Aims: Ecosystems of the Tropical Andes include plant communities above 4,000 m in elevation, associated with wetlands known as bofedales. To enhance our understanding of them, we surveyed bofedal plant communities in the Peruvian Andes. Questions: Which are the most common bofedal plant communities, and what are their main characteristics? Study area: An east-to-west 68 km megatransect in Ayacucho and Huancavelica departments in Peru, the area of influence of a gas pipeline. Methods: We surveyed 127 (1 m × 1 m) permanent plots annually between 2017 and 2019 to assess plant communities, calculated diversity metrics, and applied non-parametric hypothesis testing analysis of similarities and multivariate analyses to the data. Results: We identified 13 plant communities with 3.5 to 11.7 mean species richness. Only seven were statistically different; the other six were rare and require additional surveys to define their status as independent communities. The Distichia muscoides-dominated community was found in most sites (90%), plots (55%), and along the entire elevational range we studied. D. muscoides, Plantago tubulosa, and Rockhausenia pygmaea were the most frequent species in the studied bofedales (in 30 of 31 sites). These species are usually cushion or carpet forming, so average plant cover was high in most plant communities where they occurred (89–98%). The seven plant communities (dominated by D. muscoides, R. pygmaea, Plantago tubulosa, P. rigida, Lachemilla diplophylla, Aciachne pulvinata and Juncus stipulatus) were consistent in their structural and compositional characteristics and maintained differences between them during our three-year study. Conclusions: We show that bofedal plant communities in the southern Peruvian Andes are more heterogeneous than the four broad types previously reported. This heterogeneity occurs at local site levels but also at landscape and regional scales. We highlight the importance of considering this heterogeneity when discussing and implementing management, restoration, and conservation actions in bofedales.

Taxonomic reference: WFO (2024)

Alpine vegetation, Andes, bofedal, diversity, peatland, Peru, wetland

The tropical Andes, a global biodiversity hotspot, contain diverse ecosystems and habitats (Young et al. 2007). A major factor contributing to this diversity is the east–west humidity gradient in the Central Andes, reflected in the differentiation of the humid Puna towards the eastern flanks and the dry Puna (Josse et al. 2009) facing the Pacific slopes (Killeen et al. 2007; Espinoza et al. 2015).

Within this gradient, the Central Andes contain plant communities above 4,000 m a.s.l. which are associated with wetlands and water-saturated soils. These communities are known by several local names (“turbera” in Colombia and Ecuador; bofedal in Ecuador, Peru and Bolivia, “vegas” or “mallines” in Chile and Argentina, Hergoualc’h et al. 2022). These wetlands are patchy and island-like in nature and are usually surrounded by large swards of grasslands and drylands. At the local level, they have a consistent and characteristic set of plant species, but at regional levels, plant abundances and composition are influenced by environmental factors such as elevation, topography, hydrology, geology, and wildlife grazing (Ruthsatz 2012; Valencia et al. 2013; Salvador et al. 2014; Oropeza 2019; Portal-Quicaña 2019; Izquierdo et al. 2020; Domic et al. 2021; Monge-Salazar et al. 2022). In addition to these natural factors, human uses of the bofedales (e.g., grazing areas for native and introduced livestock, water use, peat extraction) have also been deemed important in influencing their ecological processes and plant community composition (Ruthsatz 2012; Maldonado-Fonkén 2014; Chimner et al. 2019; Yager et al. 2019; Navarro et al. 2023).

According to the National Institute for Research on Glaciers and Mountain Ecosystems (INAIGEM), the bofedales in Peru include four major types of plant formations, named after the life-form(s) of the dominant and most conspicuous species: cushions (formed by e.g. Distichia muscoides, Oxychloe andina, Plantago rigida), carpets (e.g. Plantago tubulosa, Rockhausenia pygmaea), grasses and graminoids (e.g. Festuca spp., Calamagrostis spp., Carex spp., Eleocharis spp., Phylloscirpus spp.), and mosses and shrub wetlands (formed by e.g. Sphagnum spp., Andicolea spp.). Bofedales dominated by cushions are the most frequent type, especially in central and southern Peru. INAIGEM highlighted the heterogeneity of these ecosystems at vegetation and hydrological levels. Still, little information is available on grass- and graminoid-dominated and carpet bofedales, as well as on bofedales subjected to strong seasonal water availability (saturated only in the rainy season; INAIGEM 2023).

The INAIGEM classification, the first of its kind at national level, is based on previously published information for bofedales vegetation in Peru (Cooper et al. 2010; Ruthsatz 2012; Maldonado-Fonkén 2014; Salvador et al. 2014; Maldonado-Fonkén 2018; Polk et al. 2019; Portal-Quicaña 2019), expert consultations and ongoing studies. Similar plant formations have been described for Colombia, Ecuador, Bolivia, and Argentina (Ruthsatz 2012; Benavides and Vitt 2014; Loza Herrera et al. 2015; Ruthsatz et al. 2020; Domic et al. 2021; Izquierdo et al. 2022; Suarez et al. 2022). Several plant communities or even mixed communities within a single site have also been reported (Ruthsatz 2012; Maldonado-Fonkén 2014). Nevertheless, most of these studies focused on the dominant species and physiognomy of the plant communities, so a more comprehensive description is needed.

Within the framework of the Biodiversity Monitoring and Assessment Program (BMAP), a collaboration between the Smithsonian Institution and the PERU LNG company set in the southern Andes of Peru (Dallmeier et al. 2013), we studied the vegetation of the high-Andean wetlands along an east to west 68 km-long megatransect from Ayacucho to Huancavelica departments.

This contribution aims to identify and characterize the most common bofedal plant communities along the megatransect. In doing so, we attempt to answer the following guiding questions: Is Distichia muscoides the only dominant species, as commonly treated? How do patterns of diversity, structure, and composition of plant communities change along the megatransect? Are the diversity and vegetation cover values high or low compared with other reports? Can selected environmental factors, such as soil moisture and water table depth, explain floristic and plant community patterns?

Our study area corresponds to the area of influence of the PERU LNG pipeline (LNG: liquified natural gas, Figure 1, Table 1), encompassing a variety of habitats between 4,200 and 4,900 m a.s.l. along 68 km from the southwest to the northeast of the Central Andes, in the departments of Ayacucho and Huancavelica in Peru. Precipitation follows a seasonal pattern, with 60–90% of total rainfall between December and March (Langstroth et al. 2013). For the evaluation sites the total annual precipitation values according to PISCO (Peruvian Interpolated data of SENAMHI’s Climatological and hydrological Observations), range from 480 mm to 817 mm, with site KP 164 + 500 reaching 1232 mm (Aybar et al. 2019). Average monthly precipitation in the dry season (June–August) ranges from 3.3 to 12.2 mm, and in the wet season (January–March) from 107 to 130 mm (Aybar et al. 2019). Mean air temperatures range from 9.7 °C to 19.7 °C, with minimum temperatures reaching values below 0 °C at night (Valencia et al. 2013). The study area overlaps with several rural Andean communities, where traditional husbandry of alpacas, lamas, and sheep (introduced) is common.

Figure 1. 

Location of bofedales and main ecosystems in the study area. The map inset shows the position of Peru within South America and the approximate study area, indicated by a green dot. The ecosystems shapefiles used are from Peru’s Ecosystem Map (MINAM 2019).

We surveyed plant communities of 31 bofedales along our study transect, located between 4,265 and 4,855 m a.s.l. According to previous studies using satellite images (PERU LNG, not published), bofedales were reported to have sizes between 1.25 and 43.98 ha, although we observed sites smaller than 1 ha in the field. We did the assessments annually during the austral dry season (June–July) from 2017 to 2019. In 2017, we set up randomly distributed 1 m × 1 m permanent plots in homogeneous patches of the dominant vegetation to characterize the plant communities in each bofedal. Per bofedal, we surveyed between two and seven plots (Table 1). The number was established considering the surface area of the bofedal, its heterogeneity (different plant communities), and the resources available for research. 378 plots were surveyed in the three years, 127 in 2017 and 2019, and 124 in 2018 (the missing plots could not be relocated).

We surveyed plant cover (per species) and ground cover with the point intercept method using a grid quadrat frame (Bonham 2013), considering 100 points (crossing points of the thread from the quadrat) per plot. The percentage cover for each species will equal the number of points in which the species was recorded. In subsequent years, we returned to each plot using coordinates and photographs (Suppl. material 1) and recorded the same plot area during each survey (Suppl. material 2; Linares-Palomino and Maldonado-Fonkén 2023).

We recorded cover estimates of several strata (ground cover) as useful proxies of degradation and potential habitat preferences of plant communities and species. We used the following categories: vegetation (vascular plants), moss, bare soil (or peat), dead vegetation, wildlife and livestock dung, rock, and water.

We used information on soil moisture and water table depth from the BMAP. We measured soil moisture (only in 2017) in 71 plots (2–3 records per plot) of ten plant communities (Table 2), with a ML3 ThetaProbe Soil Moisture Sensor from Delta-T Devices (UK), configured for organic soils. The water table monitoring wells were located at 11 sites (Table 1) and in six plant communities (Table 2). Using a peat sampler from Royal Ejelkamp (The Netherlands) we made a 50.8 mm diameter hole, where we inserted a PVC tube of the same size. Water table measurements were taken between July 2017 and December 2019, every one or two months in the first two years, and only in January and December of the last year. The number of sites, plots, soil moisture measurements, and monitoring wells per plant community are presented in Table 2.

We collated a list of species and morphospecies based on field collections and surveys done by the BMAP since 2009 (Valencia et al. 2013). We followed APG IV (The Angiosperm Phylogeny Group et al. 2016), and scientific names and authorship followed The World Flora Online (WFO 2024). Species were primarily identified in the field by a team of experienced bofedal botanists and ecologists (MM, HC). However, when plant material was fragmentary and/or lacked fertile structures, we collected and photographed samples and checked them against specialized literature (Tovar 1993; Gonzáles 2015; Sylvester et al. 2016) or referred them to specialists. We used the morphospecies concept on collections that were difficult to identify at the species level but otherwise had morphological characters that unequivocally differed from all the other material already identified in the area.

Since bofedal communities are associated with water-logged conditions, some species thrive in moist or saturated soils (hydrophytes and others). These were defined as moisture indicators (Suppl. material 3) based on our field observations and literature (Kahn et al. 1993; Tovar 1993; Gonzáles 2015; Meneses et al. 2015).

The bofedal plant communities we characterize here along a southern Peruvian east–west Andean transect are heterogeneous as reported in studies across the Andes from Colombia (Cleef 1981; Benavides and Vitt 2014), Ecuador (Suarez et al. 2022), Peru (Cooper et al. 2010; Maldonado-Fonkén 2014; Salvador et al. 2014; Maldonado-Fonkén 2018; Polk et al. 2019; Portal-Quicaña 2019), Bolivia (Ruthsatz 2012; Loza Herrera et al. 2015; Domic et al. 2021), Chile (Squeo et al. 2006) and Argentina (Ruthsatz et al. 2020; Izquierdo et al. 2020; Izquierdo et al. 2022).

The seven plant communities (group 1) we identified (Distichia muscoides, Rockhausenia pygmaea, Plantago rigida, Plantago tubulosa, Lachemilla diplophylla, Aciachne pulvinata and Juncus Stipulatus) were consistent in their structural and compositional characteristics and maintained differences between them during our three-year study. The remaining six potential communities or group 2 (Calamagrostis rigescens, Mixed community 1, Calamagrostis chrysantha, Distichia filamentosa, Lobelia oligophylla, Mixed community 2) require additional surveys to resolve their status as independent communities. We include them here as some have been described previously with similar structural or compositional characteristics, such as communities of Distichia filamentosa in Bolivia (Ruthsatz 2012) or communities with either Lobelia oligophylla or Calamagrostis tarmensis in northern Peru as co-dominant species (Cooper et al. 2010).

All the dominant or co-dominant species of the thirteen plant communities we describe here, have been previously reported as key components in bofedales throughout South America, as will be discussed below, although not necessarily by being the most abundant or frequent species in the community.

The Distichia muscoides hard cushion community has been reported throughout Peru (INAIGEM 2023), including in areas close to our study sites (Maldonado-Fonkén 2014, 2018; Portal-Quicaña 2019). This community is found in South America following the wide geographical distribution of its distinctive species from Colombia to northern Argentina (Ruthsatz 2012). Nevertheless, it is not the only dominant species in bofedales, as shown in our study and other works in Colombia (Cleef 1981; Benavides and Vitt 2014), Ecuador (Suarez et al. 2022), Peru (Cooper et al. 2010; Maldonado-Fonkén 2014; Salvador et al. 2014; Polk et al. 2019; Portal-Quicaña 2019), Bolivia (Ruthsatz 2012; Loza Herrera et al. 2015), Chile (Squeo et al. 2006) and Argentina (Ruthsatz et al. 2020; Izquierdo et al. 2022). This community is present in sites rarely affected by saline stress (electrical conductivity 19–713 μS cm^-1), droughts, or frost (Salvador et al. 2014). The number of species previously reported per site in Bolivia and Peru was between 16–39 (Ruthsatz 2012), while we reported 3–28 species. The lowest species richness was found in sites with almost exclusive dominance of D. muscoides, with pools retaining considerable water even in the dry season and usually with soil moisture over 80%. Since D. muscoides is an aquatic plant (Leon and Young 1996), a shallow water table (and consequently, high soil moisture) could limit the development of other species with less tolerance for saturated conditions. High water tables favor peat accumulation enabling increased carbon capture and storage compared to sites with lower water tables. D. muscoides is dominant under very stable hydrological conditions and with a water level close to the surface, not deeper than 50 cm (Oyague 2021). Our study confirmed this, as this community had the shallowest mean water table, only three monitoring wells had values that exceeded 50 cm of depth in September 2017 or 2018.

Rockhausenia pygmaea is distributed from Venezuela to Argentina (Salvador et al. 2014) and is considered a characteristic species of bofedales (Ruthsatz 2012; Ruthsatz et al. 2020). It is a peat-forming species (Benavides and Vitt 2014) that grows forming carpets (Salvador et al. 2014), low-firm cushions to hummocks (Aubert et al. 2014). It was identified as a potential dominant species in the Rockhausenia pygmaea–Pernettya prostrata assemblages in Ancash (Polk et al. 2019) and in the Plantago tubulosa–Oreobolus obtusangulus–Rockhausenia pygmaea assemblages in Cajamarca (Cooper et al. 2010), both in northern Peru. We recorded it as a dominant species and forming a recognizable community in 42% of the sites, being the second most common community after Distichia muscoides. Our records of soil moisture and water table suggest that it can thrive in drier conditions than D. muscoides.

Plantago rigida communities have been reported in Peru (Salvador et al. 2014), Ecuador (Suarez et al. 2022), and Colombia (Cleef 1981). According to Ruthsatz (2012), this species also grows at the edge of bofedales and in drier environments, and our results showed that it can thrive in drier conditions than D. muscoides. Nevertheless, Cleef (1981) reported it in wet depressions. In our study, it was the species with the highest abundance in its community and usually with fewer (less than five) companion species that can grow in the rare small openings between the leaves of this species or on the outside border of the cushions.

Plantago tubulosa is distributed along the Andes from Central America to Argentina (Cooper et al. 2010; Salvador et al. 2014) and is also considered a common species in bofedales (Ruthsatz et al. 2020). P. tubulosa communities have been reported from northern to southern Peru (Cooper et al. 2010; Salvador et al. 2014; Maldonado-Fonkén 2018), extending to Bolivia (Loza Herrera et al. 2015). However, it is most referred to as a co-dominant species in mixed communities with Distichia muscoides (Salvador et al. 2014) and Oreobolus obtusangulus (Polk et al. 2019), or in the Plantago tubulosa–Oreobolus obtusangulus–Rockhausenia pygmaea assemblage (Cooper et al. 2010). In our study, communities heavily dominated by P. tubulosa were found in 23% of the sites. This species also occurred as co-dominant with Calamagrostis vicunarum and Lachemilla pinnata in the Mixed community 1. Soil moisture was high and less variable than in the D. muscoides community; nevertheless, considering the lack of pools, we recommend additional studies about its water requirements.

Aciachne pulvinata was previously reported only as a companion species in bofedales in Peru (Cooper et al. 2010; Salvador et al. 2014) and Bolivia (Ruthsatz 2012). This species, associated with overgrazing (Salvador et al. 2014; Cochi Machaca et al. 2018), was present in eight of the 13 communities but was dominant only in one site with a low plant cover (75%) and the highest percentage of dead cushions (11%) recorded among the communities. Maldonado-Fonkén (2018) described a related community characterized by the sister taxon Aciachne acicularis in Ayacucho, where the percentage of litter and bare soil were at least 10% each, while the plant cover was 73%. We reported similar values in this study, despite the differences in survey methods. This community, together with P. rigida, had low water requirements. According to our observations, A. pulvinata increases its coverage in bofedales where the water level is decreasing.

Distichia filamentosa is distributed in Peru, Bolivia, and Chile (Ramirez 2011). This community has also been reported in those countries, growing at the upper growth limit of other cushion species (Squeo et al. 2006; Ruthsatz 2012; Loza Herrera et al. 2015). It has also been recorded in overgrazed areas in Bolivia (Cochi Machaca et al. 2018). We observed similar conditions in the only site where we identified this community: D. filamentosa was found at the extreme edge of the bofedal, next to a cryoturbated area, surrounded by D. muscoides cushions. Its water requirements seem to be high (soil moisture in dry season), but we recommend additional studies (water table measurements) in more sites. Lobelia oligophylla is distributed from Colombia to Bolivia and Chile (Cooper et al. 2010) and is considered a typical species of bofedales (Ruthsatz 2012). It was identified as a co-dominant species with the moss Drepanocladus longifolius in seasonally flooded areas in bofedales in Cajamarca (Cooper et al. 2010), and it is widely distributed in the peatlands of the Argentinean Puna (Izquierdo et al. 2020). In this study, we found it in a flat wet area but without any adjacent depressions or evidence of water courses that would indicate a potential to become a seasonally flooded area during the rainy season.

Juncus stipulatus, Calamagrostis rigescens, Calamagrostis chrysantha and Lachemilla diplophylla have been previously reported only as companion species in bofedales in Bolivia (Ruthsatz 2012) and Peru (Salvador et al. 2014). C. rigescens is also reported as a typical species of bofedales in Argentina (Ruthsatz et al. 2020) and is associated with overgrazing (Cochi Machaca et al. 2018). Species of the same genera, like Juncus arcticus or Calamagrostis tarmensis, have been previously described as co-dominant in bofedales communities in northern Peru (Cooper et al. 2010). In the case of rushes, they were found bordering lakes and ponds (Cooper et al. 2010). In our study, the Juncus stipulatus community was found in permanently waterlogged places but devoid of nearby pools. In contrast, the Lachemilla diplophylla community was usually found on wetter surfaces and moister areas, even recording the highest abundance of soil moisture indicators cover.

We encourage further study of the six preliminary communities (Group 2) to determine whether they result from local anthropogenic processes (e.g., overgrazing and draining) that increase the dominance of certain species or if more complex factors come into play (e.g., changes in water temperature and quality, regional climatic changes, etc.).

In most studies on bofedales vegetation, total species richness or mean richness per site is reported. These results are strongly influenced by the area assessed and sampling methods, which makes comparisons per area or sampling unit (e.g. square meter) difficult and shows an knowledge gap. Salvador et al. (2014) reported 56 species (vascular plants) in 24 sites, while Portal-Quicaña (2019) reported 85 species in one bofedal close to our study area. Our results (68 species in 27 sites) are similar to both studies. Other reports included 102 species in 36 sites (Cooper et al. 2010) and 119 species in 47 sites (Izquierdo et al. 2020). The number of species per 1 m² we found (1 to 15) is similar to what was reported in Argentina (1–13, Izquierdo et al. 2020). As reported in previous studies, Poaceae and Asteraceae were the families with the most species (Maldonado-Fonkén 2018). Nevertheless, studies in Argentina, northern and central Peru included Cyperaceae as a third important family (Cooper et al. 2010; Polk et al. 2019; Izquierdo et al. 2020), which is much less conspicuously represented in our study.

Water table measurements per plant community in bofedales are uncommon, and this is the first report of soil moisture values. Although mentioned, details of the water table are usually not provided in published studies (e.g. Cooper et al. 2010). According to the recent classification proposed by INAIGEM (2023), the bofedales where we installed monitoring wells are considered seasonal because the water table is deeper than 20 cm for several months of the year. This suggests that water availability in our sites is strongly influenced by rainfall seasonality. This applies to 13 of 31 sites and five communities (A. pulvinata, D. muscoides, Mixed community 2, P. rigida and R. pygmaea). However, rainfall seasonality is probably not the only water source of these bofedales. Hillslope groundwater flowing from lateral moraines, talus, colluvium, or bedrock aquifers can also be a source (Cooper et al. 2019), especially because streams, lakes, and glaciers are uncommon in the area. Further hydrological studies are required to clarify the main water source of these bofedales.

Even though bofedales can be hydrologically seasonal, the dominant plants are perennial. Some have deep roots (e.g., Distichia spp., Plantago spp.) and can withstand periods without much water. In addition, the high content of organic matter or peat facilitates water storage. Therefore, the plant cover usually does not change over time significantly. Species richness could be more sensitive. If water is unavailable (too deep) for longer periods than the species can withstand, permanent changes can occur in the plant communities (e.g., dominant species, plant cover, etc.). A La Niña event (November 2017 to March 2018; IGP 2023), usually associated with droughts in southern Peru, provided an opportunity to test this. Our results showed this was not the case within our sites, with stable values over time. Furthermore, we learned that assessing the communities in the dry season (when resources are limited for plants), can give better information of their condition over time.

The concept of bofedales necessarily includes a set of several distinctive plant communities that respond to microenvironmental site characteristics (Cooper et al. 2010; Ruthsatz 2012; Salvador et al. 2014; Polk et al. 2019). We show that the heterogeneity in plant communities occurs at local site level, and also at landscape and regional scales, with 84% of our studied sites having 2–3 plant communities. Recognizing this vegetational heterogeneity is important for conservation, ecosystem management and/or restoration activities since it allows the establishment of reasonable goals according to the diversity, structural and compositional characteristics of the sites. Given the scarcity of regional-level surveys in Peru, our results constitute the first steps towards identifying useful indicators of vegetation characteristics for biological baseline development and monitoring. These indicators are essential for setting achievable and site-specific restoration goals and distinguishing relevant contributions of ecosystem services such as carbon storage, water regulation or grazing. To achieve this, we identified three key gaps in bofedales knowledge in Peru that need to be addressed in the short to medium term. First, we need to characterize the floristic variation across latitudinal and longitudinal gradients. We believe that there are several local studies describing the floristic composition of bofedales in several regions in Peru that make it possible to attempt an initial comprehensive analysis of plot-based studies at the national scale. Second, we need to improve the accuracy of remote-based sensor mapping of bofedales. Although the currently available official map of Peru’s bofedales represents an important milestone, it inadequately describes the distribution and area of several bofedales in our study area. Some are misrepresented in terms of size or location, while others are not included at all. Third, the most complex challenge lies in understanding the various factors to which different bofedales plant communities respond, particularly those that define the structure and functioning of these unique ecosystems in a context of rapid land-use and climatic changes.

Yearly plot-based species cover data have been deposited at Figshare and are publicly available (https://doi.org/10.25573/data.24512719). All other data used and mentioned in the manuscript are provided as Suppl. materials 1–8.

Conceptualization: M.M.F. and R.L.P.; methodology, formal analysis, writing-original draft preparation: M.M.F. and H.C.; data curation, field assessment: M.M.F.; writing-review and editing: M.M.F., R.L.P., H.C. and B.V. All authors have read and agreed to the published version of the manuscript.

We thank Pablo Najarro, Marco Rivera, Andrea de la Cruz, Jaqueline Carhuapoma, Adrian Vera, Reyna Tacas, Filfredo Huamanyolli, Roque Misayme, Claudencio Galvez, Esteban Conislla S. and Luz Mabel Nuñez E. who assisted during field assessments. We thank Karim Ledesma, Dante Diaz and Cristiam Oriundo who facilitated logistics and access to the study sites. We thank Juan José Alegria and Nanette Vega for helping with species identification. Finally, we thank the editor and reviewers for their constructive comments and suggestions, which greatly improved the paper. This is contribution No. 66 of the CCS Latin American Biodiversity Programs of the Smithsonian’s National Zoo and Conservation Biology Institute.

• Mónica Maldonado-Fonkén (Corresponding author, mmaldonado@corbidi.org), ORCID: https://orcid.org/0000-0002-6735-5856
• Héctor Chuquillanqui (hchuquillanqui@gmail.com), ORCID: https://orcid.org/0000-0003-3181-128X
• Bruno Vildoso (bvildoso@huntloc.pe), ORCID: https://orcid.org/0000-0002-1075-3909
• Reynaldo Linares-Palomino (linaresr@si.edu), ORCID: https://orcid.org/0000-0002-7631-5549