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Phylogenetic Distance Among Beneficiary Species in a Cushion Plant Species Explains Interaction Outcome

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Phylogenetic Distance Among Beneficiary Species in a Cushion Plant Species Explains Interaction Outcome Zurich Open Repository and Archive University of Zurich Main Library Strickhofstrasse 39 CH-8057 Zurich www.zora.uzh.ch Year: 2015 Phylogenetic distance among beneficiary species in a cushion plant species explains interaction outcome Pistón, Nuria ; Armas, Cristina ; Schöb, Christian ; Macek, Petr ; Pugnaire, Francisco I Abstract: Determining which drivers lead to a specific species assemblage is a central issue in com- munity ecology. Although many processes are involved, plant–plant interactions are among the most important. The phylogenetic limiting similarity hypothesis states that closely related species tend to compete stronger than distantly related species, although evidence is inconclusive. We used ecological and phylogenetic data on alpine plant communities along an environmental severity gradient to assess the importance of phylogenetic relatedness in affecting the interaction between cushion plants and the whole community, and how these interactions may affect community assemblage and diversity. We first measured species richness and individual biomass of species growing within and outside the nurse cushion species, Arenaria tetraquetra. We then assembled the phylogenetic tree of species present in both com- munities and calculated the phylogenetic distance between the cushion species and its beneficiary species, as well as the phylogenetic community structure. We also estimated changes in species richness at the local level due to the presence of cushions. The effects of cushions on closely related species changed from negative to positive as environmental conditions became more severe, while the interaction with distantly related species did not change along the environmental gradient. Overall, we found an environmental context-dependence in patterns of phylogenetic similarity, as the interaction outcome between nurses and their close and distantly-related species showed an opposite pattern with environmental severity. DOI: https://doi.org/10.1111/oik.01979 Posted at the Zurich Open Repository and Archive, University of Zurich ZORA URL: https://doi.org/10.5167/uzh-117877 Journal Article Accepted Version Originally published at: Pistón, Nuria; Armas, Cristina; Schöb, Christian; Macek, Petr; Pugnaire, Francisco I (2015). Phyloge- netic distance among beneficiary species in a cushion plant species explains interaction outcome. Oikos, 124(10):1354-1359. DOI: https://doi.org/10.1111/oik.01979 1 Phylogenetic distance among beneficiary species in a cushion plant species explains 2 interaction outcome. 3 Nuria Pistón*, Cristina Armas1, Christian Schöb2, Petr Macek3, Francisco I. Pugnaire. 4 5 Pistón, N. ([email protected]) & Pugnaire, F.I.: Estación Experimental de Zonas 6 Áridas, Consejo Superior de Investigaciones Científicas, Carretera de Sacramento s/n, 7 E-04120 La Cañada de San Urbano, Almería, Spain. 8 1Armas, C.: Departamento de Biología, Universidad de La Serena, Casilla 554, La 9 Serena and Instituto de Ecología y Biodiversidad, Casilla 653, Santiago, Chile. 10 2Schöb, C.: Institute of Evolutionary Biology and Environmental Studies, University of 11 Zürich, Winterthurerstrasse 190, CH-8057, Zürich. 12 3Macek, P. current address: Faculty of Science, University of South Bohemia, 13 Branisovska 31, CZ-370 05, Ceske Budejovice. 14 15 Summary 16 Determining which drivers lead to a specific species assemblage is a central issue in 17 community ecology. Although many processes are involved, plant-plant interactions are 18 among the most important. The phylogenetic limiting similarity hypothesis states that 19 closely related species tend to compete stronger than distantly related species, although 20 evidence is inconclusive. We used ecological and phylogenetic data on alpine plant 21 communities along an environmental severity gradient to assess the importance of 22 phylogenetic relatedness in affecting the interaction between cushion plants and the 23 whole community, and how these interactions may affect community assemblage and 24 diversity. We first measured species richness and individual biomass of species growing 25 within and outside the nurse cushion species, Arenaria tetraquetra. We then assembled 26 the phylogenetic tree of species present in both communities and calculated the 27 phylogenetic distance between the cushion species and its beneficiary species, as well as 28 the phylogenetic community structure. We also estimated changes in species richness at 29 the local level due to the presence of cushions. The effects of cushions on closely 30 related species changed from negative to positive as environmental conditions became 31 more severe, while the interaction with distantly related species did not change along 32 the environmental gradient. Overall, we found an environmental context-dependence in 33 patterns of phylogenetic similarity, as the interaction outcome between nurses and their 34 close and distantly-related species showed an opposite pattern with environmental 35 severity. 36 37 Keywords: alpine habitat, competition, cushion plants, environmental filtering, 38 facilitation, phylogenetic community structure, phylogenetic limiting similarity, species 39 assemblages, stress gradient. 40 41 Introduction 42 Finding out which mechanisms interact at different spatial scales leading to a given 43 species assemblage is a central issue in community ecology. These mechanisms can be 44 separated relative to the spatial scale into external and internal filters, and involve 45 processes acting at regional scale (i.e., beyond single communities) as well as processes 46 within local communities (Ricklefs 2010, Violle et al. 2012). External filters include 47 factors such as climate, which sort species out from the regional pool and are important 48 determinants of plant community composition (Gaston 2000). Similarly, internal filters 49 such as microscale environmental heterogeneity (Fibich et al. 2013) or plant-plant 50 interactions (Armas et al. 2011) can affect community composition and structure. These 51 filters are not independent; for instance, the outcome of plant-plant interaction varies 52 depending on external filters such as climatic conditions (He et al. 2013). Internal filters 53 may interact with each other as well, e.g. when small-scale environmental heterogeneity 54 affects plant-plant interactions (Choler et al. 2001). Therefore, an approach unifying 55 different drivers, including external and internal filters, is needed to disentangle their 56 relative importance and impact on community assembly. 57 The integration of phylogenetic information in community analyses has proved 58 to be a powerful tool in understanding changes in species composition. Closely related 59 species tend to share similar trait values, hence may have similar requirements and 60 affect their microenvironment in similar ways; as a consequence, competition is 61 stronger between them than with distantly related species (Violle et al. 2011). This idea 62 was formalized as the phylogenetic limiting similarity hypothesis (PLSH; MacArthur 63 and Levins 1967) and assumes that ecological traits influencing species competition are 64 conserved along phylogenetic lineages (Blomberg et al. 2003, Wiens and Graham 65 2005). However, evidence supporting this hypothesis is inconclusive (Cahill et al. 2008, 66 Mayfield and Levine 2010). 67 The species pool under mild environmental conditions is larger and shows 68 wider range of ecological niche space than under severe conditions (Grime and Pierce 69 2012). Opposite, under harsh environmental conditions external filters (e.g., climate) 70 will severely restrict the species pool, favouring only the presence of species adapted to 71 such demanding conditions (Choler 2005). Assuming that species strategies are 72 phylogenetically conserved (Blomberg et al. 2003, but see Mayfield and Levine 2010), 73 the outcome of plant-plant interactions under mild environmental conditions could 74 render communities phylogenetically diverse, including closely related species. Hence, 75 at the local scale closely related species can show strong competition, following PLSH 76 predictions. By contrast, under harsh environmental conditions the species pool is 77 reduced and it could be expected that species will be more phylogenetically related 78 (Webb et al. 2002) as the environment selects for a given suite of traits. 79 Under harsh environmental conditions nurse plant species allow for the presence 80 of many other species in the community (Callaway 2007). In alpine environments, a 81 particular case of nurses are cushion plants, which usually ameliorate environmental 82 conditions and facilitate growth and survival of other species (beneficiaries) within 83 them (Badano and Cavieres 2006, Cavieres and Badano 2009). These nurse species can 84 create communities more phylogenetically diverse than communities in open habitats 85 (Butterfield et al. 2013). However, environmental severity restricts the species pool, 86 selecting for species sharing similar traits and more phylogenetically related (Soliveres 87 et al. 2012a). This leads to a paradox between competition intensity among 88 phylogenetically related species and environmental severity that needs more 89 exploration. 90 To test how plant-plant interactions and phylogenetic relatedness affect each 91 other and how they change along a severity gradient, we analysed the phylogenetic 92 relationships between cushions on other species in three sites along an elevation 93 gradient in the alpine belt of the Sierra Nevada Mountains,
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