Plant Systematics and Evolution (2021) 307:52 https://doi.org/10.1007/s00606-021-01773-0 ORIGINAL ARTICLE Distribution patterns, ecological niche and conservation status of endemic Tillandsia purpurea along the Peruvian coast Francisco Villasante Benavides1,2 · G. Anthony Pauca‑Tanco2 · C. R. Luque‑Fernández1,2 · Johana del Pilar Quispe‑Turpo3 · Luis N. Villegas Paredes1,2 · Alexander Siegmund5,6 · Marcus A. Koch4,5 Received: 6 March 2021 / Accepted: 22 June 2021 / Published online: 26 July 2021 © The Author(s), under exclusive licence to Springer-Verlag GmbH Austria, part of Springer Nature 2021 Abstract Species distribution modeling and assessment of the possible current conservation status for loma-forming Tillandsia purpurea Ruiz & Pavón in Peru were performed. This species is considered an epiarenic species that lives under hyperarid conditions, where its main source of water and nutrients comes from the fog of the Pacific coast. For the distribution modeling, 63 records from different sources of information were used, including a current field survey. Locations covered the whole range of the species´ known distribution along the Peruvian coast, and respective elevations lie between 0 and 2000 m a. s. l. Likewise, 27 environmental variables were used, including bioclimatic and eco-geographical ones, to determine the corresponding ecological niche and compare between actual and potential distribution. The conservation status was estimated according to the criteria recommended by the IUCN red list. High probability values were obtained predicting the occurrence of T. purpurea and describing respective environmental conditions such as altitudinal distribution between 400 and 1200 m and predominant southwest exposure of habitats. The conservation status of T. purpurea was supposed between "least concern" and near threatened, recommending that this species should be placed into the latter category and considering recurrent threats by direct anthropogenic impact and climate change verified during the field surveys. Keywords Atacama desert · IUCN status · Peru · Sechura desert · Species distribution modeling · Tillandsia purpurea Introduction Handling editor: Dietmar Quandt. * Francisco Villasante Benavides jvillasanteb@unsa.edu.pe * Marcus A. Koch marcus.koch@cos.uni-heidelberg.de 1 Departamento Académico de Biología, Universidad Nacional de San Agustín de Arequipa, Arequipa, Peru 2 Instituto de Investigación de Ciencia y Gestión Ambiental (ICIGA), Universidad Nacional de San Agustín de Arequipa, Arequipa, Peru 3 Escuela Profesional de Ingeniería Ambiental, Universidad Católica San Pablo, Arequipa, Peru 4 Centre for Organismal Studies, Heidelberg University, Heidelberg, Germany 5 Heidelberg Center for the Environment (HCE), Heidelberg University, Heidelberg, Germany 6 Department of Geography, Research Group for Earth Observation (rgeo), Heidelberg University of Education, Heidelberg, Germany Bromeliaceae is a monocot family restricted to the American continent, and it presents about 3400–2600 species, ranking seventh in species diversity, and being surpassed only by Orchidaceae, Asteraceae, Fabaceae, Rubiaceae, Poaceae and Melastomataceae (Smith and Till 1998; Ulloa-Ulloa et al. 2017). Members of the family are adapted to a wide range of habitats spanning ecological conditions from hyperarid deserts to tropical rainforests (Smith and Till 1998). Particularly, the South American deserts, known as the Atacama and Sechura (located in the coastal area of Peru and Chile), are home to conspicuous communities of bromeliads (known as tillandsiales), which are dominated by species of the genus Tillandsia, and survive water scarcity due to the fog incoming from the Pacific Ocean (Rundel et al. 1997; Pinto et al. 2006; Mostacero et al. 2007). Tillandsia plants in the desert have adapted successfully; their CAM metabolism and their morphological modifications, such as the rosette leaves (in whose armpits water is accumulated), the lack of functional roots (only as an anchorage to the substrate in the 13 Vol.:(0123456789) 52 F. Villasante Benavides et al. Page 2 of 14 earliest stages of growth) and the presence of trichomes on the leaves (which absorb water and nutrients), have made it possible for them to survive in these extreme environments (Pinto 2005). In Peru, tillandsiales are either monospecific or are represented by two or even more species of Tillandsia, being T. purpurea Ruiz & Pavón, T. latifolia Meyen, T. marconae Till & Vitek, T. capillaris Ruiz & Pavón, T. paleacea C.Presl, T. landbeckii Philippi and T. recurvata (L.) L., the ones registered for these communities (Ono 1986; Mostacero et al 2007). Among these species, T. purpurea and T. latifolia are the most widely distributed loma-forming taxa in Peru (Ono 1986; Rundel et al. 1991; Arakaki and Cano 2003; Pinto 2005; Pinto et al. 2006; Mostacero et al. 2007; Aponte and Flores 2013; Pauca-Tanco et al. 2020). Tillandsia purpurea is a common species in tillandsiales; it is endemic in Peru and is distributed along the coast ranging over 2250 km from the departments of La Libertad in the north to Tacna in the south (Smith and Downs 1977; Mostacero et al. 2007; Ulloa-Ulloa et al. 2017; Zizka et al. 2020; Pauca-Tanco et al. 2020). Its taxonomy is a controversial issue. T. purpurea has been associated with T. straminea Kunth (Smith and Downs 1977; Barfuss et al. 2016), although it is distributed at higher altitudes in the Andes of Peru and Ecuador. However, some authors suggest that both species should be treated as different taxa (UlloaUlloa et al. 2017; Zizka et al. 2020) and it is argued that T. straminea is probably the epiphytic ancestor of T. purpurea (T. purpurea is epiarenic) (Smith and Downs 1977). Consequently, Barfuss et al. (2016) introduced a T. purpurea species complex with at least four different species. At present, the communities of Tillandsia are poorly studied such as for the individual species, despite being some endemic to Peru (e.g., T. purpurea) (Ulloa-Ulloa et al. 2017; Zizka et al. 2020) or even local endemics (T. werdermannii) (Brako and Zarucchi 1993). Most aspects of Peruvian epiarenic Tillandsias systematics, evolutionary history, distribution, ecology or threatening, e.g., by climate change are largely unknown. The conservation status of T. purpurea has currently been qualified as near threatened or of least concern (NT or LC, respectively) (Zizka et al. 2020), and some threats have been listed, the main ones being human activities (conflicts of the sites with human recreation, construction work, environmental pollution) and also climate change (Pinto et al. 2006; Schulz et al. 2010; Pauca-Tanco et al. 2020). Ecological niche models (ENMs), also known as species distribution models (SDMs), are tools that help to understand how a taxonomic entity is related to the geographic space considering its environmental requirements, which, in general, is most often simplified and reduced to temperature- and precipitation-related variables (Peterson et al. 2011; Guisan and Thuiller 2005; Ibarra-Montoya et al. 2012). Evaluation of the ecological niche and its distribution in space helps to predict the real distribution (species 13 distribution modeling, SDM), e.g., if field surveys are difficult, and might also provide first ideas about potential distribution ranges suitable for conservation and restoration activities. ENM may also help to predict future distribution scenarios and, thereby, test for the impact of climate change scenarios (Ferrier 2002). Currently, there are some applications for ENMs, and one of the most used tools is MaxEnt (maximum entropy) (Phillips et al. 2006), which uses occurrence data and bioclimatic variables. Over time, the use of MaxEnt has increased, giving rise to concepts and methodologies allowing a more rigorous statistical application, thus ensuring efficient generation of valid and biologically meaningful models (Muscarella et al. 2014; Kass et al. 2018; Cobos et al. 2019). ENM has been applied in Bromeliaceae with few studies (e.g., Zizka et al. 2009; Judith et al. 2013; Aguiar-Melo et al. 2019) and contributing substantially to our understanding of Bromeliaceae ecology and evolution. There is a major need for efforts to understand the various aspects of the ecology of biological communities in the desert of Peru. In particular, the communities of Tillandsia, including those made up by Peruvian endemic species such as T. purpurea, need respective attention, since their habitats are currently being severely altered and populations are decreasing rapidly (Pauca-Tanco et al. 2020). Here we aim to close some of these questions and apply the ENM in combination with a field study along the entire Peruvian coast. Ecological preferences for the endemic Tilladsia purpurea are described, and a reliable IUCN (2014) conservation status for this species has been elaborated based on its distribution area and pressures observed in the field and literature. Material and methods Study area The study region stretches along the entire coastal area of Peru, ranging from latitude 3° 23′ (Tumbes) to 18° 21′ (Tacna), extending approximately 2250 km in length and with an altitude ranging from 0 to 2000 m a. s. l. (Fig. 1). Most of Peru’s coastal zone is characterized by desert and is defined as one of the driest in the world (Rundel et al 1991; Dillon et al. 2009). Rainfall is very limited due to three main conditions that occur during the southern hemisphere winter (the Humboldt current, the Pacific anticyclone and a thermal inversion layer) with low altitude stratocumulus clouds (from 800 to 1200 m) entering the inland from the coast. While rising up the coastal mountains, the fog condenses and gives rise to the so-called oases of life, which are locally known as coastal lomas or lomas (Ferreyra 1993; Jiménez et al. 1997). The soil in this area is mostly clay sand, with scattered rocks; the terrain is rugged close to the coastal Distribution patterns, ecological niche and conservation of Tillandsia purpurea Page 3 of 14 52 re-presented with inadequate coordinates (farmland, urban centers or at open sea). It was also taken into account that the distance between individual records was at least 1 km in order to eliminate the autocorrelation of the records with respective environmental variables (Peterson et al. 2011; Boria et al. 2014). The delimitation of a calibration area (potential distribution range) was defined considering the influence of the fog coming from the ocean, and the dispersal capacity of T. purpurea (exploration capacity and potential colonization), since it is known that an incorrect choice of the extension of the area can result in severe artifacts, which further negatively influence final results (Ono 1986, Ogawa 1986, Ferreyra 1993, Barve et al. 2011, Romero-Álvarez et al. 2020, Martinez-Mendez et al. 2016, Soberón and Peterson 2005). The zone was also defined using an altitudinal range from 0 to 2000 m a. s. l., on the basis that Galán de Mera et al (2010) bioclimatically identify that zone as hyperarid and ultra-hyperarid, being T. purpurea an indicator species. The boundaries were taken to the north (Tumbes) and south (Tacna) of Peru. Treatment of environmental variables Fig. 1 Study area located along the coast of Peru between sea level up to 2000 m a. s. l. mountain range, and the remainder are extensive plains, only interrupted by valleys and streams. Occurrence data of Tillandsia purpurea and calibration area For the delimitation of the species T. purpurea, we followed Smith (1936), since some authors currently consider T. straminea as a synonym of T. purpurea (Smith and Downs 1977). In this taxonomical sense, T. purpurea is found in arid coastal environments only, whereas T. straminea is found above 2000 m a. s. l. In total, 63 geographical records of the species under study were obtained, 26 of which were retrieved from GBIF digital database (GBIF 2020), and most of these corresponded to samples preserved in herbaria and some field observations. Records from 1778 to 2018 were temporarily covered, and the database was downloaded in June 2020. From the total amount, 10 are sourced from bibliographic reviews (Smith and Downs 1977; Pinto 2005; Whaley et al. 2019) and 27 are based on field surveys made during the years 2018 and 2019. The filtering of the geographical locations was performed with NicheToolBox package (OsorioOlvera et al. 2020) in RStudio (RStudio Team 2020) attempting to eliminate records that were found to be duplicated or A total of 27 variables were obtained, which had approximately 1 km2 (30 arcsen) of resolution, of which 19 correspond to bioclimatic layers (Fick and Hijmans 2017) and the others to ecogeographic variables. Within the ecogeographic variables, slope and orientation were obtained from the digital elevation model ASTER GDEM (30 m resolution), retrieved from the MINAM server (http://geoservido rperu.minam.gob.pe) and using the QGIS program ver. 3.14 (QGIS 2020). The aridity index was obtained from Trabucco and Zomer (2019). Variables of solar radiation and water vapor pressure were obtained from Fick and Hijmans (2017), too. A human footprint was extracted from SECAD-NASA (Venter et al. 2016), and values for vegetation cover were generated from the vector information provided by MINAM. In order to eliminate the correlation between the variables obtained and to prevent models with over-fits from being produced (Peterson and Nakazawa 2008), a bivariate correlation analysis was performed using the NicheToolBox package (Osorio-Olvera et al. 2016) and discarding those variables with redundant information (threshold = 0.85). As a consequence of this analysis, 18 independent variables were obtained, which were used for the subsequent modeling of T. purpurea. Modeling and evaluation The modeling of the ecological niche and the potential distribution range were carried out with the Wallace v1.0.6.1 package (Kass et al. 2018) running RStudio (RStudio Team 13 52 F. Villasante Benavides et al. Page 4 of 14 2020). The choice of the package was due to the attributes it has: intuitive, flexible and offering substantial reproducibility while evaluating the choice of the most appropriate model used. For the processing, excluding the previously cleaned points, a posterior verification was made using the tool Thinning distance (1 km) for the separation between individual occurrence records; the number of sample Background Point was set to 5000; the non-spatial method was selected with the Random K-Fold option (Folds = 3) for the partition of the occurrence points. The model was calculated using the Maxent algorithm v.3.4.1 (Phillips et al 2006). For this procedure, all the entity’s classes and their combinations were selected in features (there are a total of 4 classes and 13 combinations), and for the Regularization multipliers values 1 and 2 were established. For the evaluation of the obtained models and their selection, the values of AUC (Area under curve), AIC (Akaike information criterion) and omission rate were used, which are provided by the ENMeval package (Muscarella et al. 2014) and implemented in Wallace. Additionally, 13 validation points (not used in the model) were used for evaluation through the partialROC parameter implemented with the NicheToolBox package (Osorio 2020). State of conservation The conservation status of T. purpurea was estimated according to the criteria recommended by the IUCN Red List (Standards and Requests of the IUCN Subcommittee 2014), for which the same georeferences of the distribution model training and the records of validation (76 records in total) were used, calculating the occupation area (AOO, area of occupation) and the degree of occurrence (EOO, extent of occurrence) (Gaston and Fuller 2009). Extent of occurrence (EOO) is that area which lies within the outermost geographic limits to the occurrence of a species, and the area of occupancy (AOO) is that within those outermost limits over which it actually occurs. This means that EOO values are always higher than AOO. All these analyzes were performed with the ConR package (Dauby et al. 2017) in Rstudio (RStudio Team 2020). For the parameters used, the Researchers’ own knowledge of the biology of the species was applied, since many records do not come from databases, but from our field inspection and verification visits. This knowledge includes, for example, a subpopulation distance (radius = 7 km) and using a grid resolution of 2 km2 in size as recommended by the IUCN. Finally, when the recommendation of the conservation status differed between the AOO and EOO evaluations, we followed the criteria used by Knapp (2013), where the highest value was chosen, i.e., most threatened. This was followed as a criterion for conservation action, which will also be supported by the threats observed on the field for the populations evaluated along the coast. Additionally, a shapefile showing the Protected Areas of Peru was incorporated (obtained from World Database on Protected Areas, https://www.protectedplanet.net) aiming to quantify how many of the estimated subpopulations are nationally protected. Results A total of 63 occurrence data points were used for the creation of the models, where the best result corresponded to LQHP_1 (AUC = 0.962, PartialROC = 0.68, α = 0.05), for which it is established that the predicted area for T. purpurea (Threshold = 0.31) corresponds to 16,786.3 km2. The main variables that defined the model were altitude, aridity index, orientation, evapotranspiration, temperature seasonality, max temperature of warmest month, annual precipitation and precipitation of warmest quarter (Table 1), where the distribution of T. purpurea was found along a wide altitudinal range (66–1952 m a. s. l.) and showing an average elevation around 786 m a. s. l., likewise, the ranges of aridity and orientation where they can present are widely variable, preferring in this last one the orientation to the southwest (SW). On the variables of temperature and precipitation, it occurred in areas with temperatures above 20 °C and low rainfall characteristics with on average 50 mm per year and where there were high values of daily evapotranspiration (range 880–2171 mm*day−1). The distribution of T. purpurea is located mainly along the coast of Peru, ranging from Lambayeque in the north (6° Table 1 Main environmental variables that defined the distribution model of Tillandsia purpurea in Peru Variable Altitude m a. s. l. Aridity index Orientation (degrees) Evapotranspira- bio 4 (°C) tion (mm*day−1) bio 5 (°C) bio 12 (mm) bio 18 (mm) Mean (SD) Max Min IC (95%) 786.1 ± 373.3 1952 66 5.07 124.1 ± 137.9 1144 0 1.87 202.8 ± 80.8 359.9 0 1.10 1639.8 ± 276.5 2171 880 3.76 24.8 ± 1.5 28.1 20.2 0.02 50.3 ± 45.4 258 3 0.62 38.4 ± 45.4 272 0 0.62 10.8 ± 2.6 18.3 5 0.03 bio 4 temperature seasonality, bio 5 max temperature of warmest month, bio 12 annual precipitation, bio 18 precipitation of warmest quarter 13 Distribution patterns, ecological niche and conservation of Tillandsia purpurea 52′ S) to Tacna (18° 12′ S) in the south; however, the resulting area of distribution is not continuous. An isolated area is observed in Cerro Cantarilla-Saña in the extreme north (Fig. 2), approximately 152 km apart from the closest T. purpurea population close to the town Chicama (Trujillo). The entire distribution range continues to the south of Lima (San Page 5 of 14 52 Vicente de Cañete), is interrupted through almost the entire department of Ica and continues again toward the south near the town Marcona (Fig. 2a, b). In southern Peru, in the departments of Arequipa, Moquegua and Tacna (Fig. 2c, d), the distribution is continuous and suitable habitats are also found along the coast with the most important geographical Fig. 2 Potential distribution of Tillandsia purpurea in coastal regions of Peru (green dots are presences). a Northern Peru (Lambayeque, La Libertad and Ancash), b central and southern Peru (Lima and Ica), c southern Peru (Arequipa), d southern Peru (Moquegua and Tacna) 13 52 F. Villasante Benavides et al. Page 6 of 14 Table 2 Main characteristics of the environmental variables that determined the different areas of distribution (north and south) of Tillandsia purpurea in Peru Variable Altitude (m a. s. l.) Mean (SD) Median (Q1, Q3) Range Aridity index Mean (SD) Median (Q1, Q3) Range Orientation (degrees) Mean (SD) Median (Q1, Q3) Range Evapotranspiration Mean (SD) Median (Q1, Q3) Range bio 4 (°C) Mean (SD) Median (Q1, Q3) Range bio 5 (°C) Mean (SD) Median (Q1, Q3) Range bio 12 (mm) Mean (SD) Median (Q1, Q3) Range bio 18 (mm) Mean (SD) Median (Q1, Q3) Range North (N = 2000) South (N = 2000) 531.0 (313.7) 439.0 (272.0, 745.5) 69.0–1537.0 978.5 (293.0) 1001.0 (800.0, 1158.0) 191.0–1943.0 247.0 (136.9) 212.0 (161.0, 289.0) 24.0–1115.0 35.8 (27.8) 28.0 (16.0, 49.0) 0.0–273.0 202.9 (82.9) 215.2 (148.4, 263.4) 0.1–359.8 203.0 (76.7) 207.5 (152.9, 255.4) 0.0–359.5 1358.0 (134.8) 1392.0 (1284.0, 1454.0) 905.0–1657.0 1843.6 (139.6) 1852.0 (1736.0, 1954.0) 1476.0–2171.0 879.4 (120.6) 867.0 (784.0, 962.0) 610.0–1259.0 1216.7 (234.5) 1223.0 (989.0, 1403.0) 836.0–1784.0 251.4 (12.6) 255.0 (243.0, 261.0) 214.0–276.0 244.8 (16.2) 246.0 (232.0, 258.0) 203.0–281.0 90.4 (45.1) 82.0 (57.0, 119.0) 9.0–250.0 22.5 (14.1) 18.0 (12.0, 30.0) 3.0–129.0 78.4 (47.0) 66.0 (46.0, 104.0) 6.0–263.0 10.3 (6.6) 9.0 (5.0, 14.0) 0.0–73.0 P value < 0.001 < 0.001 0.244 < 0.001 < 0.001 < 0.001 < 0.001 < 0.001 Applied statistics: Kruskal–Wallis test, with a 95% confidence interval based on 2000 randomized subsamples for each area bio 4 temperature seasonality, bio 5 max temperature of warmest month, bio 12 annual precipitation, bio 18 precipitation of warmest quarter feature, namely presence of coastal mountain ranges being only interrupted by valleys. An important characteristic in the distribution of T. purpurea in southern Peru is that populations occur at greater distances from the sea coast, especially in Tacna, with distances greater than 44 km. Through the comparison between the southern zone (Ica, Arequipa, Moquegua and Tacna) and northern Peru (Lambayeque, La Libertad, Ancash and Lima), in terms of the ideal zones within the variables that defined the model (Table 2, Fig. 3), it is possible to observe some patterns. It is observed that in the southern zone, the aridity index presents lower values than the northern zone. The orientation is similar in both zones. Evapotranspiration is notably higher in 13 the southern zone. Toward the north, T. purpurea is located in low areas, while toward the south it prefers higher areas. In the case of the temperature seasonality (bio 4), toward the south it is frequent to find higher values than toward the north. For the maximum temperature in the warmest month (bio 5), there is some overlap in the values for the areas evaluated; however, the south presents low values more often than the north. The annual precipitation (bio 12) shows higher values for the northern zone as opposed to the southern zone. In the case of bio 18 (precipitation of warmest quarter), the northern zone frequently presents higher values than the southern zone. Finally, the statistical comparison between the northern and southern zones (Table 2) shows Distribution patterns, ecological niche and conservation of Tillandsia purpurea Page 7 of 14 52 Fig. 3 Comparative boxplot of the environmental conditions which defined the presence of the Tillandsia purpurea population on the coast of Peru. ***Indicates significant different means with p < 0.001 (see Table 2) that there are significant differences (p < 0.001) for the most suitable areas of T. purpurea in the south and north of Peru. Conservation status The conservation status of T. purpurea according to the evaluation with the ConR package, would be considered between the conservation categories of "Least Concern" (LC) or "Near Threatened" (NT). The characteristics for this species resulted in an AOO of 272 km2, and an EOO of 200,611 km2. A total of 29 subpopulations were found along the Peruvian coast, based on an estimated spacing for this parameter of 7 km. In addition, 46 localities are reported based on a spacing resolution of 80.3 km. Regarding the number of records, there is a greater abundance towards the extreme 13 52 F. Villasante Benavides et al. Page 8 of 14 Fig. 4 Maps generated by IUCN.eval function for T. purpurea occurrences in coastal regions of Peru. The top map shows a convex hull used for calculating the EOO (Shown for the species as a gray polygon with the occurrences points as black dots). Protected areas (PA) are shown as polygons within Peru, where all occurrence points are outside of PA. The delimitation of locations is displayed with pink squares and sub-populations with black circles near the occurrences point (zoom image). The number of records of T. purpurea per 1° sample units is also shown with the small inserted map south of Peru (more than 12 records). On the other hand, the evaluation regarding the protected areas for Peru shows that no population system of T. purpurea is found within a conservation area. (Fig. 4). Discussion Observed and suitable but not realized, potential distribution ranges The AUC value for the selected model is high (0.962), being considered reliable to predict the existence of adequate 13 Distribution patterns, ecological niche and conservation of Tillandsia purpurea environmental conditions and indicates a good performance with low levels of commission errors, so it identifies quite well all the locations where these species have been reported or observed (Phillips et al. 2006; Ortíz-Yusty et al. 2014). Nevertheless, for the additional validation of the selected model (with presence points independent of the modeling), a test was applied using the partial ROC curve (Peterson et al. 2008), being satisfactory, since the model is statistically different from a process performed by chance (p < 0.05). The partial ROC curve is recommended in ecological niche models, since the error of omission is sufficiently low and is compensated by the adjustments established by the user (Lobo et al. 2008). As expected, the potential distribution areas of T. purpurea are located along the Cordillera Costera of Peru and sometimes reaching the lower slopes of the Cordillera Occidental of the Andes. Many of the modeled areas are shown as considerably large, elongated, and relatively continuous areas, except for the presence of small potential areas in central Peru. The first is located between the north of the department of La Libertad and the south of the department of Lambayeque, and the second is located in the center of the Peruvian coast, between the departments of Lima and Ica (Fig. 2 and 3a, b), of which we will talk later. A singular result in the distribution is what happens in Tacna, in southern Peru, where the modeled potential areas are pushed inland, because the coastal cliff disappears and the slope gently ascends from the coast allowing the entry of a greater amount of fog inland reaching up to about 60 km from the coast (Pinto et al. 2006). The potential distribution obtained is explained by the variables that contributed the most to the model, especially the low aridity index, which according to the UNEP (1997) generalized climate classification scheme places the entire study area as hyperarid, this due to low precipitation and high evapotranspiration, the latter being several times higher than precipitation (Table 1); this extremely arid climatic condition has been described by various authors (Rundel and Dillon 1998; Hesse 2012; Aponte and Flores 2013; Koch et al. 2019). Although hyperaridity is the most outstanding condition of the climate in the study area, it should also be noted that the main contribution of water for this species comes from the fog (variable which has not been considered in the model as there is no data for it). This is why some researchers have pointed out that T. purpurea, as in most species of the genus Tillandsia, is dependent on fog water (Rundel and Dillon 1998; Pinto et al. 2006; Zizka et al. 2009; Hesse 2012). The existence of the coastal mountain range so close to the Pacific Ocean favors that the altitude, the orientation and the slope can create the optimal conditions to predict, with high probability, the presence of T. purpurea (Cereceda 1999; Pinto et al. 2006; Hesse 2012; Pauca-Tanco et al. Page 9 of 14 52 2020). In general, the communities of Tillandsia are preferably located in places with little slope and oriented toward the southwest. It is in such places that the wind currents which carry the humidity of the Pacific Ocean will collide with the Tillandsia (Borthagaray et al. 2010; Hesse 2012; Koch et al. 2019; Pauca-Tanco et al. 2020). These topographic characteristics are decisive for the maintenance of T. purpurea populations, since these plants depend on the water that they trap from the fog, the most important environmental condition along the Peruvian coast. Therefore, it is paramount to know the conditions of habitat and niche necessary for this species, along with the orientation of the hills (S and SW) where the tillandsiales are preferentially located (PaucaTanco et al. 2020). It is known that, morphologically, the plants of these communities present efficient adaptations to the arid conditions of their environment. The presence of fogs is the reason they have developed some strategies such as the lack of functional roots and presenting small scales on their stems and leaves, which provide protection against radiation and help take advantage of the surrounding moisture (Rauh 1985; Ono 1986; Haslam et al. 2003). The greater extension of the distribution area obtained by the model, in comparison with the real distribution of the species, shows in the form of isolated and distant patches observed in the field and described by several authors (Ono, 1986; Rundel et al. 1991; Arakaki and Cano, 2003; Pinto et al. 2006; Mostacero et al. 2007; Aponte and Flores, 2013), this may be due to the fact that the ecological niche models are explicitly designed to predict areas of potential distribution, so they are generally broader than the actual ranges (Peterson et al. 2008). However, the fact that they present a greater extension does not indicate an error. Instead, it indicates that these would be the places that meet the environmental conditions for the development of the species, but there could be some limiting factor still unknown (Hesse 2012), so that they may not be colonized; or if they were occupied in the past, some environmental change may have occurred that locally extinguished the populations of these places, as shown by the discovery of a fossil of Tillandsia in the dunes of Pampa Blanca in Ica. However, currently there are no more tillandsiales to be found in this location (Hesse 2012). In relation to the above, during visits to the field along the Peruvian coast, several sites with dead tillandsiales populations have been observed, which could indicate that environmental conditions could be changing, especially in relation to the availability of water from fog (Schulz et al. 2010). The significant differences between the environmental variables in the northern distribution areas of Peru (Lambayeque, La Libertad, Ancash and Lima) and those of the south (Ica, Arequipa, Moquegua and Tacna) (Table 2, Fig. 3) can be explained due to the greater aridity in the south (aridity index = 36.3 in the south vs. 247.0 in the 13 52 Page 10 of 14 F. Villasante Benavides et al. Fig. 5 a Closeup of a flower of Tillandsia purpurea, b emerging capsules of T. purpurea, c vital tillandsiales of T. purpurea in Camaná (Arequipa), d die back of T. purpurea in the Pampa Clemesí (Moquegua), e decorative “figure” made with T. purpurea plants north), therefore, less annual precipitation (22.1 mm in the south vs. 89.6 mm in the north) and greater evapotranspiration (1841.8 mm*day−1 vs. 1356.8 mm*day−1). The difference in altitude above the sea is also notable, since in the north it reaches a lower average (526.1 m a. s. l.) than in the south (971.8 m a. s. l.). All these climatic and topographic characteristics could determine a greater dependence on water from the fog in populations of T. purpurea 13 in the south than in the north, since, according to Rundel et al. (1991), the formation of dense stratocumulus clouds occurs particularly in the central and southern coast of Peru, in addition to populations of T. purpurea being preferably located above 900 m a. s. l. In relation to the discontinuity in the distribution of T. purpurea in central Peru—south of Lima and almost the entire department of Ica—except for the presence Distribution patterns, ecological niche and conservation of Tillandsia purpurea of small areas of distribution, it could be due to the fact that the central zone of the coast Peru (Pisco—San Juan, ~ 14–15° S) presents the most intense coastal surface winds (Gutiérrez et al. 2011), with typical speeds that exceed 10–15 m s−1 and that blow from the south–southeast (S–SE), and which are known locally as Vientos Paracas causing episodic wind erosion events (dust storms) (Briceño-Zuluaga 2017). These winds would prevent the deposit of seeds and the anchoring of these plants in the ground, and these winds have the form of a low-level atmospheric jet along the coast leaving a coastal strip without clouds that extends from 20 to 30 km sea indoors (Briceño-Zuluaga, 2017), causing a lower presence of fogs and, as a consequence, little or no water availability for the tillandsias, leading to extreme aridity (Whaley et al. 2019), in addition to the intensity, frequency and interannual reliability of the fog vary widely and are probably insufficient for the development of tillandsiales (Hesse 2012). This adverse condition for T. purpurea can be considered as a biogeographic barrier that could have decreased or eliminated the genetic flow between the populations located in the north of Peru in relation to those in the south, for which we propose the need to carry out studies on genetic diversity among these populations. In the area between La Libertad and Lambayeque, it seems that humidity and topography would also be conditioning the presence of T. purpurea, since geographically it has few elevations close to the sea (lomas). Indeed, between the Trujillo and Chiclayo areas, the flat terrain is the most common topographic feature, so the extensive plains in this area would prevent the presence of this species since it does not meet the necessary slope, orientation and altitude conditions for the presence of T. purpurea, or for its colonization, especially due to the inability of plants to obtain water from the fogs from the Pacific. Implications of distribution patterns on the conservation status We suggest that T. purpurea be placed in the near threatened (NT) category. Although the categories obtained in our evaluation were similar to those of Zizka et al. (2020), some differences can be observed in terms of AOO (272 km 2 for this study, vs. 140 km 2), and EOO (200,611 km2 for this study, vs. 461 541 km2). It is possible that these differences are explained by the possibility that Zizka et al. (2020) used free GBIF data for their evaluation, so it is possible that they have not been properly cleaned and verified (see, on his map you can see some records of T. purpurea in the Peruvian jungle). In our study, the GBIF records were also used, but with a thorough review, verification and careful cleaning in the location of the points, some of them being confirmed in Page 11 of 14 52 the field visits between February and March 2020; therefore, quite accurate and reliable information was available. Furthermore, the IUCN has not considered T. purpurea in any threat category, probably due to insufficient information on this taxon and its taxonomic implications. Given that, in this study and according to Ulloa et al. (2017) and Zizka et al. (2020), T. purpurea is different from T. straminea, the former would be considered endemic to Peru and restricted to the coastal zone. Therefore, according to the threats identified in its habitat, T. purpurea deserves to have a threat category. In addition, assuming the conservative criterion of Knapp (2013) by which we consider T. purpurea as near threatened (NT), and given that during field verifications it was observed that, like Pauca-Tanco et al. (2020) and Pinto (2006), the most common negative impacts on T. purpurea communities were habitat destruction (due to urban or agricultural expansion), intentional burning, removal or relocation of individuals for decorative purposes and the death of populations (Fig. 5e, d) and accumulation of solid waste such as plastics and paper was observed (especially in areas near roads). Finally, considering that the distribution of this species covers the entire Peruvian coast and depends essentially on the fog for its subsistence, this variation in the layers of fog is a latent threat of greater impact in a climate change scenario (Klemm and Lin, 2016), as has already been demonstrated in an investigation on the Peruvian-Chilean Pacific coast on the variation of fog and its relationship with fog-dependent vegetation (Muñoz et al. 2016; Koch et al. 2020), and in this context emphasizes the use of T. purpurea as an indicator of climate change (Rundel et al. 1997) for this variable. Currently, there are indications that humidity in the coastal zone is changing its regime (Schulz et al. 2010), which can alter the subsistence of the populations of this species, especially due to phenological changes such as in the flowering and fruiting periods, as documented for other species (Molau et al. 2005; Miller-Rushing and Primak, 2008). Acknowledgements We thank Mr. Jorge Ignacio for his collaboration in the field work in Tacna (Peru). Author contributions FV, MAK and AS conceived the project and the experiments. FV, CL, AP, JP and LV conducted the experiment(s). AP and CL analyzed the results. FV, AP and CL drafted the manuscript. All authors contributed to the final manuscript draft. Funding F. Villasante Benavides and collaborators acknowledge the financial support of the Consejo Nacional de Ciencia, Tecnología e Innovación Tecnológica (Concytec) through its executing unit the Fondo Nacional de Desarrollo Científico, Tecnológico y de Innovación Tecnológica (Fondecyt) for this research work. 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