�ۿ۴�ý

All applicants must be:

• US citizens or permanent residents;
• At least 18 years old by May 15 of the year of travel;
• Full time students at their home institution and have at least one more semester at their home institution after their research experience in Spain;
• Curious about science, learning about other cultures, and open to new ideas.

��

Additionally, applicants should have:

• >3.0 cumulative GPA;
• Completed at least 30 academic credits by the beginning of their trip to Spain;
• Taken introductory coursework emphasizing evolution, ecology, environmental science, and/or systematics; coursework in statistics is recommended but not required;��
• An openness to language learning, althougha minimum proficiency level is not required to apply. If selected, students should plan to enroll in a Spanish language course in the spring semester, particularly if they have little to no Spanish language proficiency.

��

Previous research experience is not required.

��

Each of the Spanish partner institutions offers a rich diversity of potential research experiences for US students. They all have extensive research facilities, a long history of gypsum research involving multiple scientists interested in varying aspects of gypsum ecology and evolution, and are located very near important gypsum outcrops in different parts of Spain. The three participating institutions are world leaders in the study of gypsum ecosystem ecology, with complementary expertise in floristics, conservation, ecological restoration, evolutionary and functional ecology, plant and biological soil crust community ecology, phylogeography, population genetics, plant-herbivore interactions, and ecophysiology of gypsum plants and lichens.��

is a public research university in Madrid that is located near the extensive gypsum exposures of central Spain. The URJC Biodiversity and Conservation group was founded in 1996 and is one of the most productive research groups in ecology in Spain (ranked #2 of Spanish institutions, 2021 Shanghai Global Ranking of Academic Subjects). Composed of 107 members (44 with teaching and research responsibilities), the program also hosts 25 PhD and 15 postdoctoral students, providing a dynamic environment where US students can interact with biologists at all professional stages.��

Founded in 1942, the is a state-run research institute located in northeast Spain, near the extensive gypsum exposures of the Ebro Valley. It is one of the oldest research institutes of the Spanish National Research Council (CSIC) and the only one entirely devoted to ecological studies in mountainous areas. Research undertaken at IPE seeks to identify the processes and mechanisms underlying the biodiversity, functioning and structure of mountain ecosystems, from semi-arid lowlands (often covered by gypsum soils in Spain) to alpine summits. With over 56 highly multidisciplinary research staff members (ranging from economists and geologists to ecologists, molecular biologists, climatologists, and geographers), the institution has extensive experience in the study of ecological processes in gypsum soils. This vibrant academic community includes some of the most highly cited researchers in ecology and climate change.��

is a public university in the southeastern city of ���������� and is located adjacent to the Sorbas and Tabernas gypsum exposures, which contain the highest degree of gypsum endemism in Europe. The RNM344 Conservation Biology research group at UA has a long history of research into plants growing on unusual soils, including gypsum, as well as dolomites (soils rich in magnesium carbonate). Composed of nineteen scientists, the group has interests in conservation, restoration, ecophysiology, and systematics of plants growing on unique soil types, particularly gypsum and dolomite.

��

IPE hosts numerous postdocs and graduate students and has also been involved in undergraduate mentorship, including a recent �ۿ۴�ý undergraduate alumnus and Fulbright scholar. IPE/IRES mentors Drs. Palacio (who also coordinates the project), Pueyo, and Tejero have made extensive contributions to nutrient ecophysiology, community ecology, and evolutionary ecology in gypsum plants (e.g., Palacio et al. 2007; Pueyo and Alados 2007; Pueyo et al. 2007; Pueyo et al. 2008; Palacio et al. 2014a, b; Escudero et al. 2015; Palacio et al. 2017).

• Dr. Sara Palacio specializes in the mechanisms that plants use to survive in extreme environments and how these traits have influenced plant evolution. Her research ranges from high mountain peaks to the driest deserts on the planet, with a special emphasis on gypsum soils.��

• Dr. Yolanda Pueyo studies the functioning and dynamics of terrestrial ecosystems, with a particular emphasis on grazed systems in semi-arid, subalpine, and gypsum environments. Her research focuses on the interplay between plants, soils, and human activity in determining ecosystem function, with the ultimate goal of providing recommendations for sustainable grazing management and tools for ecological restoration of degraded lands.

• Dr. Pablo Tejero is the herbarium curator at the Instituto Pirenaico de Ecologia in Jaca, Spain, the most important alpine herbarium in the country. His primary research interest is plant evolution, adaptation, and systematics, with particular foci on mountain and gypsum floras.

As at IPE, the IRES mentors at URJC have extensive mentorship experience, having collectively supervised more than 20 doctoral theses and more than 35 Master's and undergraduate theses. IRES mentors Drs. García-Cervigón, Luzuriaga, Matesanz, Prieto, and Sánchez are URJC faculty members who are able to provide a wide variety of research projects ranging from community ecology and the evolution of phenotypic plasticity to phylogeography and population genetics (e.g., Escudero et al. 2007, Matesanz et al. 2017, Luzuriaga et al. 2018, Blanco-Sánchez et al. 2021, García-Cervigón et al. 2021, Sánchez et al. 2021, López-Rubio et al. 2022).

• Dr. Maria Prieto��Álvaro is a lichenologist interested in understanding the phylogenetic, taxonomic, and functional diversity of lichen communities in relation to environmental conditions, including soil factors. She is also interested in understanding how the environment influences the functional attributes of lichens.

• Dr. Ana García-Cervigón is a plant ecologist interested in understanding the response of natural systems to global warming and human impacts. She uses dendrochronology and quantitative wood anatomy with Mediterranean and tropical dry forests, and with shrub communities from high mountains to semiarid gypsum environments.

• Dr. Silvia Matesanz is an evolutionary ecologist interested in micro‐evolutionary processes that occur in plant populations of gypsum endemics, and how phenotypic variation is shaped by genetic and environmental factors.

• Dr.��Arantzazu L. Luzuriaga is community ecologist interested in species assembly processes. Her research spans annual and perennial plant communities in arid and semiarid systems, particularly on restricted soils such as gypsiferous substrates. She is interested in soil seed bank dynamics, secondary succession, and the implementation of functional and phylogenetic diversity as tools to disentangle species assembly mechanisms.

• Dr. Ana Sánchez Álvarez is a community ecologist interested in how drought impacts species diversity, particularly in gypsum ecosystems.

IRES mentors Drs. Mota, Salmerón, Martinez, Mendoza, and Merlo are broadly interested in gypsum evolution, ecology, and conservation and have led many important recent studies into gypsum plant phylogeography, phylogeny, and floristics (Martínez-Nieto et al. 2013; Mota et al. 2017; Salmerón-Sánchez et al. 2014, 2017; Pérez-García et al. 2017, 2018). Dr. Mota and colleagues have also conducted key studies of conservation reserve design, including the ecological restoration of gypsum quarries and conservation of the unique plant communities that develop on gypsum and other atypical soils (Cerrillo et al. 2002; Mota et al. 2003, 2004; Mendoza-Fernández et al. 2010; Pérez-García et al. 2021; Bobo-Pinilla et al. 2021; Salmerón-Sánchez et al. 2021).

• Dr. Antonio Mendoza-Fernández is a botanist, whose research interests include distribution patterns, conservation, restoration and management of the endangered flora in Andalusia (southern Spain) and in the Mediterranean Basin.

• Dr. Encarna Merlo is a plant physiologist who investigates the stress responses of plants living on special soils such as gypsum, dolomite and serpentine

• Dr. Juan Mota studies the flora of unique soil types, such as gypsum, dolomite, and serpentine, with research projects ranging from microbiomes to ecophysiology to entire vascular plant communities. Given that these specialized soils are derived from rocks that are often mined, much of his research focuses on applying our knowledge of how these systems function to the restoration of gypsum mines and the uses of endemic species for phytoremediation.

• Dr. Esteban Salmerón Sánchez is a plant biologist interested in the study of endemic species associated with special substrates such as gypsum, dolomite or serpentine. His studies are focused on conservation biology; ecophysiology, phylogeography, molecular phylogenetics and conservation biology.

��

Below are example projects US undergraduates will participate in at Spanish partner institutions, with other potential projects possible, depending on student interests. Projects are grouped by the four major research objectives for the GYP-NEXTGEN project, with abbreviations given for Spanish institutions (IPE: Instituto Pirenaico de Ecología; UA: Universidad de ���������í��; URJC: Universidad Rey Juan Carlos) and research mentors [AL: Arantzazu Luzuriaga (URJC); AG: Ana García Cervigón (URJC); AM: Antonio Mendoza (UA); AS: Ana Sánchez (URJC); EM: Encarnación Merlo (UA); ES: Esteban Salmerón (UA); FM: Fabián Martínez (UA); JM: Juan Mota (UA); MP: María Prieto (URJC); PT: Pablo Tejero (IPE); SM: Silvia Matesanz (URJC); SP: Sara Palacio (IPE); YP: Yolanda Pueyo (IPE)].

Research objective 1: Investigate the mechanistic links between soil chemistry, plant elemental composition, defense chemistry, and phylogeny (IPE, UA, URJC). One of the most striking responses of plants to gypsum soils is the enhanced ability of gypsum endemic plants to accumulate high S and Ca concentrations in their tissues. Students will select from a diversity of projects investigating the ecological and evolutionary role of this mechanism on gypsum plants.

• Leveraging a global dataset of plant and soil chemical composition from gypsum ecosystems to compare trends worldwide, probing the links between gypsum adaptation, plant ionic composition, and phylogeny, and investigating patterns among plants from atypical substrates (gypsum, dolomite, serpentine). PIs: JM, SP, PT
• Analyzing S metabolism in gypsophile and gypsovag plants by evaluating differences in gene expression under different levels of S availability. PIs: SP, PT
• Understanding the ecological role of glucosinolates in gypsum Brassicaceae in response to environmental pressures (e.g., drought, grazing). PIs: SP, YP, AS

Research objective 2: Investigate the processes leading to community assembly in gypsum plant communities and the roles of intraspecific trait variation, phenotypic plasticity, genetic variation, and competition in gypsum habitats (URJC). Students will focus on the biotic and abiotic processes that define plant community assembly in gypsum ecosystems, selecting from a suite of potential projects and PIs.��

• Competitive response and effects of annual plant species growing in semiarid gypsum systems of central Spain. PI: AL
• Unraveling the role of phenotypic plasticity, population differentiation and adaptive trait variation under ecologically realistic multivariate environments. PI: SM, AS
• Disentangling rapid evolution on quantitative traits related to drought and fitness in a dominant gypsum shrub through a resurrection experiment. PI: SM
• Understanding phenotypic variation and plasticity of wood anatomy in gypsum chamaephytes in response to drought. PI: AGC, AS

Research objective 3: Investigate phylogenetic relationships among key gypsum endemic taxa in the Mediterranean to understand trait evolution and genetic diversity (UA, IPE). Students will work to understand the evolutionary processes that have allowed some plants to thrive on gypsum, using a mixture of molecular and bioinformatics tools.

• Resolving phylogenetic relationships of gypsum endemic species of Teucrium (Lamiaceae) and Gypsophila (Caryophyllaceae) to explore the evolution of plant elemental composition and gypsum affinity. PI: JM, ES
• Investigating hybridization among gypsophile and non-gypsophile taxa in Helianthemum. PI: EM, ES
• Exploring the species identities and phylogenetic relationships of cryptogamic organisms from gypsum and non-gypsum biological soil crusts. PI: MP����

Research objective 4: Investigate succession and restoration processes in gypsum landscapes (UA). Spain has a long history of gypsum mining and is a major exporter of gypsum. Gypsum ecosystems are now considered a priority habitat for protection by the EU. Students will work to evaluate gypsum ecosystem restoration as part of sustainable development in a changing world.

• Evaluating plant community responses as a function of time since restoration. PI: JM, FM

• Characterizing biological soil crusts across gypsum landscape successional series. PI: EM
• Investigating primary succession in gypsum quarries. PI: EM, AM

International travel will be booked by the US university representatives. Participants will travel via economy class from their representative US locations (Cleveland for �ۿ۴�ý and OC students; El Paso for NMSU students) to Madrid, Spain, where all participants will participate in a multi-day orientation session. Those conducting their research at URJC will move into their long-term accommodations, and those at IPE and UA will stay in local hotels. Following the orientation conference, IPE and UA participants will travel by train (IPE) or air (UA) to their research locations. Once in Jaca and ���������í��, participants will move into their long-term accommodations. All arrangements will be made by the US and Spanish partners for the participants.

��