A leaf-height-seed (LHS) plant ecology strategy scheme

“…a common language for comparing species and vegetation types worldwide, seems to have dropped below the horizon, with the notable exception of a sustained campaign from the Unit for Comparative Plant Ecology at Sheffield (e.g. Grime, 1974, 1977, 1979; Grime et al., 1988, 1997), of which more later.”

“High SLA indicates high light capture by leaf tissue mass, whereas low SLA reflects mass allocation that increases longevity, robustness, and defense (Westoby, 1998).”

“and function in plants, has generated a great deal of new interest in traits (Reich, 2014; Westoby, 1998).”

“Within the adult palm community, three functional traits representing variation in light acquisition strategies (Chazdon, 1991; Westoby, 1998; Svenning, 2000) were associated with varied regeneration stages: more mature forests were characterized by species with decreased (1) maximum stem height,…”

“Another recent study examining intraspecific trait variation along a steep aridity gradient found no evidence of ecotypic differentiation with respect to Leaf‐Height‐Seed (LHS) traits (Westoby, 1998; Westoby et al., 2002), suggesting that such traits may be relatively unimportant for climate adaptation compared to phe…”

“To calculate these functional diversity metrics, we used four main traits representing key axes of ecological strategies among plant species: height, SLA, seed mass and annual clonal spread (Hejda & de Bello, 2013; Westoby, 1998).”

“On the453 K side are species with dense wood and slow growth (Westoby, 1998; Poorter et al., 2005).454 Low foliar nitrogen and low SLA can align with low foliar Rubisco content, low photosynthetic455 capacity, and, thus, low maintenance respiration in low light (Reich et al., 1998; Poorter,…”

“The importance of the tree height variable may be related to higher tolerance of taller trees to drought (Westoby 1998).”

“…nitrogen concentration (%), specific leaf area (SLA; cm 2 g −1 ) and leaf tissue density (LTD; cm 3 g −1 ) are key traits that reflect physiological constraints associated with the leaf economic spectrum and other resource use strategies (Pierce et al., 2007 ; Poorter & Evans, 1998; Westoby, 1998).”

“Also, larger-seeded species tend to mobilize their metabolic resources over a longer period (Westoby 1998; Kidson and Westoby 2000; Leishman et al. 2000).”

“Seed weight and size have been identified as plant traits strongly influencing the dispersal, establishment, survival and growth of seedlings (Harper et al. 1970; Harper 1977; Westoby 1998; Weiher et al. 1999; Moles & Westoby 2004), particularly at early seedling stages (Leishman et al.”

“Average height at maturity was chosen, because a tree's height influences its ability to compete for light, resist wind, and support leaves and chemical transport (Westoby 1998).”

“We analyzed lignin by the van Soest (1963) procedure and total soluble phenolic concentration by the Folin–Ciocalteu method using 50 % methanol as extract solution and tannic acid as standard (Waterman and Mole 1994).”

“All of these traits are related to plants’ ecological strategies (for details see Supplementary Table S3), furthermore H and LA are also informative about competitive ability and plant persistence (Westoby, 1998; Díaz et al., 2016).”

“Plant height is a surrogate for plant body size and is a key functional trait relating to the ecological strategies of species (Westoby 1998).”

“We focused on five key traits, which are considered to capture the plant strategy schemes: specific leaf area, wood density, maximum height, seed mass and shade tolerance (Westoby 1998).”

“The traits utilized are chosen because they represent a common currency that can be measured and compared across systems (Westoby, 1998; Westoby et al., 2002; Reich, 2014) and because they are believed to represent fundamental life history trade-offs that should be related to performance (e.g.…”

“distance) and CH at maturity (competition for light with neighbouring plants) as effect traits, according to leaf–height–seed strategy scheme proposed by Westoby [39].”

“LMA and nutrient concentrations are important indicators of plant resource capture and usage strategies, and are fundamental for the growth and regeneration of species ( Westoby, 1998; Wilson et al., 1999; Vendramini et al., 2002; Hallik et al., 2009; He et al., 2009; Poorter et al., 2009).”

“…to be fundamental components of the ecological strategies of plants, relating to how plants acquire and use resources (Givnish, 1987; Reich et al., 1997; Westoby, 1998; Diaz et al., 2004), as well as playing an important role in influencing the properties of ecosystems (Lavorel and Garnier, 2002).”

“The three traits were selected to test the accuracy of the Leaf-height-seed (LHS)-scheme (Westoby, 1998) and to analyze whether a few core traits identified as crucial for plant survival can be used as response traits in changing grasslands.”

“The statistical parameters (i.e., estimates, MS, F, and p) were showed. advantage to achieve light resource when competing with neighbours, which confers a higher species biomass (Valladares & Niinemets, 2008; Westoby, 1998).”

“Tall plant heights are advantageous for dispersing seed and intercepting light (Gaudet and Keddy, 1988; Westoby, 1998; Westoby et al., 2002; Muller-Landau et al., 2008; Moles et al., 2009), although construction and maintenance costs are incurred along with higher risks of breaking (Niklas, 1993;…”

“These are commonly described by the leaf-height-seed (LHS) strategy scheme [80,81].”

“Together, these results suggest that invasiveness represents a general ecological strategy that can be assessed quantitatively and along which multiple species may be aligned (i.e. Pianka 1970; Grime 1977; Westoby 1998).”

“Reproductive height is a proxy of the competitive ability for light (Westoby 1998).”

“Plant height and Specific Leaf Area (SLA, the ratio of leaf surface to leaf dry mass) were both expected to respond to intensity of disturbance, with disturbed environments being characterized by shorter plants with higher SLA, associated with faster resource use strategies sensu Westoby (1998).”

“…function (Oksanen et al. 2016) for passive projection of environmental factors: DIFN, litter mass, disturbance indices, and CWMs of specific leaf area (SLA), canopy height, and seed mass—functional traits describing plant life strategies, according to LHS concept (leaf-height-seed; Westoby 1998).”

“The negative relationship between seed mass and surface area, either SLA or SRL, suggests that light-seeded populations may be better suited for aboveground and belowground resource acquisition in nutrient-rich environments (Westoby, 1998; Poorter and Jong, 1999).”

“We focused on three key traits that capture important resource investment tradeoffs and affect plant competitive abilities: SLA, maximum height and seed weight (Westoby, 1998).”

“The CSR strategy scheme provides a system for classifying herbaceous plants as having strategies adapted to intense competition (C), abiotic stress (S) and frequent disturbance (R) (Grime 1974; McIntyre et al. 1995; Lavorel et al. 1997; Westoby 1998).”

“…leaf, and whole-plant traits); we chose these traits according to their presumed or demonstrated relations with their growth strategies and ecological requirements (Westoby, 1998; Wahl and Ryser, 2000; Roumet et al., 2006; Poorter and Markesteijn, 2008; Mommer et al., 2011; Fort et al., 2015).”

“2003), has resulted in a number of strategy schemes being developed (e.g. Grime 1977; Gitay and Noble 1997; Westoby 1998; Westoby et al. 2002) and, as a result, was considered as a valid starting point for this project.”

“The full procedures for specific leaf area (SLA) and leaf dry matter content (LDMC) determination followed those of Westoby (1998) and Garnier et al. (2001).”

“…for instance, by modelling LARF of dominant and rare species separately, and d) the consideration of different plant ecological strategies in the functionaltrait-centred approach, preferably considering important axes of trait variation (Westoby 1998, Reich et al. 2003, Westoby and Wright 2006).”

“Consistently with LHS and LES schemes, we expect individuals occurring on exposed habitats of outcrops (when compared with individuals of the same species in sheltered habitats in surrounding woodlands) should: H1 – exhibit functional traits indicative of more resource-conservative strategies (e.g. lower leaf area, sp…”

“(e) Key parts of the CSR strategy concept (Grime 1979; Pierce et al. 2014) and the leaf-heightseed concept (LHS, Westoby 1998) are seed traits such as seed mass and volume, where seed mass is thought to be an indicator for seedling survival.”

“…2000; Poorter et al., 2005; Westoby, 1998; Westoby et al., 2002), wood density (Chave et al., 2009; Enquist et al., 1999) and seed mass (Rees, 1996; Westoby, 1998) typically explain the greatest amount of variance in tree performance in tropical forests when compared to leaf traits like SLA…”

“A key question in community ecology has been the extent to which these fundamental tradeoff axes or trait spectra are coordinated (Díaz et al., 2016; Westoby, 1998; Westoby et al., 2002).”

“Moreover, if functional traits are explicitly incorporated into assembly models, we can infer the characteristics determining species-specific responses (e.g. resource acquisition strategies, fecundity, growth rates; Diaz et al., 2004; Westoby, 1998).”

“Reproductive traits are expected to form independent strategy axes (Loehle 2000; Sutherland et al. 2000; Westoby 1998; Westoby and Wright 2006; Westoby et al. 2002; but see Deng et al. 2008).”

“SLA is correlated with other plant traits such as photosynthetic leaf N content, leaf life-span and relative growth rate, representing, therefore, a trade-off between the leaf carbon acquisition rate and the longevity of plant tissues (Westoby 1998).”

“Although often measured as a major trait accounting for above- ground competition (Westoby, 1998), we also considered maximum height as the closest proxy for grazing effects (see Díaz, Noy- Meir, & Cabido, 2001) and as an indicator of above- ground biomass (Catchpole & Wheeler, 1992).”

“These include increased height (Westoby 1998; Feng and van Kleunen 2016) and leaf area (Daehler 2003; Liu et al. 2015), lower seed mass (Thuiller et al. 2006; Schmidt and Drake 2011), leaf dry mass (Fynn et al. 2009; Matzek 2011) and leaf thickness (Niinemets 2001; Funk and Throop 2010), and the…”

“We opted to use maximum vegetative height as our functional trait of interest be‐ cause it is known to be an important proxy for resource partitioning and competitive ability in plants (Cornwell et al., 2014; Weiher et al., 1999; Westoby, 1998).”

“We focused on specific leaf area (SLA; cm2/g), adult tree height (H; m), oven-dried seed mass (SM; mg) and wood density (WD; g/cm3).”

“…traits, our results therefore show that biotic and abiotic drivers can independently alter successional trajectories by shifting community composition, and that community membership depends on specific traits related to dispersal and resource acquisition and allocation strategies (Westoby, 1998).”

“Here, we summarize three key dimensions of functional niches based on plant functional traits: leaf economy, mechanical support, and reproductive phenology (Table 1), which have been frequently discussed in previous studies (Westoby, 1998; Diaz et al., 2016; Messier et al., 2017).”

“…and life-cycle. ntroduction Biomass, growth rate and the suite of traits associated, are cenral to models in plant ecology because they are key determinants of lant performance, including reproduction, survival, and competiive ability (Tilman, 1988; Westoby, 1998; Grime, 2001; Pino et al., 002).”

“Previous work has shown that the traits included in this study are related to different strategies of plant dispersal, establishment and persistence (Westoby 1998; Weiher et al. 1999; Westoby et al. 2002; Laughlin 2014).”

“The plant functional traits studied most are those described by Westoby ( 1998 ): specific leaf area (i.e. the ratio of leaf area to leaf dry mass), plant height and seed mass, which describe three axes of plant variation related to their re - source management, competitive ability and regenerative…”

“…has progressively enriched ecology: Grime (1974) positioned species within a triangular space of competition, stress tolerance, and ruderality; Westoby (1998) formalized orthogonal axes of leaf investment, size, and regenerative capacity; and the leaf economics spectrum (LES; Wright et al .,…”

“…associated to the second axis of our PCA but had an intermediate position in the first PC plan and indicated that the seed dimension axis defined by Westoby (1998), as an expression of the potential of dispersal and successful establishment of a species43–45, was marginally correlated to plant and…”

“…species had consistently smaller seeds 508 than locally-persistent species, which is thought to increase dispersal distance (Greene and 509 Johnson 1993, Westoby 1998), and may also decrease performance upon arrival due to their 510 smaller maternal subsidies (Moles and Westoby 2006).”

“The leaf- height-seed scheme proposed by Westoby ( 1998 ) is very useful to describe key aspects of plant ecological strategies in response to resource availability and disturbances.”

“We included cardinal traits representing main components of plant economic spectrum (Westoby 1998; Díaz et al. 2016): two traits representing the leaf economic spectrum: specific leaf area (SLA, cm 2 g −1 ) and leaf dry matter content (LDMC, g g −1 ), and two traits representing plant size: plant…”