was calculated to identify the contribution of different species to the composition of the communities studied. To evaluate whether certain species are indicators of a site, IndVal analysis was performed using the “indicspecies” package in R (Dufrêne and Legendre 1997 ). This analysis focuses on specificity and fidelity, where values near 1 signify that species effectively indicate the habitat, while very low (near 0) values denote rare species (Martín-Regalado 2019 ). RESULTS Species richness In a total sampling of 1 hectare, 103 species were recorded across 58 genera and 22 families (Annex 1). Of these, 55 are epiphytic species (35 are angiosperms, while the rest are ferns), and 48 species represent related groups found on land. Among the ten best-represented families in the entire study, six are ferns. The families with the highest richness are as follows: Polypodiaceae (16), Orchidaceae (15), and Bromeliaceae (11); with the genus Tillandsia being the most represented with nine species, followed by Thelypteris with seven, and the genera Peperomia and Pleopeltis , each with six species. Therefore, the most abundant group in terms of the number of records consists of ferns (58%), including both epiphytic and terrestrial species, followed by orchids (15%) and bromeliads (11%). Of the total number of species, four were registered in the NOM-059-SEMARNAT-2010, three were in the IUCN Red List, and 14 were endemic to Mexico. The richness estimated from the sampling coverage obtained with the iNEXT software indicates that the sampling effort in the five urban forests studied is adequate (Fig. 2 a). Additionally, Hill's numbers show that Clavijero (CLA) had the highest richness (q0) and diversity (q1, q2), while two areas within the city (TEJ and CAD) exhibited the lowest values (Fig. 2 b). The comparison of richness among the study sites indicates that Clavijero Park is the peri-urban forest with the highest richness (64 species), followed by La Martinica with 49 species, while the CAD Campus had the lowest richness (Table 1 ). These data were supported by the results of the Kruskal-Wallis test, which demonstrated significant differences among the sampled urban forests (H = 17.65, Hc = 24.03, p < 0.05), indicating that at least one site significantly differs in richness. A post-hoc Dunn's test with Bonferroni correction was then performed to identify differences between the sites. The results indicated that Clavijero Park has significantly higher richness than other sites (p < 0.05). However, no significant differences in richness were found among the other sampled forests (Table 1 , Fig. 3 ). Table 1 Species richness by site: Urban areas include Tejar Garnica, CAD Campus, and Macuiltépetl Park; peri-urban areas encompass Clavijero Park in the Municipality of Xalapa and La Martinica in the Municipality of Banderilla, Veracruz, Mexico. Site Species richness (Ephytes/Terrestrials) Family richness Most representative families Location Tejar Garnica 35 (20/15) 11 Polypodiaceae (7), Orchidaceae (7), Bromeliaceae (4) y Araceae (4) Urban Macuiltépetl Park 41 (23/18) 17 Polypodiaceae (11), Orchidaceae (4) y Bromeliaceae (4) Urban Campus CAD 34 (16/18) 14 Polypodiaceae (5), Thelypteridaceae(5) y Bromeliaceae (5) Urban La Martinica 49 (30/19) 17 Polypodiaceae (10), Bromeliaceae (6), Orchidaceae (5) y Piperaceae (5) Peri-urban Clavijero Park 64 (38/26) 20 Polypodiaceae (12), Orchidaceae (10) y Bromeliaceae (9) Peri-urban Species turnover The Whittaker index results indicate that the peri-urban area of La Martinica has the highest species turnover compared to urban areas. Additionally, the peri-urban areas show index values exceeding 50% among themselves, and this notable species turnover is also observed between the peri-urban and urban areas. Urban areas exhibit smaller differences in species composition when compared to one another (Table 2 ). Regarding the floristic similarity among the analyzed urban forests, the similarity dendrogram shows the division into three groups. One group includes the inner-city sites Campus CAD and El Tejar Garnica, while the other group consists of the two peri-urban areas. The species from Macuiltépetl Park, which is situated in the center of the urban area, are partly grouped with the peri-urban forests, creating a subgroup with Clavijero Park, while another part forms a separate group (Fig. 4 ). On the other hand, the ANOSIM performed with the Bray-Curtis similarity index indicates differences in the floristic composition of epiphytes and their terrestrial counterparts among the sampled urban forests (R = 0.8398; P < 0.05). Paired ANOSIM comparisons show differences in composition at all sites, with values of P < 0.05. Table 2 Whittaker's beta diversity index for the sampled sites of the Xalapa-Banderilla conurbation, Veracruz, Mexico. Tejar Garnica Macuiltépetl Park Campus CAD La Martinica Clavijero Park Tejar Garnica 0 0.474 0.450 0.643 0.535 Macuiltépetl Park 0 0.545 0.622 0.429 Campus CAD 0 0.518 0.490 La Martinica 0 0.522 Clavijero Park 0 Values of 0 represent equality and 1 signifies complete difference . According to the SIMPER analysis, the first 20 species that contribute the most to dissimilarity (representing 19% of the total registered species) account for 39% of the variation in species composition recorded across the five sites. Meanwhile, 40 species represent 68% of the contribution (Table 3 ). Ferns are the most numerous group in the study overall, but they do not significantly contribute to the changes in composition. In contrast, the Bromeliaceae family, mainly species from the genus Tillandsia , does have a substantial impact; among the 20 species with the greatest contribution, they account for 16% of the richness. Table 3 Species that, according to the SIMPER analysis, contribute more to the dissimilarity of epiphytic plant communities and their terrestrial allied in the urban forests of Xalapa-Banderilla conurbation, Veracruz, Mexico. Species Average dissimilarity Contribution (%) Cumulative (%) Tillandsia schiedeana 1.467 2.258 2.258 Tillandsia deppeana 1.457 2.244 4.502 Pteris quadriaurita 1.405 2.163 6.665 Syngonium sp. 1.396 2.149 8.815 Catopsis nutans 1.374 2.116 10.93 Polypodium conterminans 1.373 2.115 13.05 Tillandsia butzii 1.352 2.082 15.13 Tillandsia multicaulis 1.328 2.046 17.17 Tillandsia juncea 1.31 2.017 19.19 Pteris orizabae 1.294 1.993 21.18 Cyclopogon luteo-albus 1.29 1.987 23.17 Peperomia glabella 1.283 1.976 25.15 Pleopeltis polypodioides 1.266 1.949 27.1 Alsophila firma 1.209 1.861 28.96 Selaginella stellata 1.207 1.859 30.82 Monstera deliciosa 1.196 1.842 32.66 Campylocentrum schiedei 1.194 1.839 34.5 Tillandsia polystachia 1.147 1.766 36.26 Selaginella martensii 1.141 1.756 38.02 Scaphyglottis minutiflora 1.119 1.724 39.74 Among the total number of species documented in the five urban forests, only 27 (26%) demonstrated an indication value based on the IndVal (Indicator Value Method) analysis. Of these, 10 species were associated with more than three urban and peri-urban forests (37%). Macuiltépetl Park and La Martinica had the highest number of indicator species, totaling four, while only one species was recorded as unique to CLA (Table 4 ). Table 4 Indicator species based on the IndVal analysis of epiphyte species and their allied terrestrial between urban (Tejar Garnica: TEJ; CAD Campus: CAD; Macuiltépetl Park: MAC) and peri-urban (Clavijero Park: CLA; La Martinica: MAR) forests in Xalapa-Banderilla conurbation, Veracruz, Mexico. In all instances, the species exhibited values of P<0.05. Species Site IndVal* Syngonium podophyllum TEJ, MAC, CAD 1 Tillandsia punctulata MAR 1 Scaphyglottis minutiflora MAC 0.966 Christella dentata TEJ, MAC, CAD, CLA 0.934 Tillandsia schiedeana TEJ, MAC, CLA 0.931 Tillandsia deppeana CAD, MAR, CLA 0.931 Tillandsia butzii MAR, CLA 0.929 Selaginella stellata CAD 0.918 Campylocentrum schiedei CAD, CLA 0.894 Polypodium conterminans MAC, MAR, CLA 0.894 Selaginella martensii MAC, CLA 0.894 Catopsis nutans TEJ, MAR, CLA 0.879 Pteris quadriaurita TEJ, MAC, CLA 0.879 Tillandsia juncea CAD, CLA 0.869 Tillandsia multicaulis MAC, MAR, CLA 0.856 Peperomia san-joseana MAC, MAR 0.837 Monstera deliciosa MAC 0.775 Adiantopsis radiata TEJ, CAD 0.775 Alsophila firma CAD, MAR, CLA 0.775 Catopsis sessiliflora MAC, CAD 0.775 Thelypteris pilosohispida MAR 0.775 Polypodium eatonii MAR 0.775 Blechnum schiedeanum MAR 0.775 Phanerophlebia remotispora MAC 0.775 Ctenitis melanosticta MAC 0.775 Tillandsia polystachia TEJ, MAC, CLA 0.730 Polypodium rhodopleuron CLA 0.725 * IndVal values indicate affinity to a specific forest type, ranging from 1 (indicating it is only recorded in that forest type) to 0 (indicating it is not recorded at all) . Discussion Out of the total species richness of epiphytic angiosperms reported for the state of Veracruz (569 species), only 6% was documented in this study (Krömer et al. 2020 ). Given that a high diversity of epiphytes (285 species) exists in the humid montane forests of the state, the species documented in the urban forests of Xalapa represent 12.2% of the total diversity found in the humid montane ecosystems of Veracruz (Krömer et al. 2020 ). Among the most well-represented groups, ferns, orchids, and bromeliads are notable as they contribute the highest number of epiphytic species in floristic inventories across the Neotropics (Zotz, 2013 ; Krömer et al. 2014 ; Furtado and Menini-Neto 2015; Villaseñor 2016; Santana et al. 2017 ). One aspect to highlight in this study is the low presence of species from groups such as orchids and bromeliads, which are typically the most diverse group in inventories of epiphytic plants (Benzing 1990 ; Kreft et al. 2004 ; Flores-Palacios and García-Franco 2006; Zotz 2013 ; Alzate-Q et al. 2019). In contrast, ferns were the group with the highest representation in our results, excluding terrestrial species. This pattern aligns with findings by Krömer et al. ( 2021 ) in central Veracruz, who observed that ferns dominate in quantity and cover in conserved forest areas, followed by orchids and bromeliads. In the urban and peri-urban forests studied, the presence of fourteen epiphytes endemic to Mexico was noted, constituting 0.12% of the endemic diversity of vascular plants identified in the country (Villaseñor 2016). Of these, four are protected under Mexican laws by being included in the Norma Oficial Mexicana, which oversees the protection of wild plant and animal species (NOM-059-SEMARNAT-2010), and three are listed in the IUCN Red List under some risk category. This finding shows that, despite human pressure from urban growth, these areas continue to serve as important refuges for native and endemic biodiversity. Epiphytes and related terrestrial groups (especially ferns) require high humidity conditions to establish themselves, which they find in forests with a higher degree of conservation; in this case, these conditions are maintained in peri-urban areas forests. A recent study indicates that sites like Clavijero Park exhibit more temperate microclimatic conditions, showing less daily variation in temperature and relative humidity throughout the year compared to other evaluated forests, particularly those located in the city's interior, which tend to be warmer and show greater fluctuations in these environmental variables (Landeros-López 2025 ). Therefore, the observed patterns of richness could be explained by these changes in the microclimate. Moreover, vegetation structure may also be an important factor. A previous study revealed that the forests within the interior of the city (TEJ and CAD) and one peri-urban forest (MAR) exhibited a higher dominance of secondary trees with smaller diameters. In contrast, the Clavijero peri-urban area predominantly featured species typical of mature forests in the region, characterized by larger diameters (Jara-Toto et al. 2023 ). Similarly, Macuiltépetl Park, situated in the center of the urban area, features trees with larger diameters and primary species (Figueroa-Solis et al. 2023). This contrasting situation is explained by the unique transformation histories of each site. In the most transformed areas, there were previously pastures or abandoned crops, whereas Clavijero Park contained remnants of larger vegetation with less anthropogenic impact. However, it is important to note that all sites exhibit some degree of anthropogenic influence due to their proximity to the city. The above situation is significant for interpreting the results of epiphytes and their terrestrial relatives because tree species and the structure they create in the forest play a crucial role in the establishment and persistence of species. Host characteristics, such as diameter, bark type, and branching, are essential for epiphyte establishment and influence the microclimatic conditions they create in the understory (Bonnet et al. 2007 ; Caglioni et al. 2012 ; González and Ceballos 2021 ). Additionally, tree age is a crucial factor in epiphyte diversity. Adhikari et al. ( 2021 ) suggest that tree age may serve as a significant indicator of epiphyte diversity, as older and larger diameter trees provide better conditions for the establishment of these species. A study conducted in the urban area of Xalapa revealed that the proportion of woody species is linked to the distribution of epiphytes, particularly bromeliads (Aoki-Gonçalves et al. 2023 ). This helps elucidate the pattern observed in the studied forests. At a site like Clavijero, having a better tree structure, along with its peripheral location, enhances connectivity and ecosystem continuity with other forest fragments, allowing for the movement and establishment of propagules. Furthermore, as demonstrated in other studies, structural diversity, along with improved microclimatic conditions and reduced anthropogenic pressure, promotes the establishment of these botanical groups, which could lead to greater species richness (Krömer et al. 2014 ; Krömer et al. 2021 ; Von Thaden et al. 2021; Parra-Sánchez and Banks-Leite 2022). In this peri-urban forest, which resembles a primary forest, epiphytes and their terrestrial relatives are finding optimal conditions for establishment, making this site an important source for species and germplasm conservation in the city's surrounding area Xalapa. In the case of La Martinica, which is a peri-urban area that currently experiences less anthropogenic pressure than the forests in the interior of the city, a greater richness was expected; however, the number of species is low, a situation that was also previously reported by Krömer and collaborators (2021). This may be due to the smaller diameter trees present in this forest, indicating that it is undergoing succession. This factor may influence the establishment and abundance of these plants (Lahoti et al. 2020 ; Jara-Toto et al. 2023 ), which supports the hypothesis of Woods and De Walt (2012), who argues that an increase in epiphyte species will be observed as forest age increases. Another factor that can be determinant in species richness in urban areas is the extraction of certain plants for ornamental purposes. In Xalapa, it's common to find orchids and bromeliads in local markets due to collector demand, which intensifies the pressure on these epiphytes (Flores-Palacios and Valencia-Díaz 2007 ; Toledo-Aceves et al. 2014 ; Krömer et al. 2020 ). Although the study sites are protected natural areas by official decree and extraction activities are prohibited, the entry of people for various extraction activities (such as soil, firewood, and plants) has been observed in some urban forests (personal observation). This phenomenon may partially explain the disappearance of certain groups, with ferns dominating their place, as some species are infrequently used for these purposes in the area due to current human activities and past extraction practices. On the other hand, urban forests (Macuiltépetl Park, Tejar Garnica, and Campus CAD) exhibit lower epiphyte richness. This could be attributed to their location within the urban infrastructure of Xalapa, as they are subject to territorial and functional simplification of their habitats (Acebey et al. 2003 ; Mickinney 2008; de Juana Aranzana 2015 ; Parra-Sánchez and Banks-Leite 2022). However, it is essential to understand the life history of these forests—the historical processes of degradation and their environmental impact—to properly grasp the low rates of observed species richness (Alvey 2006 ). For example, Macuiltepetl Park, despite being surrounded by urban sprawl, ranks third in epiphyte richness with 41 species, surpassing the CAD Campus and Tejar Garnica. This may be due to restoration activities, such as those carried out in 1977, which contributed to the arboreal enrichment of the forest. Likewise, conserving a slope in the northeastern area that maintains the original arboreal vegetation of the HMF could have been an important source of propagules for plant dispersal and ecosystem regeneration (Pavón 2019 ). These past actions may have preserved microclimatic conditions and facilitated the subsequent establishment of epiphytes prior to the rapid growth of the city’s outskirts. Regarding Tejar Garnica and the CAD Campus, which had the lowest epiphyte richness (35 and 34 species), both areas feature secondary vegetation derived from the HMF. Tejar Garnica was originally a coffee plantation and cattle pasture (Jara-Toto et al. 2023 ), while the CAD Campus was also a cattle ranching pasture, where less than half of the area was left untouched to follow its natural secondary succession process, while the remaining area was altered with the establishment of gardens mainly composed of exotic flora (Teodosio-Faustino et al. 2021 ). The tree structure of the succession and intervention processes likely affects species richness, as it lacks the structural complexity of the forest, which reduces the formation of microclimates vital for the establishment of epiphytes and their allied terrestrials. Recambio de especies The Whittaker index and cluster analysis results indicate species turnover between peri-urban and urban forests, particularly regarding TEJ and CAD, which show smaller differences between them. In line with this, the analysis of percentage similarity (SIMPER) enabled us to identify ferns and bromeliads as the primary groups contributing to the epiphyte community structure of these forests, highlighting their role in species differentiation between sites. Consequently, the urban forests studied exhibited differences in species composition. This can be attributed to the variations in the composition and tree structure of the sites. Additionally, the HMF naturally shows species heterogeneity, as research has demonstrated high beta diversity among forest fragments for certain plant groups, including ferns (Hernández-Rojas 2010 ; Carvajal-Hernández et al. 2014 ). These results indicate that the urban forests examined are heterogeneous in their epiphytic and allied terrestrial composition. It is important to note that urbanization tends to reduce overall diversity, especially in epiphytes (McKinney 2008 ; Alvim et al. 2021 ). While previous studies have focused on smaller urban green spaces, our data was collected from urban forests covering more than 5 hectares of wooded area. This larger scale supports greater richness and diversity of epiphytic and related terrestrial species, underscoring the necessity of preserving large fragments of vegetation within cities. Due to their sensitivity to environmental changes caused by disturbances like deforestation and human activities, many epiphytes are considered bioindicators of habitat quality because they respond to variations in humidity and temperature. This sensitivity makes them vulnerable to human-induced disturbances that alter community structure and composition (Larrea and Werner 2010 ; Köster et al. 2011 ; Cach et al. 2014; Zotz 2016 ; Carvajal-Hernández et al. 2017 ; Aoki-Gonçalves et al. 2023 ). While some species that manage to survive the pressures of urbanization may exhibit effective strategies under stressful conditions, ferns of the genus Pleopeltis demonstrate poikilohydry, allowing them to reduce their exposed leaf area and minimize water loss in extreme conditions. This enables them to survive with only 25% of their water content until they receive a water pulse. (Moran 2004 ; Hietz 2010 ). Species of the genus Thelypteris thrive in warm climates and open areas. Examples include Macrothelypteris torresiana and Christella dentata , introduced species that have naturalized in tropical regions worldwide, demonstrating how these plants flourish in disturbed environments (Mehltreter 2008 ; Fawcett and Smith 2021 ). Other adaptations encompass reduced size and the presence of leaf scales, which help repel solar radiation and absorb nutrients and water from the environment. This adaptation has enabled species such as Tillandsia juncea, Tillandsia schiedeana, Pleopeltis furfuracea , and Pleopeltis thyssanolepis to survive in these altered environments (Page 2002 ; Watkins et al. 2006 ; Hietz 2010 ; Benzing 2012 ; Zotz 2016 ). Environmental modifications in the forests of Xalapa have resulted in a greater presence of generalist species adapted to the stressful conditions of the urban environment, which are characterized by high resilience to water fluctuations and tolerance to sunlight (e.g., Pleopeltis spp. and atmospheric Tillandsia species) (Hietz 2010 ; Benzing 2012 ; Zotz 2016 ). This pattern has been documented in other studies related to urbanization and habitat fragmentation, where environmental conditions become more favorable for generalist species, thereby displacing disturbance-sensitive species (McKinney, 2008 ; Krömer et al. 2014 ; Reyes-García et al. 2023 ). This indicates that, although the urban forests of Xalapa maintain considerable diversity, the current conditions restrict the presence of species with specific ecological requirements, which rely on less disturbed environments with greater stability in their microclimates. Conversely, in peri-urban forests like Clavijero and La Martinica, which face fewer current environmental pressures, species regarded as indicators for the conservation of mountain rainforests ( Vittaria graminifolia, Alsophila firma, Elaphoglossum sartorii, Elaphoglossum vestitum, Lophosoria quadripinnata, Trichomanes capillaceum, Trichomanes reptans ) were documented (Carvajal-Hernández et al. 2017 ). However, in this study, when applying the IndVal to link the species to a specific site, it was found that only 26% of the total exhibit an indication value, with ten classified as generalists, as they were identified in at least three of the examined forests. This indicates that they flourish in their living environment, having adjusted to overcome the existing environmental pressures. The physiological (CAM metabolism) and morphological adaptations (such as the presence of scales, elongated leaves, or tank-like structures for water storage) of generalist species enable them to thrive and colonize areas that may present environmental stressors that other species cannot tolerate (Gámez-Virués et al. 2015 ; Aronson et al. 2016 ; Alvim et al. 2021 ). Conclusion Urban forests can help mitigate conditions that negatively impact urban areas, such as the homogenization of diversity. The variation in vascular epiphyte richness and turnover found in peri-urban forests, compared to the lower variation in inner-city forests, indicates that changing environmental conditions in the latter favor generalist and stress-tolerant species. This dynamic underscore urbanization's role as a selective filter for biological diversity, limiting the presence of species with specific ecological needs. Conversely, areas on the city's periphery serve as vital reservoirs for the diversity of epiphytic species and their terrestrial relatives, particularly promoting the persistence of rare and conservation indicator species. This suggests that current environmental conditions support overall diversity and are likely to enhance the ecosystem services it provides. Declarations Acknowledgments: To the Secretaría de Medio Ambiente del Estado de Veracruz, for the facilities to enter the Natural Protected Areas. To Ana María Aquino Zapata, Rodrigo Carral Domínguez for their support in the field work. Funding This work was funded by the Consejo Nacional de Humanidades Ciencia y Tecnología (CONAHCYT) through Frontier Science project in the group modality (64358). The first author (MBJP) received funding for her postdoctoral stay from 2023 to 2025 (4816369) through CONAHCYT. Competing Interests The authors have no relevant financial or non-financial interests to disclose. Author contributions S.A.M. and C.I.C.H. contributed to the study conception and design. Fieldwork and data collection were conducted by M.B.J.P., S.A.M., C.I.C.H. and J.C.L.; S.A.M. prepared the map. M.B.J.P. and C.I.C.H. performed the analyses. The first draft of the manuscript was written by M.B.J.P. and C.I.C.H. All authors contributed to drafting and improving the manuscript. Each author read and approved the final version of the manuscript. References Acebey A, Gradstein S, Krömer T (2003) Species richness and hábitat diversification of bryophytes in submontane rain forest and fallows of Bolivia. J Trop Ecol 19(1):9–18. https://doi:10.1017/S026646740300302X Adhikari YP, Hoffmann S, Kunwar RM, Bobrowski M, Jentsch A, Beierkuhnlein C (2021) Vascular epiphyte diversity and host tree architecture in two forest management types in the Himalaya. 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Biol J Linn Soc 171(3):453-481. https://doi.org/10.1111/boj.12010 Zotz G (2016) Plants on Plants-The Biology of Vascular Epiphytes. Vol. 15. Springer International Publishing, Cham. Additional Declarations No competing interests reported. Supplementary Files AnnexI.docx Cite Share Download PDF Status: Published Journal Publication published 26 Jun, 2025 Read the published version in Urban Ecosystems → Version 1 posted Editorial decision: Revision requested 24 Apr, 2025 Reviews received at journal 23 Apr, 2025 Reviewers agreed at journal 05 Apr, 2025 Reviewers agreed at journal 31 Mar, 2025 Reviews received at journal 29 Mar, 2025 Reviewers agreed at journal 09 Mar, 2025 Reviewers invited by journal 07 Mar, 2025 Editor assigned by journal 02 Mar, 2025 Submission checks completed at journal 28 Feb, 2025 First submitted to journal 26 Feb, 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. As a division of Research Square Company, we’re committed to making research communication faster, fairer, and more useful. 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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-6116120","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":422857075,"identity":"f522303b-fe61-4e2b-aeca-4260c0cc55be","order_by":0,"name":"Maria Berenice Jarquin-Pacheco","email":"","orcid":"","institution":"Universidad Veracruzana","correspondingAuthor":false,"prefix":"","firstName":"Maria","middleName":"Berenice","lastName":"Jarquin-Pacheco","suffix":""},{"id":422857076,"identity":"123a0a51-7958-49d5-b88e-73e79de58606","order_by":1,"name":"Samaria Armenta-Montero","email":"","orcid":"","institution":"Universidad Veracruzana","correspondingAuthor":false,"prefix":"","firstName":"Samaria","middleName":"","lastName":"Armenta-Montero","suffix":""},{"id":422857077,"identity":"9a6f1359-eef2-4fb5-9c70-49ffb2107ec3","order_by":2,"name":"Jazmín Contreras-López","email":"","orcid":"","institution":"Universidad Veracruzana","correspondingAuthor":false,"prefix":"","firstName":"Jazmín","middleName":"","lastName":"Contreras-López","suffix":""},{"id":422857078,"identity":"21c43585-2ec7-4c68-b6c6-65b7825c40db","order_by":3,"name":"César Isidro Carvajal-Hernández","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAAtElEQVRIiWNgGAWjYPACNgY+9gZStbDxHCDZHokEIlXytx9+9pmnhk+eTfKNmQRjzmHCWiTOpBnP5jnGZtgmnQPUsi2NsBYDCQZjxhlsbIxALcYGjNtsiNHC/plxxj82+zbJMyAtEsRo4TFm+NjGltgmwWP4gChbJM7kFDN87GNLbuNJK3yQSIxf+NuPb2ZI+HbMtp/98IYDH7cREWJQcAxCJRCtgYGhhgS1o2AUjIJRMOIAAO/rLvzZzJFpAAAAAElFTkSuQmCC","orcid":"","institution":"Universidad Veracruzana","correspondingAuthor":true,"prefix":"","firstName":"César","middleName":"Isidro","lastName":"Carvajal-Hernández","suffix":""}],"badges":[],"createdAt":"2025-02-26 21:38:13","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-6116120/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-6116120/v1","draftVersion":[],"editorialEvents":[{"content":"https://doi.org/10.1007/s11252-025-01756-w","type":"published","date":"2025-06-26T15:56:56+00:00"}],"editorialNote":"","failedWorkflow":false,"files":[{"id":77628607,"identity":"6ceb2f6e-76b8-4a70-8ea9-eae55531c264","added_by":"auto","created_at":"2025-03-03 17:05:41","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":1018918,"visible":true,"origin":"","legend":"\u003cp\u003eGeographic locations of urban forests in the Xalapa-Banderilla conurbation, Veracruz, Mexico.\u003c/p\u003e","description":"","filename":"Fig1.png","url":"https://assets-eu.researchsquare.com/files/rs-6116120/v1/1262a23e55d8c61ca4f8a35e.png"},{"id":77629514,"identity":"b4d375f0-61cf-43c8-bb71-84f1ac416801","added_by":"auto","created_at":"2025-03-03 17:13:41","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":801711,"visible":true,"origin":"","legend":"\u003cp\u003ea)\u003cstrong\u003e \u003c/strong\u003eRarefaction curve and extrapolation of estimated richness. b) Diversity of epiphytes and their terrestrial equivalents in the urban forests of the Xalapa-Banderilla conurbation, Veracruz, Mexico, assessed using Hill numbers. CAD = Campus CAD; CLA = Clavijero Park; MAC = Macuiltépetl Park; MAR = La Martinica; TEJ = Tejar Garnica.\u003c/p\u003e","description":"","filename":"22.png","url":"https://assets-eu.researchsquare.com/files/rs-6116120/v1/e0ced490837d1a18e9012656.png"},{"id":77628608,"identity":"3842565c-607b-4d48-9042-30fd4ab803e8","added_by":"auto","created_at":"2025-03-03 17:05:41","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":79812,"visible":true,"origin":"","legend":"\u003cp\u003eEpiphyte species richness and their terrestrial counterparts in the urban (Tejar Garnica: TEJ; CAD Campus: CAD; Macuiltépetl Park: MAC) and peri-urban (Clavijero Park: CLA; La Martinica: MAR) forests of Xalapa-Banderilla conurbation, Veracruz, Mexico.\u003c/p\u003e","description":"","filename":"Fig3.png","url":"https://assets-eu.researchsquare.com/files/rs-6116120/v1/3b5e63235a7b482137ca21d1.png"},{"id":77628610,"identity":"9c9d0b4d-374c-4fc8-8a33-15c2932a2c31","added_by":"auto","created_at":"2025-03-03 17:05:41","extension":"png","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":124374,"visible":true,"origin":"","legend":"\u003cp\u003eDendrogram illustrating floristic similarity based on Jaccard's index of epiphytes and their terrestrial counterparts between urban (Tejar Garnica: TEJ; CAD Campus: CAD; Macuiltépetl Park: MAC) and peri-urban (Clavijero Park: CLA; La Martinica: MAR) forests in Xalapa-Banderilla conurbation, Veracruz, Mexico.\u003c/p\u003e","description":"","filename":"Fig4.png","url":"https://assets-eu.researchsquare.com/files/rs-6116120/v1/bdaf76a4408f9b91f871425c.png"},{"id":85686144,"identity":"79018ede-1a39-4939-9b7d-acc8c7c5f09e","added_by":"auto","created_at":"2025-06-30 16:03:50","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":3131942,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-6116120/v1/afd59c44-11b8-422d-8edd-651921250992.pdf"},{"id":77628609,"identity":"08494a57-5d95-40da-862b-4923637e7fea","added_by":"auto","created_at":"2025-03-03 17:05:41","extension":"docx","order_by":1,"title":"","display":"","copyAsset":false,"role":"supplement","size":54573,"visible":true,"origin":"","legend":"","description":"","filename":"AnnexI.docx","url":"https://assets-eu.researchsquare.com/files/rs-6116120/v1/47f85082027ae30ad147ce93.docx"}],"financialInterests":"No competing interests reported.","formattedTitle":"Urban forests as habitats for vascular epiphytes and allied terrestrial plants","fulltext":[{"header":"Introduction","content":"\u003cp\u003eEpiphytic plants are an important component of plant communities in the tropics, especially in humid mountain forests. These host a great diversity, accounting for up to 50% of the leaf biomass, and, in some forests, can represent up to 25% of the total vascular species (Cach-P\u0026eacute;rez et al. \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Gotsh et al. 2015). Vascular epiphytes are mostly represented by species of ferns, peperomias, bromeliads, aroids and orchids (Zotz \u003cspan citationid=\"CR89\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Kr\u0026ouml;mer et al. \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). These main groups can occasionally grow terrestrially in ecosystems, mainly ferns, and to a lesser extent, Araceae, orchids, and Peperomia. These plants provide habitat for organisms such as insects, birds, and amphibians, offering resources such as food and shelter; they also participate in canopy interactions (Zotz \u003cspan citationid=\"CR89\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Gonz\u0026aacute;lez and Ceballos \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Mendoza et al. \u003cspan citationid=\"CR58\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). However, these types of plants depend on atmospheric conditions like humidity and temperature; that is, they are sensitive to changes in their habitat, which can lead to shifts in the composition of their communities and a reduction in their richness. Therefore, they can serve as bioindicators of climate change and ecological damage (K\u0026ouml;ster et al. \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2009\u003c/span\u003e; Larrea and Werner \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; K\u0026ouml;ster et al. \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Reina-Rodr\u0026iacute;guez et al. \u003cspan citationid=\"CR69\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Carvajal-Hern\u0026aacute;ndez et al. \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2017\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eVascular epiphytes are one of the most important elements in humid mountain forests (HMF) (Gotsh et al. 2015). In Mexico, this type of vegetation supports a significant diversity of plants due to the transition zone between two biogeographic regions that converge in the country (Rzedowski \u003cspan citationid=\"CR72\" class=\"CitationRef\"\u003e1991\u003c/span\u003e; \u003cspan citationid=\"CR73\" class=\"CitationRef\"\u003e2006\u003c/span\u003e). One of the most vital ecosystem services provided by this type of vegetation is the availability of water resources, derived from precipitation and fog, which is essential for the establishment of groups dependent on humid conditions, such as vascular epiphytes (Kr\u0026ouml;mer et al. \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Mu\u0026ntilde;oz-Villers et al. \u003cspan citationid=\"CR61\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). Despite its ecological importance, this forest is one of those currently presents strong anthropogenic pressure, primarily due to changes in land use for creating agricultural land, livestock, and urban development (Challenger and Caballero \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e1998\u003c/span\u003e; Mu\u0026ntilde;oz-Villers et al. \u003cspan citationid=\"CR61\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Rosas et al. \u003cspan citationid=\"CR71\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). Recent studies have revealed that over a period of 17 years (1999\u0026ndash;2016), a 47.2% reduction in this type of vegetation has been observed, highlighting the current issues faced by the HMF (Espinoza-Guzm\u0026aacute;n et al. \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). The loss, reduction, and/or fragmentation of this forest type leads to the decline of species in general and vascular epiphytes in particular (Paredes et al. \u003cspan citationid=\"CR64\" class=\"CitationRef\"\u003e2021\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eOne of the areas in Mexico that still maintains remnants of the HMF within the city and in its surroundings is the conurbation of Xalapa, located in the central part of the State of Veracruz, along the slope of the Atlantic Ocean (Williams-Linera et al. \u003cspan citationid=\"CR84\" class=\"CitationRef\"\u003e2002\u003c/span\u003e). However, urban growth in Xalapa has occurred at the expense of rural areas and natural ecosystems (Von Thaden et al. 2021), modifying 90% of the original HMF (Williams-Linera et al. \u003cspan citationid=\"CR84\" class=\"CitationRef\"\u003e2002\u003c/span\u003e). These transformations have altered the structure and functioning of ecosystems, creating habitat fragmentation, causing a lack of connectivity between them (Von Thaden et al. 2021), and inducing the loss of plant species. However, the presence of forest fragments within urban areas can still sustain some biodiversity in small islands of vegetation (Rosas et al. \u003cspan citationid=\"CR71\" class=\"CitationRef\"\u003e2019\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eGiven the threat posed by biodiversity loss and its combination with overall urban growth, priority has been given to preserving areas with some type of vegetation, such as urban forests (UF), which are defined as areas with native and secondary vegetation, existing within transformed ecosystems in cities (Zewdie and Tegegne \u003cspan citationid=\"CR88\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). In that sense, the city of Xalapa is still recognized as a \u0026ldquo;green city\" because of the presence of areas with remnant forests of native vegetation and/or protected natural areas that exist in a fragmented manner and at various stages of ecological succession (Williams-Linera et al. \u003cspan citationid=\"CR84\" class=\"CitationRef\"\u003e2002\u003c/span\u003e; Von Thaden et al. 2021), which, on a smaller scale, also include parks and gardens medians. UF provides a variety of benefits to cities, including the conservation of environmental services (such as carbon sequestration, microclimate regulation, and mitigation of weather events), protection of flora and fauna, and space for recreation (Heidt and Neef \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; WHO 2017; Lemoine-Rodriguez et al. 2019).\u003c/p\u003e \u003cp\u003eUrban forests can support high levels of biodiversity, including endangered species (Alvey \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2006\u003c/span\u003e). This richness is highly dependent on the conservation status of the UF (Pesola et al. \u003cspan citationid=\"CR67\" class=\"CitationRef\"\u003e2017\u003c/span\u003e) and the existence of remnants of native vegetation (Kowarik \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e2011\u003c/span\u003e). For instance, forests adjacent to cities tend to support a greater diversity of medium-sized mammals than more urbanized areas (Tee et al. \u003cspan citationid=\"CR76\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). Similarly, a 10\u0026ndash;30% increase in native vegetation cover has been found to support a rise in the presence of bats, birds, and insects (Threlfall et al. \u003cspan citationid=\"CR78\" class=\"CitationRef\"\u003e2017\u003c/span\u003e) by providing shelter and food (Mohamad et al \u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). However, different taxa respond distinctly to urban spaces (MacGregor-Fors et al. \u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e2016\u003c/span\u003e), as evident in the case of epiphytes (Santana et al. \u003cspan citationid=\"CR74\" class=\"CitationRef\"\u003e2017\u003c/span\u003e), whose presence is associated with trees of larger diameter and height (Izuddin and Webb \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). Considering that the conurbation of Xalapa contains forest fragments surrounded by urban development on its periphery, there is an opportunity to understand the patterns of vascular epiphyte diversity under various anthropogenic pressures typical of urban dynamics, including isolation, modifications to tree structure, biodiversity extraction, microclimatic changes, soil degradation, and the introduction of exotic flora. Therefore, the aim is to examine the patterns of vascular epiphytes in urban forests with varying transformation histories in Xalapa, situated in central Veracruz, Mexico. This study contributes to recognizing the essential role of vegetation remnants in the city and their significance as a refuge for the diversity of vascular epiphytes and associated terrestrial plants.\u003c/p\u003e"},{"header":"Materials and Methods","content":"\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e \u003ch2\u003eStudy area\u003c/h2\u003e \u003cp\u003eThe study was carried out in the conurbation of the municipalities of Xalapa and Banderilla, located in the central region of the State of Veracruz in southeastern Mexico (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). The area contains remnants of humid mountain forest, consisting of patches of vegetation of various sizes. The surface area of the urban region of Xalapa encompasses approximately 60 km\u0026sup2;, where a mixture of native and exotic species has been recently observed, similar to what occurs in various cities (Falf\u0026aacute;n and MacGregor-Fors \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). The altitude ranges from 1,120 to 1,700 meters, with the predominant climates being semi-warm humid, characterized by abundant summer rainfall, and semi-warm humid with year-round rainfall. The temperature range oscillates between 18 and 24\u0026deg;C, while precipitation levels vary from 1,100 to 1,600 mm annually (SIEGVER 2020; Jara-Toto et al. \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2023\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eThe study was conducted in five urban forests, three of which are located within the city of Xalapa: 1) Protected Natural Area \u0026ldquo;El Tejar Garnica\u0026rdquo; (TEJ) (133 ha, altitude of 1310 m), corresponding to the polygon known as Parque Natura; 2) Protected Natural Area \u0026ldquo;Macuilt\u0026eacute;petl Ecological Park\u0026rdquo; (MAC) (31 ha; 1600 masl); 3) \u0026ldquo;Campus para la Cultura, las Artes y el Deporte\u0026rdquo; (CAD) primarily in its section known as Agrobosque Kan\u0026aacute; (28 ha; 1366 masl); 4) \u0026ldquo; Francisco Xavier Clavijero Ecological Reserve\u0026rdquo; (CLA) (59.85 ha; 1362 masl), hereafter referred to as Clavijero Park; and 5) \u0026ldquo;La Martinica\u0026rdquo; Natural Protected Area (MAR) (52.30 ha; 1599 masl). The last two sites are situated on the periphery of the urban areas of the municipalities of Xalapa and Banderilla. Of all the study sites, only the CAD Campus is not officially classified by the government as a protected natural area; however, it is under the protection of the Universidad Veracruzana as an environmental management unit. The remaining sites are protected natural areas administered by different levels of government; the vegetation they harbor are remnants of humid mountain forest with different degrees of anthropogenic disturbance and different transformation histories (Jara-Toto et al. \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2023\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eIn the forest fragments within the city (El Tejar Garnica, Campus CAD, and Parque Macuilt\u0026eacute;petl), the vegetation structure is mainly composed of tree species from secondary vegetation derived from the HMF, such as \u003cem\u003eDendropanax arboreus\u003c/em\u003e, \u003cem\u003ePiper amalago\u003c/em\u003e, and \u003cem\u003eHeliocarpus appendiculatus\u003c/em\u003e (Jara-Toto et al. et al. 2023), where the presence of native and introduced plant species is also recorded. In the peri-urban area of Clavijero Park, typical HMF species exhibit a higher state of maturity, including \u003cem\u003eClethra macrophylla, Liquidambar styraciflua, Quercus lancifolia\u003c/em\u003e, and \u003cem\u003eTurpinia insignis\u003c/em\u003e (Williams-Linera et al. \u003cspan citationid=\"CR85\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). Meanwhile, in La Martinica, there is a mix of primary and secondary HMF species at various levels of conservation.\u003c/p\u003e \u003c/div\u003e\n\u003ch3\u003eField work\u003c/h3\u003e\n\u003cp\u003eSampling was conducted by establishing five plots measuring 20 x 20 m (400 m\u0026sup2;) at each of the five sites mentioned earlier, following the methodology proposed by Kessler and Bach (\u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e1999\u003c/span\u003e). As a result, a total of 25 plots (1 ha) were sampled across the five areas. Each plot was selected to maintain a distance from water bodies and prevent the establishment of a well-defined tree and shrub layer. In each plot, data on the incidence of understory species were recorded, including both holoepiphytes and related groups found on the ground or on fallen trunks and branches (e.g., ferns, Araceae, peperomias, and terrestrial orchids). This approach was adopted to ensure adequate representation of specific groups seen as indicators of mountain ecosystem quality, such as certain ferns found in terrestrial habitats. However, some species are almost exclusively epiphytes, like the genus \u003cem\u003eTillandsia\u003c/em\u003e. This strategy aimed to increase the number of species recorded, providing a more comprehensive view of the plant community within the studied ecosystem. Furthermore, certain plants, such as specific ferns, are recognized as indicators of habitat quality (Della and Falkenberg, \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). Including both epiphytes and terrestrial species allowed for better identification of the groups that reflect the condition and status of the ecosystems.\u003c/p\u003e \u003cp\u003eThe observed species were collected and herborized according to the methodology of Lot and Chiang (\u003cspan citationid=\"CR53\" class=\"CitationRef\"\u003e1986\u003c/span\u003e) and Lorea and Riba (\u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e1990\u003c/span\u003e). Subsequently, dichotomous keys were used for their identification, and the specimens were deposited in the herbarium of the Universidad Veracruzana in Xalapa (XALU), with duplicates stored in the National Herbarium (MEXU) of the Institute of Biology at the Universidad Aut\u0026oacute;noma de M\u0026eacute;xico. The resulting floristic list was organized at the family level following APG IV (\u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2016\u003c/span\u003e) for angiosperms and PPGI (\u003cspan citationid=\"CR68\" class=\"CitationRef\"\u003e2016\u003c/span\u003e) for ferns. The scientific names of each species were verified using the Tropics.org database (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://www.tropicos.org/home\u003c/span\u003e\u003cspan address=\"https://www.tropicos.org/home\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e). The IUCN Red List was consulted to compile species in any risk category. At the national level, the Official Mexican Standard (NOM-059-SEMARNAT-2010) was reviewed to identify species that fall into a risk category within Mexican territory.\u003c/p\u003e \u003cdiv id=\"Sec5\" class=\"Section2\"\u003e \u003ch2\u003eData analysis\u003c/h2\u003e \u003cp\u003eTo assess alpha diversity, the data were analyzed using the online version of iNEXT software to generate rarefaction curves that evaluate sampling effort and calculate Hill numbers to measure diversity at each study site (Chao et al. \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). The Kruskal-Wallis test was conducted to confirm differences in total species richness across all sites, followed by Dunn's post hoc test to assess the differences between each site. For beta diversity, Whittaker's species turnover index was calculated. These analyses were conducted using Past 4.04 (Hammer et al. \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2001\u003c/span\u003e; Koleff et al. \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e2003\u003c/span\u003e; Tuomisto \u003cspan citationid=\"CR80\" class=\"CitationRef\"\u003e2010\u003c/span\u003e). A similarity dendrogram was also created to group the various urban forests based on their floristic composition, utilizing the Bray-Curtis distance matrix through the Vegan package in RStudio. Additionally, a one-way similarity analysis (ANOSIM) was conducted using the Bray-Curtis similarity index to confirm statistical differences in species composition among the sampled communities. The similarity percentage (SIMPER) was calculated to identify the contribution of different species to the composition of the communities studied. To evaluate whether certain species are indicators of a site, IndVal analysis was performed using the \u0026ldquo;indicspecies\u0026rdquo; package in R (Dufr\u0026ecirc;ne and Legendre \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e1997\u003c/span\u003e). This analysis focuses on specificity and fidelity, where values near 1 signify that species effectively indicate the habitat, while very low (near 0) values denote rare species (Mart\u0026iacute;n-Regalado \u003cspan citationid=\"CR55\" class=\"CitationRef\"\u003e2019\u003c/span\u003e).\u003c/p\u003e \u003c/div\u003e"},{"header":"RESULTS","content":"\u003cdiv id=\"Sec7\" class=\"Section2\"\u003e \u003ch2\u003eSpecies richness\u003c/h2\u003e \u003cp\u003eIn a total sampling of 1 hectare, 103 species were recorded across 58 genera and 22 families (Annex 1). Of these, 55 are epiphytic species (35 are angiosperms, while the rest are ferns), and 48 species represent related groups found on land. Among the ten best-represented families in the entire study, six are ferns. The families with the highest richness are as follows: Polypodiaceae (16), Orchidaceae (15), and Bromeliaceae (11); with the genus \u003cem\u003eTillandsia\u003c/em\u003e being the most represented with nine species, followed by \u003cem\u003eThelypteris\u003c/em\u003e with seven, and the genera \u003cem\u003ePeperomia\u003c/em\u003e and \u003cem\u003ePleopeltis\u003c/em\u003e, each with six species. Therefore, the most abundant group in terms of the number of records consists of ferns (58%), including both epiphytic and terrestrial species, followed by orchids (15%) and bromeliads (11%). Of the total number of species, four were registered in the NOM-059-SEMARNAT-2010, three were in the IUCN Red List, and 14 were endemic to Mexico.\u003c/p\u003e \u003cp\u003eThe richness estimated from the sampling coverage obtained with the iNEXT software indicates that the sampling effort in the five urban forests studied is adequate (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003ea). Additionally, Hill's numbers show that Clavijero (CLA) had the highest richness (q0) and diversity (q1, q2), while two areas within the city (TEJ and CAD) exhibited the lowest values (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003eb).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eThe comparison of richness among the study sites indicates that Clavijero Park is the peri-urban forest with the highest richness (64 species), followed by La Martinica with 49 species, while the CAD Campus had the lowest richness (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). These data were supported by the results of the Kruskal-Wallis test, which demonstrated significant differences among the sampled urban forests (H\u0026thinsp;=\u0026thinsp;17.65, Hc\u0026thinsp;=\u0026thinsp;24.03, p\u0026thinsp;\u0026lt;\u0026thinsp;0.05), indicating that at least one site significantly differs in richness. A post-hoc Dunn's test with Bonferroni correction was then performed to identify differences between the sites. The results indicated that Clavijero Park has significantly higher richness than other sites (p\u0026thinsp;\u0026lt;\u0026thinsp;0.05). However, no significant differences in richness were found among the other sampled forests (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e, Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\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\u003eSpecies richness by site: Urban areas include Tejar Garnica, CAD Campus, and Macuilt\u0026eacute;petl Park; peri-urban areas encompass Clavijero Park in the Municipality of Xalapa and La Martinica in the Municipality of Banderilla, Veracruz, Mexico.\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"6\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" 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 \u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eSite\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eSpecies richness (Ephytes/Terrestrials)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eFamily richness\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colspan=\"2\" nameend=\"c5\" namest=\"c4\"\u003e \u003cp\u003eMost representative families\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003eLocation\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eTejar Garnica\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e35 (20/15)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e11\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003ePolypodiaceae (7),\u003c/p\u003e \u003cp\u003eOrchidaceae (7), Bromeliaceae (4) y Araceae (4)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c6\" namest=\"c5\"\u003e \u003cp\u003eUrban\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eMacuilt\u0026eacute;petl Park\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e41 (23/18)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e17\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003ePolypodiaceae (11),\u003c/p\u003e \u003cp\u003eOrchidaceae (4) y\u003c/p\u003e \u003cp\u003eBromeliaceae (4)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c6\" namest=\"c5\"\u003e \u003cp\u003eUrban\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eCampus CAD\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e34 (16/18)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e14\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003ePolypodiaceae (5), Thelypteridaceae(5) y Bromeliaceae (5)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c6\" namest=\"c5\"\u003e \u003cp\u003eUrban\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eLa Martinica\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e49 (30/19)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e17\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003ePolypodiaceae (10),\u003c/p\u003e \u003cp\u003eBromeliaceae (6),\u003c/p\u003e \u003cp\u003eOrchidaceae (5) y\u003c/p\u003e \u003cp\u003ePiperaceae (5)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c6\" namest=\"c5\"\u003e \u003cp\u003ePeri-urban\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eClavijero Park\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e64 (38/26)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e20\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003ePolypodiaceae (12),\u003c/p\u003e \u003cp\u003eOrchidaceae (10) y\u003c/p\u003e \u003cp\u003eBromeliaceae (9)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c6\" namest=\"c5\"\u003e \u003cp\u003ePeri-urban\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 \u003c/div\u003e \u003cdiv id=\"Sec8\" class=\"Section2\"\u003e \u003ch2\u003eSpecies turnover\u003c/h2\u003e \u003cp\u003eThe Whittaker index results indicate that the peri-urban area of La Martinica has the highest species turnover compared to urban areas. Additionally, the peri-urban areas show index values exceeding 50% among themselves, and this notable species turnover is also observed between the peri-urban and urban areas. Urban areas exhibit smaller differences in species composition when compared to one another (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). Regarding the floristic similarity among the analyzed urban forests, the similarity dendrogram shows the division into three groups. One group includes the inner-city sites Campus CAD and El Tejar Garnica, while the other group consists of the two peri-urban areas. The species from Macuilt\u0026eacute;petl Park, which is situated in the center of the urban area, are partly grouped with the peri-urban forests, creating a subgroup with Clavijero Park, while another part forms a separate group (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e). On the other hand, the ANOSIM performed with the Bray-Curtis similarity index indicates differences in the floristic composition of epiphytes and their terrestrial counterparts among the sampled urban forests (R\u0026thinsp;=\u0026thinsp;0.8398; P\u0026thinsp;\u0026lt;\u0026thinsp;0.05). Paired ANOSIM comparisons show differences in composition at all sites, with values of P\u0026thinsp;\u0026lt;\u0026thinsp;0.05.\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab2\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 2\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eWhittaker's beta diversity index for the sampled sites of the Xalapa-Banderilla conurbation, Veracruz, Mexico.\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"6\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" 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 \u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\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\u003eTejar Garnica\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eMacuilt\u0026eacute;petl Park\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eCampus CAD\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003eLa Martinica\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003eClavijero Park\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eTejar Garnica\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.474\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.450\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.643\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.535\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eMacuilt\u0026eacute;petl Park\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.545\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.622\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.429\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eCampus CAD\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.518\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.490\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eLa Martinica\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.522\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eClavijero Park\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0\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 \u003csup\u003eValues of 0 represent equality and 1 signifies complete difference\u003c/sup\u003e.\u003c/p\u003e \u003cp\u003eAccording to the SIMPER analysis, the first 20 species that contribute the most to dissimilarity (representing 19% of the total registered species) account for 39% of the variation in species composition recorded across the five sites. Meanwhile, 40 species represent 68% of the contribution (Table\u0026nbsp;\u003cspan refid=\"Tab3\" class=\"InternalRef\"\u003e3\u003c/span\u003e). Ferns are the most numerous group in the study overall, but they do not significantly contribute to the changes in composition. In contrast, the Bromeliaceae family, mainly species from the genus \u003cem\u003eTillandsia\u003c/em\u003e, does have a substantial impact; among the 20 species with the greatest contribution, they account for 16% of the richness.\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab3\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 3\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eSpecies that, according to the SIMPER analysis, contribute more to the dissimilarity of epiphytic plant communities and their terrestrial allied in the urban forests of Xalapa-Banderilla conurbation, Veracruz, Mexico.\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"4\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" 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 \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eSpecies\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eAverage dissimilarity\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eContribution (%)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eCumulative (%)\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eTillandsia schiedeana\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e1.467\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e2.258\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e2.258\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eTillandsia deppeana\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e1.457\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e2.244\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e4.502\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003ePteris quadriaurita\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e1.405\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e2.163\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e6.665\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eSyngonium\u003c/em\u003e sp.\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e1.396\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e2.149\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e8.815\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eCatopsis nutans\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e1.374\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e2.116\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e10.93\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003ePolypodium conterminans\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e1.373\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e2.115\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e13.05\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eTillandsia butzii\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e1.352\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e2.082\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e15.13\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eTillandsia multicaulis\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e1.328\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e2.046\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e17.17\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eTillandsia juncea\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e1.31\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e2.017\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e19.19\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003ePteris orizabae\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e1.294\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e1.993\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e21.18\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eCyclopogon luteo-albus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e1.29\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e1.987\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e23.17\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003ePeperomia glabella\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e1.283\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e1.976\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e25.15\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003ePleopeltis polypodioides\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e1.266\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e1.949\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e27.1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eAlsophila firma\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e1.209\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e1.861\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e28.96\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eSelaginella stellata\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e1.207\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e1.859\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e30.82\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eMonstera deliciosa\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e1.196\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e1.842\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e32.66\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eCampylocentrum schiedei\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e1.194\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e1.839\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e34.5\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eTillandsia polystachia\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e1.147\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e1.766\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e36.26\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eSelaginella martensii\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e1.141\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e1.756\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e38.02\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eScaphyglottis minutiflora\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e1.119\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e1.724\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e39.74\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\u003eAmong the total number of species documented in the five urban forests, only 27 (26%) demonstrated an indication value based on the IndVal (Indicator Value Method) analysis. Of these, 10 species were associated with more than three urban and peri-urban forests (37%). Macuilt\u0026eacute;petl Park and La Martinica had the highest number of indicator species, totaling four, while only one species was recorded as unique to CLA (Table\u0026nbsp;\u003cspan refid=\"Tab4\" class=\"InternalRef\"\u003e4\u003c/span\u003e).\u003c/p\u003e\u003cdiv class=\"gridtable\"\u003e\n \u003cdiv align=\"left\" class=\"colspec\"\u003e\u003c/div\u003e\n \u003cdiv align=\"left\" class=\"colspec\"\u003e\u003cstrong\u003eTable 4\u003c/strong\u003e Indicator species based on the IndVal analysis of epiphyte species and their allied terrestrial between urban (Tejar Garnica: TEJ; CAD Campus: CAD; Macuilt\u0026eacute;petl Park: MAC) and peri-urban (Clavijero Park: CLA; La Martinica: MAR) forests in Xalapa-Banderilla conurbation, Veracruz, Mexico. In all instances, the species exhibited values of P\u0026lt;0.05.\u003c/div\u003e\n\u003c/div\u003e\n\u003cdiv\u003e\n \u003ctable border=\"0\" cellspacing=\"0\" cellpadding=\"0\" width=\"88%\"\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 47.4747%;\"\u003e\n \u003cp\u003e\u003cstrong\u003eSpecies\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 37.3737%;\"\u003e\n \u003cp\u003e\u003cstrong\u003eSite\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 15.1515%;\"\u003e\n \u003cp\u003e\u003cstrong\u003eIndVal*\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 47.4747%;\"\u003e\n \u003cp\u003e\u003cem\u003eSyngonium podophyllum \u0026nbsp; \u0026nbsp;\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 37.3737%;\"\u003e\n \u003cp\u003eTEJ, MAC, CAD\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 15.1515%;\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 47.4747%;\"\u003e\n \u003cp\u003e\u003cem\u003eTillandsia punctulata\u0026nbsp;\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 37.3737%;\"\u003e\n \u003cp\u003eMAR\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 15.1515%;\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 47.4747%;\"\u003e\n \u003cp\u003e\u003cem\u003eScaphyglottis minutiflora \u0026nbsp;\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 37.3737%;\"\u003e\n \u003cp\u003eMAC\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 15.1515%;\"\u003e\n \u003cp\u003e0.966\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 47.4747%;\"\u003e\n \u003cp\u003e\u003cem\u003eChristella dentata\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 37.3737%;\"\u003e\n \u003cp\u003eTEJ, MAC, CAD, CLA\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 15.1515%;\"\u003e\n \u003cp\u003e0.934\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 47.4747%;\"\u003e\n \u003cp\u003e\u003cem\u003eTillandsia schiedeana\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 37.3737%;\"\u003e\n \u003cp\u003eTEJ, MAC, CLA\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 15.1515%;\"\u003e\n \u003cp\u003e0.931\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 47.4747%;\"\u003e\n \u003cp\u003e\u003cem\u003eTillandsia deppeana\u0026nbsp;\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 37.3737%;\"\u003e\n \u003cp\u003eCAD, MAR, CLA\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 15.1515%;\"\u003e\n \u003cp\u003e0.931\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 47.4747%;\"\u003e\n \u003cp\u003e\u003cem\u003eTillandsia butzii \u0026nbsp; \u0026nbsp;\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 37.3737%;\"\u003e\n \u003cp\u003eMAR, CLA\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 15.1515%;\"\u003e\n \u003cp\u003e0.929\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 47.4747%;\"\u003e\n \u003cp\u003e\u003cem\u003eSelaginella stellata\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 37.3737%;\"\u003e\n \u003cp\u003eCAD\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 15.1515%;\"\u003e\n \u003cp\u003e0.918\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 47.4747%;\"\u003e\n \u003cp\u003e\u003cem\u003eCampylocentrum schiedei \u0026nbsp;\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 37.3737%;\"\u003e\n \u003cp\u003eCAD, CLA\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 15.1515%;\"\u003e\n \u003cp\u003e0.894\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 47.4747%;\"\u003e\n \u003cp\u003e\u003cem\u003ePolypodium conterminans\u0026nbsp;\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 37.3737%;\"\u003e\n \u003cp\u003eMAC, MAR, CLA\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 15.1515%;\"\u003e\n \u003cp\u003e0.894\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 47.4747%;\"\u003e\n \u003cp\u003e\u003cem\u003eSelaginella martensii\u0026nbsp;\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 37.3737%;\"\u003e\n \u003cp\u003eMAC, CLA\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 15.1515%;\"\u003e\n \u003cp\u003e0.894\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 47.4747%;\"\u003e\n \u003cp\u003e\u003cem\u003eCatopsis nutans\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 37.3737%;\"\u003e\n \u003cp\u003eTEJ, MAR, CLA\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 15.1515%;\"\u003e\n \u003cp\u003e0.879\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 47.4747%;\"\u003e\n \u003cp\u003e\u003cem\u003ePteris quadriaurita\u0026nbsp;\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 37.3737%;\"\u003e\n \u003cp\u003eTEJ, MAC, CLA\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 15.1515%;\"\u003e\n \u003cp\u003e0.879\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 47.4747%;\"\u003e\n \u003cp\u003e\u003cem\u003eTillandsia juncea \u0026nbsp;\u0026nbsp;\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 37.3737%;\"\u003e\n \u003cp\u003eCAD, CLA\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 15.1515%;\"\u003e\n \u003cp\u003e0.869\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 47.4747%;\"\u003e\n \u003cp\u003e\u003cem\u003eTillandsia multicaulis \u0026nbsp;\u0026nbsp;\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 37.3737%;\"\u003e\n \u003cp\u003eMAC, MAR, CLA\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 15.1515%;\"\u003e\n \u003cp\u003e0.856\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 47.4747%;\"\u003e\n \u003cp\u003e\u003cem\u003ePeperomia san-joseana \u0026nbsp;\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 37.3737%;\"\u003e\n \u003cp\u003eMAC, MAR\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 15.1515%;\"\u003e\n \u003cp\u003e0.837\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 47.4747%;\"\u003e\n \u003cp\u003e\u003cem\u003eMonstera deliciosa\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 37.3737%;\"\u003e\n \u003cp\u003eMAC\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 15.1515%;\"\u003e\n \u003cp\u003e0.775\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 47.4747%;\"\u003e\n \u003cp\u003e\u003cem\u003eAdiantopsis radiata \u0026nbsp;\u0026nbsp;\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 37.3737%;\"\u003e\n \u003cp\u003eTEJ, CAD\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 15.1515%;\"\u003e\n \u003cp\u003e0.775\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 47.4747%;\"\u003e\n \u003cp\u003e\u003cem\u003eAlsophila firma \u0026nbsp; \u0026nbsp;\u0026nbsp;\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 37.3737%;\"\u003e\n \u003cp\u003eCAD, MAR, CLA\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 15.1515%;\"\u003e\n \u003cp\u003e0.775\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 47.4747%;\"\u003e\n \u003cp\u003e\u003cem\u003eCatopsis sessiliflora \u0026nbsp;\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 37.3737%;\"\u003e\n \u003cp\u003eMAC, CAD\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 15.1515%;\"\u003e\n \u003cp\u003e0.775\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 47.4747%;\"\u003e\n \u003cp\u003e\u003cem\u003eThelypteris pilosohispida\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 37.3737%;\"\u003e\n \u003cp\u003eMAR\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 15.1515%;\"\u003e\n \u003cp\u003e0.775\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 47.4747%;\"\u003e\n \u003cp\u003e\u003cem\u003ePolypodium eatonii \u0026nbsp; \u0026nbsp; \u0026nbsp;\u0026nbsp;\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 37.3737%;\"\u003e\n \u003cp\u003eMAR\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 15.1515%;\"\u003e\n \u003cp\u003e0.775\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 47.4747%;\"\u003e\n \u003cp\u003e\u003cem\u003eBlechnum schiedeanum \u0026nbsp;\u0026nbsp;\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 37.3737%;\"\u003e\n \u003cp\u003eMAR\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 15.1515%;\"\u003e\n \u003cp\u003e0.775\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 47.4747%;\"\u003e\n \u003cp\u003e\u003cem\u003ePhanerophlebia remotispora\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 37.3737%;\"\u003e\n \u003cp\u003eMAC\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 15.1515%;\"\u003e\n \u003cp\u003e0.775\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 47.4747%;\"\u003e\n \u003cp\u003e\u003cem\u003eCtenitis melanosticta \u0026nbsp;\u0026nbsp;\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 37.3737%;\"\u003e\n \u003cp\u003eMAC\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 15.1515%;\"\u003e\n \u003cp\u003e0.775\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 47.4747%;\"\u003e\n \u003cp\u003e\u003cem\u003eTillandsia polystachia \u0026nbsp;\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 37.3737%;\"\u003e\n \u003cp\u003eTEJ, MAC, CLA\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 15.1515%;\"\u003e\n \u003cp\u003e0.730\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 47.4747%;\"\u003e\n \u003cp\u003e\u003cem\u003ePolypodium rhodopleuron\u0026nbsp;\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 37.3737%;\"\u003e\n \u003cp\u003eCLA\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 15.1515%;\"\u003e\n \u003cp\u003e0.725\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n \u003c/table\u003e\n\u003c/div\u003e\n\u003cp\u003e\u003csup\u003e* IndVal values indicate affinity to a specific forest type, ranging from 1 (indicating it is only recorded in that forest type) to 0 (indicating it is not recorded at all)\u003c/sup\u003e.\u003c/p\u003e"},{"header":"Discussion","content":"\u003cp\u003eOut of the total species richness of epiphytic angiosperms reported for the state of Veracruz (569 species), only 6% was documented in this study (Kr\u0026ouml;mer et al. \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Given that a high diversity of epiphytes (285 species) exists in the humid montane forests of the state, the species documented in the urban forests of Xalapa represent 12.2% of the total diversity found in the humid montane ecosystems of Veracruz (Kr\u0026ouml;mer et al. \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Among the most well-represented groups, ferns, orchids, and bromeliads are notable as they contribute the highest number of epiphytic species in floristic inventories across the Neotropics (Zotz, \u003cspan citationid=\"CR89\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Kr\u0026ouml;mer et al. \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Furtado and Menini-Neto 2015; Villase\u0026ntilde;or 2016; Santana et al. \u003cspan citationid=\"CR74\" class=\"CitationRef\"\u003e2017\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eOne aspect to highlight in this study is the low presence of species from groups such as orchids and bromeliads, which are typically the most diverse group in inventories of epiphytic plants (Benzing \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e1990\u003c/span\u003e; Kreft et al. \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e2004\u003c/span\u003e; Flores-Palacios and Garc\u0026iacute;a-Franco 2006; Zotz \u003cspan citationid=\"CR89\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Alzate-Q et al. 2019). In contrast, ferns were the group with the highest representation in our results, excluding terrestrial species. This pattern aligns with findings by Kr\u0026ouml;mer et al. (\u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e2021\u003c/span\u003e) in central Veracruz, who observed that ferns dominate in quantity and cover in conserved forest areas, followed by orchids and bromeliads.\u003c/p\u003e \u003cp\u003eIn the urban and peri-urban forests studied, the presence of fourteen epiphytes endemic to Mexico was noted, constituting 0.12% of the endemic diversity of vascular plants identified in the country (Villase\u0026ntilde;or 2016). Of these, four are protected under Mexican laws by being included in the Norma Oficial Mexicana, which oversees the protection of wild plant and animal species (NOM-059-SEMARNAT-2010), and three are listed in the IUCN Red List under some risk category. This finding shows that, despite human pressure from urban growth, these areas continue to serve as important refuges for native and endemic biodiversity.\u003c/p\u003e \u003cp\u003eEpiphytes and related terrestrial groups (especially ferns) require high humidity conditions to establish themselves, which they find in forests with a higher degree of conservation; in this case, these conditions are maintained in peri-urban areas forests. A recent study indicates that sites like Clavijero Park exhibit more temperate microclimatic conditions, showing less daily variation in temperature and relative humidity throughout the year compared to other evaluated forests, particularly those located in the city's interior, which tend to be warmer and show greater fluctuations in these environmental variables (Landeros-L\u0026oacute;pez \u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). Therefore, the observed patterns of richness could be explained by these changes in the microclimate.\u003c/p\u003e \u003cp\u003eMoreover, vegetation structure may also be an important factor. A previous study revealed that the forests within the interior of the city (TEJ and CAD) and one peri-urban forest (MAR) exhibited a higher dominance of secondary trees with smaller diameters. In contrast, the Clavijero peri-urban area predominantly featured species typical of mature forests in the region, characterized by larger diameters (Jara-Toto et al. \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). Similarly, Macuilt\u0026eacute;petl Park, situated in the center of the urban area, features trees with larger diameters and primary species (Figueroa-Solis et al. 2023). This contrasting situation is explained by the unique transformation histories of each site. In the most transformed areas, there were previously pastures or abandoned crops, whereas Clavijero Park contained remnants of larger vegetation with less anthropogenic impact. However, it is important to note that all sites exhibit some degree of anthropogenic influence due to their proximity to the city.\u003c/p\u003e \u003cp\u003eThe above situation is significant for interpreting the results of epiphytes and their terrestrial relatives because tree species and the structure they create in the forest play a crucial role in the establishment and persistence of species. Host characteristics, such as diameter, bark type, and branching, are essential for epiphyte establishment and influence the microclimatic conditions they create in the understory (Bonnet et al. \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2007\u003c/span\u003e; Caglioni et al. \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2012\u003c/span\u003e; Gonz\u0026aacute;lez and Ceballos \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Additionally, tree age is a crucial factor in epiphyte diversity. Adhikari et al. (\u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2021\u003c/span\u003e) suggest that tree age may serve as a significant indicator of epiphyte diversity, as older and larger diameter trees provide better conditions for the establishment of these species. A study conducted in the urban area of Xalapa revealed that the proportion of woody species is linked to the distribution of epiphytes, particularly bromeliads (Aoki-Gon\u0026ccedil;alves et al. \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). This helps elucidate the pattern observed in the studied forests.\u003c/p\u003e \u003cp\u003eAt a site like Clavijero, having a better tree structure, along with its peripheral location, enhances connectivity and ecosystem continuity with other forest fragments, allowing for the movement and establishment of propagules. Furthermore, as demonstrated in other studies, structural diversity, along with improved microclimatic conditions and reduced anthropogenic pressure, promotes the establishment of these botanical groups, which could lead to greater species richness (Kr\u0026ouml;mer et al. \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Kr\u0026ouml;mer et al. \u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Von Thaden et al. 2021; Parra-S\u0026aacute;nchez and Banks-Leite 2022). In this peri-urban forest, which resembles a primary forest, epiphytes and their terrestrial relatives are finding optimal conditions for establishment, making this site an important source for species and germplasm conservation in the city's surrounding area Xalapa.\u003c/p\u003e \u003cp\u003eIn the case of La Martinica, which is a peri-urban area that currently experiences less anthropogenic pressure than the forests in the interior of the city, a greater richness was expected; however, the number of species is low, a situation that was also previously reported by Kr\u0026ouml;mer and collaborators (2021). This may be due to the smaller diameter trees present in this forest, indicating that it is undergoing succession. This factor may influence the establishment and abundance of these plants (Lahoti et al. \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Jara-Toto et al. \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2023\u003c/span\u003e), which supports the hypothesis of Woods and De Walt (2012), who argues that an increase in epiphyte species will be observed as forest age increases.\u003c/p\u003e \u003cp\u003eAnother factor that can be determinant in species richness in urban areas is the extraction of certain plants for ornamental purposes. In Xalapa, it's common to find orchids and bromeliads in local markets due to collector demand, which intensifies the pressure on these epiphytes (Flores-Palacios and Valencia-D\u0026iacute;az \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2007\u003c/span\u003e; Toledo-Aceves et al. \u003cspan citationid=\"CR79\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Kr\u0026ouml;mer et al. \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Although the study sites are protected natural areas by official decree and extraction activities are prohibited, the entry of people for various extraction activities (such as soil, firewood, and plants) has been observed in some urban forests (personal observation). This phenomenon may partially explain the disappearance of certain groups, with ferns dominating their place, as some species are infrequently used for these purposes in the area due to current human activities and past extraction practices.\u003c/p\u003e \u003cp\u003eOn the other hand, urban forests (Macuilt\u0026eacute;petl Park, Tejar Garnica, and Campus CAD) exhibit lower epiphyte richness. This could be attributed to their location within the urban infrastructure of Xalapa, as they are subject to territorial and functional simplification of their habitats (Acebey et al. \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2003\u003c/span\u003e; Mickinney 2008; de Juana Aranzana \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Parra-S\u0026aacute;nchez and Banks-Leite 2022). However, it is essential to understand the life history of these forests\u0026mdash;the historical processes of degradation and their environmental impact\u0026mdash;to properly grasp the low rates of observed species richness (Alvey \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2006\u003c/span\u003e). For example, Macuiltepetl Park, despite being surrounded by urban sprawl, ranks third in epiphyte richness with 41 species, surpassing the CAD Campus and Tejar Garnica. This may be due to restoration activities, such as those carried out in 1977, which contributed to the arboreal enrichment of the forest. Likewise, conserving a slope in the northeastern area that maintains the original arboreal vegetation of the HMF could have been an important source of propagules for plant dispersal and ecosystem regeneration (Pav\u0026oacute;n \u003cspan citationid=\"CR66\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). These past actions may have preserved microclimatic conditions and facilitated the subsequent establishment of epiphytes prior to the rapid growth of the city\u0026rsquo;s outskirts.\u003c/p\u003e \u003cp\u003eRegarding Tejar Garnica and the CAD Campus, which had the lowest epiphyte richness (35 and 34 species), both areas feature secondary vegetation derived from the HMF. Tejar Garnica was originally a coffee plantation and cattle pasture (Jara-Toto et al. \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2023\u003c/span\u003e), while the CAD Campus was also a cattle ranching pasture, where less than half of the area was left untouched to follow its natural secondary succession process, while the remaining area was altered with the establishment of gardens mainly composed of exotic flora (Teodosio-Faustino et al. \u003cspan citationid=\"CR77\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). The tree structure of the succession and intervention processes likely affects species richness, as it lacks the structural complexity of the forest, which reduces the formation of microclimates vital for the establishment of epiphytes and their allied terrestrials.\u003c/p\u003e\n\u003ch3\u003eRecambio de especies\u003c/h3\u003e\n\u003cp\u003eThe Whittaker index and cluster analysis results indicate species turnover between peri-urban and urban forests, particularly regarding TEJ and CAD, which show smaller differences between them. In line with this, the analysis of percentage similarity (SIMPER) enabled us to identify ferns and bromeliads as the primary groups contributing to the epiphyte community structure of these forests, highlighting their role in species differentiation between sites. Consequently, the urban forests studied exhibited differences in species composition. This can be attributed to the variations in the composition and tree structure of the sites. Additionally, the HMF naturally shows species heterogeneity, as research has demonstrated high beta diversity among forest fragments for certain plant groups, including ferns (Hern\u0026aacute;ndez-Rojas \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Carvajal-Hern\u0026aacute;ndez et al. \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). These results indicate that the urban forests examined are heterogeneous in their epiphytic and allied terrestrial composition. It is important to note that urbanization tends to reduce overall diversity, especially in epiphytes (McKinney \u003cspan citationid=\"CR56\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Alvim et al. \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). While previous studies have focused on smaller urban green spaces, our data was collected from urban forests covering more than 5 hectares of wooded area. This larger scale supports greater richness and diversity of epiphytic and related terrestrial species, underscoring the necessity of preserving large fragments of vegetation within cities.\u003c/p\u003e \u003cp\u003eDue to their sensitivity to environmental changes caused by disturbances like deforestation and human activities, many epiphytes are considered bioindicators of habitat quality because they respond to variations in humidity and temperature. This sensitivity makes them vulnerable to human-induced disturbances that alter community structure and composition (Larrea and Werner \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; K\u0026ouml;ster et al. \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Cach et al. 2014; Zotz \u003cspan citationid=\"CR90\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Carvajal-Hern\u0026aacute;ndez et al. \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Aoki-Gon\u0026ccedil;alves et al. \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). While some species that manage to survive the pressures of urbanization may exhibit effective strategies under stressful conditions, ferns of the genus \u003cem\u003ePleopeltis\u003c/em\u003e demonstrate poikilohydry, allowing them to reduce their exposed leaf area and minimize water loss in extreme conditions. This enables them to survive with only 25% of their water content until they receive a water pulse. (Moran \u003cspan citationid=\"CR60\" class=\"CitationRef\"\u003e2004\u003c/span\u003e; Hietz \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2010\u003c/span\u003e). Species of the genus \u003cem\u003eThelypteris\u003c/em\u003e thrive in warm climates and open areas. Examples include \u003cem\u003eMacrothelypteris torresiana\u003c/em\u003e and \u003cem\u003eChristella dentata\u003c/em\u003e, introduced species that have naturalized in tropical regions worldwide, demonstrating how these plants flourish in disturbed environments (Mehltreter \u003cspan citationid=\"CR57\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Fawcett and Smith \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Other adaptations encompass reduced size and the presence of leaf scales, which help repel solar radiation and absorb nutrients and water from the environment. This adaptation has enabled species such as \u003cem\u003eTillandsia juncea, Tillandsia schiedeana, Pleopeltis furfuracea\u003c/em\u003e, and \u003cem\u003ePleopeltis thyssanolepis\u003c/em\u003e to survive in these altered environments (Page \u003cspan citationid=\"CR63\" class=\"CitationRef\"\u003e2002\u003c/span\u003e; Watkins et al. \u003cspan citationid=\"CR83\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Hietz \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Benzing \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2012\u003c/span\u003e; Zotz \u003cspan citationid=\"CR90\" class=\"CitationRef\"\u003e2016\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eEnvironmental modifications in the forests of Xalapa have resulted in a greater presence of generalist species adapted to the stressful conditions of the urban environment, which are characterized by high resilience to water fluctuations and tolerance to sunlight (e.g., \u003cem\u003ePleopeltis\u003c/em\u003e spp. and atmospheric \u003cem\u003eTillandsia\u003c/em\u003e species) (Hietz \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Benzing \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2012\u003c/span\u003e; Zotz \u003cspan citationid=\"CR90\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). This pattern has been documented in other studies related to urbanization and habitat fragmentation, where environmental conditions become more favorable for generalist species, thereby displacing disturbance-sensitive species (McKinney, \u003cspan citationid=\"CR56\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Kr\u0026ouml;mer et al. \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Reyes-Garc\u0026iacute;a et al. \u003cspan citationid=\"CR70\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). This indicates that, although the urban forests of Xalapa maintain considerable diversity, the current conditions restrict the presence of species with specific ecological requirements, which rely on less disturbed environments with greater stability in their microclimates.\u003c/p\u003e \u003cp\u003eConversely, in peri-urban forests like Clavijero and La Martinica, which face fewer current environmental pressures, species regarded as indicators for the conservation of mountain rainforests (\u003cem\u003eVittaria graminifolia, Alsophila firma, Elaphoglossum sartorii, Elaphoglossum vestitum, Lophosoria quadripinnata, Trichomanes capillaceum, Trichomanes reptans\u003c/em\u003e) were documented (Carvajal-Hern\u0026aacute;ndez et al. \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). However, in this study, when applying the IndVal to link the species to a specific site, it was found that only 26% of the total exhibit an indication value, with ten classified as generalists, as they were identified in at least three of the examined forests. This indicates that they flourish in their living environment, having adjusted to overcome the existing environmental pressures. The physiological (CAM metabolism) and morphological adaptations (such as the presence of scales, elongated leaves, or tank-like structures for water storage) of generalist species enable them to thrive and colonize areas that may present environmental stressors that other species cannot tolerate (G\u0026aacute;mez-Viru\u0026eacute;s et al. \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Aronson et al. \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Alvim et al. \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2021\u003c/span\u003e).\u003c/p\u003e"},{"header":"Conclusion","content":"\u003cp\u003eUrban forests can help mitigate conditions that negatively impact urban areas, such as the homogenization of diversity. The variation in vascular epiphyte richness and turnover found in peri-urban forests, compared to the lower variation in inner-city forests, indicates that changing environmental conditions in the latter favor generalist and stress-tolerant species. This dynamic underscore urbanization's role as a selective filter for biological diversity, limiting the presence of species with specific ecological needs. Conversely, areas on the city's periphery serve as vital reservoirs for the diversity of epiphytic species and their terrestrial relatives, particularly promoting the persistence of rare and conservation indicator species. This suggests that current environmental conditions support overall diversity and are likely to enhance the ecosystem services it provides.\u003c/p\u003e \u003c/div\u003e"},{"header":"Declarations","content":"\u003cp\u003e\u003cstrong\u003eAcknowledgments:\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eTo the Secretar\u0026iacute;a de Medio Ambiente del Estado de Veracruz, for the facilities to enter the Natural Protected Areas. To Ana Mar\u0026iacute;a Aquino Zapata, Rodrigo Carral Dom\u0026iacute;nguez for their support in the field work.\u003c/p\u003e\u003cp\u003e\u003cstrong\u003eFunding\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThis work was funded by the Consejo Nacional de Humanidades Ciencia y Tecnolog\u0026iacute;a (CONAHCYT) through \u0026nbsp;Frontier Science project in the group modality (64358). The first author (MBJP) received funding for her postdoctoral stay from 2023 to 2025 (4816369) through CONAHCYT.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;\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\u003cp\u003e\u003cstrong\u003eAuthor contributions\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eS.A.M. and C.I.C.H. contributed to the study conception and design. Fieldwork and data collection were conducted by M.B.J.P., S.A.M., C.I.C.H. and J.C.L.; S.A.M. prepared the map. M.B.J.P. and C.I.C.H. performed the analyses. The first draft of the manuscript was written by M.B.J.P. and C.I.C.H. All authors contributed to drafting and improving the manuscript. Each author read and approved the final version of the manuscript.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\n\u003cli\u003eAcebey A, Gradstein S, Kr\u0026ouml;mer T (2003) Species richness and h\u0026aacute;bitat diversification of bryophytes in submontane rain forest and fallows of Bolivia. J Trop Ecol 19(1):9\u0026ndash;18. https://doi:10.1017/S026646740300302X \u003c/li\u003e\n\u003cli\u003eAdhikari YP, Hoffmann S, Kunwar RM, Bobrowski M, Jentsch A, Beierkuhnlein C (2021) Vascular epiphyte diversity and host tree architecture in two forest management types in the Himalaya. Glob Ecol Conserv 27:e01544. https://doi.org/10.1016/j.gecco.2021.e01544 \u003c/li\u003e\n\u003cli\u003eAlvey AA (2006) Promoting and preserving biodiversity in the urban forest. 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Springer International Publishing, Cham.\u003c/li\u003e\n\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":false,"highlight":"","institution":"","isAcceptedByJournal":true,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"urban-ecosystems","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"ueco","sideBox":"Learn more about [Urban Ecosystems](https://www.springer.com/journal/11252)","snPcode":"11252","submissionUrl":"https://submission.nature.com/new-submission/11252/3","title":"Urban Ecosystems","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false},"keywords":"bromeliads, ecological indicators, ferns, peri-urban forests, urban green areas","lastPublishedDoi":"10.21203/rs.3.rs-6116120/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-6116120/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eEpiphytic plants and allied terrestrial groups, which are sensitive to changes in humidity and temperature, play a crucial role in understanding the dynamics of ecosystem disruption caused by human activity impact. In the conurbation of Xalapa-Banderilla, Veracruz, Mexico, urban and peri-urban forests have different levels of disturbance conditions. This study aimed to analyze the patterns of vascular epiphytes and related terrestrial plants in urban forests with varying transformation histories in Xalapa, located in the central region of Veracruz Mexico. Five sampling plots were established in each forest (three urban and two peri-urban) where the richness of these groups was recorded. In a sample of 1 ha, 103 species distributed among 58 genera and 22 families were recorded, with ferns being the most represented. The peri-urban forest \u0026ldquo;Clavijero\u0026rdquo; exhibited the highest species richness compared to the others. Overall, heterogeneity in species composition was observed between sites, being lower when comparing only urban forests. In peri-urban forests, species considered indicator species were recorded, while in urban forests, mostly generalist species adapted to stressful conditions were recorded. The urban forests studied are reservoirs of diversity, highlighting the importance of forests in the periphery that shelter rare and conservation indicator species, suggesting that the original environmental conditions are still being maintained for the benefit of diversity in general.\u003c/p\u003e","manuscriptTitle":"Urban forests as habitats for vascular epiphytes and allied terrestrial plants","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2025-03-03 17:05:37","doi":"10.21203/rs.3.rs-6116120/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"decision","content":"Revision requested","date":"2025-04-24T07:32:10+00:00","index":"","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2025-04-23T19:34:20+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"129327082182676837585279597456373587407","date":"2025-04-05T15:19:14+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"67887090168257709444902725341665165682","date":"2025-03-31T12:53:24+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2025-03-29T18:27:32+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"228899757306241796453093075277438980009","date":"2025-03-09T12:05:31+00:00","index":"hide","fulltext":""},{"type":"reviewersInvited","content":"","date":"2025-03-07T08:27:18+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2025-03-02T08:58:46+00:00","index":"","fulltext":""},{"type":"checksComplete","content":"","date":"2025-02-28T06:29:20+00:00","index":"","fulltext":""},{"type":"submitted","content":"Urban Ecosystems","date":"2025-02-26T21:29:15+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"urban-ecosystems","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"ueco","sideBox":"Learn more about [Urban Ecosystems](https://www.springer.com/journal/11252)","snPcode":"11252","submissionUrl":"https://submission.nature.com/new-submission/11252/3","title":"Urban Ecosystems","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false}}],"origin":"","ownerIdentity":"39f146f9-10d5-41d9-8020-b5e6d2fec239","owner":[],"postedDate":"March 3rd, 2025","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"published-in-journal","subjectAreas":[],"tags":[],"updatedAt":"2025-06-30T15:59:19+00:00","versionOfRecord":{"articleIdentity":"rs-6116120","link":"https://doi.org/10.1007/s11252-025-01756-w","journal":{"identity":"urban-ecosystems","isVorOnly":false,"title":"Urban Ecosystems"},"publishedOn":"2025-06-26 15:56:56","publishedOnDateReadable":"June 26th, 2025"},"versionCreatedAt":"2025-03-03 17:05:37","video":"","vorDoi":"10.1007/s11252-025-01756-w","vorDoiUrl":"https://doi.org/10.1007/s11252-025-01756-w","workflowStages":[]},"version":"v1","identity":"rs-6116120","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-6116120","identity":"rs-6116120","version":["v1"]},"buildId":"8U1c8b4HqxoKbykW_rLl7","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}