Seeing the fields through the weeds: introducing the WeedEco R package for comparing past and present arable farming systems using functional weed ecology.

The functional ecology of arable weeds provides a way of comparing present-day and past farming regimes. This paper presents the R package WeedEco, an open-source resource which allows users to compare their archaeobotanical dataset against three previously published arable weed models to understand fertility, disturbance or a combination of both. The package provides functions for data organisation, classification and visualisation, allowing users to enter raw archaeobotanical data, obtain trait values from the functional trait dataset, conduct discriminant analysis and plot the results against the relevant present-day model. Using data from the early medieval site of Stafford in the UK, the paper provides a detailed example of the use of the package, demonstrating its different functions, as well as how the results can be interpreted. Supplementary Information: The online version contains supplementary material available at 10.1007/s00334-023-00964-8.

A new functional ecological model reveals the nature of early plant management in southwest Asia.

The protracted domestication model posits that wild cereals in southwest Asia were cultivated over millennia before the appearance of domesticated cereals in the archaeological record. These 'pre-domestication cultivation' activities are widely understood as entailing annual cycles of soil tillage and sowing and are expected to select for domestic traits such as non-shattering ears. However, the reconstruction of these practices is mostly based on indirect evidence and speculation, raising the question of whether pre-domestication cultivation created arable environments that would select for domestic traits. We developed a novel functional ecological model that distinguishes arable fields from wild cereal habitats in the Levant using plant functional traits related to mechanical soil disturbance. Our results show that exploitation practices at key pre-domestication cultivation sites maintained soil disturbance conditions similar to untilled wild cereal habitats. This implies that pre-domestication cultivation did not create arable environments through regular tillage but entailed low-input exploitation practices oriented on the ecological strategies of the competitive large-seeded grasses themselves.

Seed size, number and strategies in annual plants: a comparative functional analysis and synthesis.

BACKGROUND AND AIMS: Plants depend fundamentally on establishment from seed. However, protocols in trait-based ecology currently estimate seed size but not seed number. This can be rectified. For annuals, seed number should simply be a positive function of vegetative biomass and a negative function of seed size. METHODS: Using published values of comparative seed number as the 'gold standard' and a large functional database, comparative seed yield and number per plant and per m2 were predicted by multiple regression. Subsequently, ecological variation in each was explored for English and Spanish habitats, newly calculated C-S-R strategies and changed abundance in the British flora. KEY RESULTS: As predicted, comparative seed mass yield per plant was consistently a positive function of plant size and competitive ability, and largely independent of seed size. Regressions estimating comparative seed number included, additionally, seed size as a negative function. Relationships differed numerically between regions, habitats and C-S-R strategies. Moreover, some species differed in life history over their geographical range. Comparative seed yield per m2 was positively correlated with FAO crop yield, and increasing British annuals produced numerous seeds. Nevertheless, predicted values must be viewed as comparative rather than absolute: they varied according to the 'gold standard' predictor used. Moreover, regressions estimating comparative seed yield per m2 achieved low precision. CONCLUSIONS: For the first time, estimates of comparative seed yield and number for >800 annuals and their predictor equations have been produced and the ecological importance of these regenerative traits has been illustrated. 'Regenerative trait-based ecology' remains in its infancy, with work needed on determinate vs. indeterminate flowering ('bet-hedging'), C-S-R methodologies, phylogeny, comparative seed yield per m2 and changing life history. Nevertheless, this has been a positive start and readers are invited to use estimates for >800 annuals, in the Supplementary data, to help advance 'regenerative trait-based ecology' to the next level.

Trait-based plant ecology a flawed tool in climate studies? The leaf traits of wild olive that pattern with climate are not those routinely measured.

Climate-related studies have generally focussed upon physiologically well-defined 'mechanistic' traits rather than 'functional' ones relating indirectly to resource capture. Nevertheless, field responses to climate are likely to typically include both 'mechanistic' specialization to climatic extremes and 'functional' strategies that optimize resource acquisition during less climatically-severe periods. Here, this hypothesis was tested. Seventeen traits (six 'functional', six 'mechanistic' and five 'intermediate') were measured from 19 populations of oleaster (wild olive) along a climatic gradient in Morocco. Principal components analysis of the trait dataset identified size and the 'worldwide leaf economics spectrum' as PCA axes 1 and 2. However, contrary to our prediction, these axes, and commonly-measured 'functional' traits, were little correlated with climate. Instead, PCA 3, perhaps relating to water-use and succulence, together stomatal density, specific leaf water content and leaf shape, patterned with altitude, aridity, rainfall and temperature. We concluded that, at least for slow-growing species, such as oleaster, 'mechanistic' traits are key to identifying mechanisms of climatic restriction. Meaningful collaboration between 'mechanistic' and 'functional' disciplines provides the best way of improving our understanding of the global impacts of climate change on species distribution and performance.

Trade-offs between seed and leaf size (seed-phytomer-leaf theory): functional glue linking regenerative with life history strategies and taxonomy with ecology?

Background and Aims: While the 'worldwide leaf economics spectrum' (Wright IJ, Reich PB, Westoby M, et al. 2004. The worldwide leaf economics spectrum. Nature : 821-827) defines mineral nutrient relationships in plants, no unifying functional consensus links size attributes. Here, the focus is upon leaf size, a much-studied plant trait that scales positively with habitat quality and components of plant size. The objective is to show that this wide range of relationships is explicable in terms of a seed-phytomer-leaf (SPL) theoretical model defining leaf size in terms of trade-offs involving the size, growth rate and number of the building blocks (phytomers) of which the young shoot is constructed. Methods: Functional data for 2400+ species and English and Spanish vegetation surveys were used to explore interrelationships between leaf area, leaf width, canopy height, seed mass and leaf dry matter content (LDMC). Key Results: Leaf area was a consistent function of canopy height, LDMC and seed mass. Additionally, size traits are partially uncoupled. First, broad laminas help confer competitive exclusion while morphologically large leaves can, through dissection, be functionally small. Secondly, leaf size scales positively with plant size but many of the largest-leaved species are of medium height with basally supported leaves. Thirdly, photosynthetic stems may represent a functionally viable alternative to 'small seeds + large leaves' in disturbed, fertile habitats and 'large seeds + small leaves' in infertile ones. Conclusions: Although key elements defining the juvenile growth phase remain unmeasured, our results broadly support SPL theory in that phytometer and leaf size are a product of the size of the initial shoot meristem (≅ seed mass) and the duration and quality of juvenile growth. These allometrically constrained traits combine to confer ecological specialization on individual species. Equally, they appear conservatively expressed within major taxa. Thus, 'evolutionary canalization' sensu Stebbins (Stebbins GL. 1974. Flowering plants: evolution above the species level . Cambridge, MA: Belknap Press) is perhaps associated with both seed and leaf development, and major taxa appear routinely specialized with respect to ecologically important size-related traits.

An evolutionary perspective on leaf economics: phylogenetics of leaf mass per area in vascular plants.

In plant leaves, resource use follows a trade-off between rapid resource capture and conservative storage. This "worldwide leaf economics spectrum" consists of a suite of intercorrelated leaf traits, among which leaf mass per area, LMA, is one of the most fundamental as it indicates the cost of leaf construction and light-interception borne by plants. We conducted a broad-scale analysis of the evolutionary history of LMA across a large dataset of 5401 vascular plant species. The phylogenetic signal in LMA displayed low but significant conservatism, that is, leaf economics tended to be more similar among close relatives than expected by chance alone. Models of trait evolution indicated that LMA evolved under weak stabilizing selection. Moreover, results suggest that different optimal phenotypes evolved among large clades within which extremes tended to be selected against. Conservatism in LMA was strongly related to growth form, as were selection intensity and phenotypic evolutionary rates: woody plants showed higher conservatism in relation to stronger stabilizing selection and lower evolutionary rates compared to herbaceous taxa. The evolutionary history of LMA thus paints different evolutionary trajectories of vascular plant species across clades, revealing the coordination of leaf trait evolution with growth forms in response to varying selection regimes.

Long-term resistance to simulated climate change in an infertile grassland.

Climate shifts over this century are widely expected to alter the structure and functioning of temperate plant communities. However, long-term climate experiments in natural vegetation are rare and largely confined to systems with the capacity for rapid compositional change. In unproductive, grazed grassland at Buxton in northern England (U.K.), one of the longest running experimental manipulations of temperature and rainfall reveals vegetation highly resistant to climate shifts maintained over 13 yr. Here we document this resistance in the form of: (i) constancy in the relative abundance of growth forms and maintained dominance by long-lived, slow-growing grasses, sedges, and small forbs; (ii) immediate but minor shifts in the abundance of several species that have remained stable over the course of the experiment; (iii) no change in productivity in response to climate treatments with the exception of reduction from chronic summer drought; and (iv) only minor species losses in response to drought and winter heating. Overall, compositional changes induced by 13-yr exposure to climate regime change were less than short-term fluctuations in species abundances driven by interannual climate fluctuations. The lack of progressive compositional change, coupled with the long-term historical persistence of unproductive grasslands in northern England, suggests the community at Buxton possesses a stabilizing capacity that leads to long-term persistence of dominant species. Unproductive ecosystems provide a refuge for many threatened plants and animals and perform a diversity of ecosystem services. Our results support the view that changing land use and overexploitation rather than climate change per se constitute the primary threats to these fragile ecosystems.

Specific leaf area and dry matter content estimate thickness in laminar leaves.

BACKGROUND AND AIMS: Leaf thickness plays an important role in leaf and plant functioning, and relates to a species' strategy of resource acquisition and use. As such, it has been widely used for screening purposes in crop science and community ecology. However, since its measurement is not straightforward, a number of estimates have been proposed. Here, the validity of the (SLA x LDMC)(-1) product is tested to estimate leaf thickness, where SLA is the specific leaf area (leaf area/dry mass) and LDMC is the leaf dry matter content (leaf dry mass/fresh mass). SLA and LDMC are two leaf traits that are both more easily measurable and often reported in the literature. METHODS: The relationship between leaf thickness (LT) and (SLA x LDMC)(-1) was tested in two analyses of covariance using 11 datasets (three original and eight published) for a total number of 1039 data points, corresponding to a wide range of growth forms growing in contrasted environments in four continents. KEY RESULTS AND CONCLUSIONS: The overall slope and intercept of the relationship were not significantly different from one and zero, respectively, and the residual standard error was 0.11. Only two of the eight datasets displayed a significant difference in the intercepts, and the only significant difference among the most represented growth forms was for trees. LT can therefore be estimated by (SLA x LDMC)(-1), allowing leaf thickness to be derived from easily and widely measured leaf traits.