Agriculture has played a pivotal role in shaping landscapes, soils and vegetation. Developing a better understanding of early farming practices can contribute to wider questions regarding the long-term impact of farming and its nature in comparison with present-day traditional agrosystems. In this study we determine stable carbon and nitrogen isotope values of barley grains from a series of present-day traditionally managed farming plots in Morocco, capturing a range of annual rainfall and farming practices. This allows a framework to be developed to refine current isotopic approaches used to infer manuring intensity and crop water status in (semi-)arid regions. This method has been applied to charred crop remains from two early farming sites in the eastern Mediterranean: Abu Hureyra and 'Ain Ghazal. In this way, our study enhances knowledge of agricultural practice in the past, adding to understanding of how people have shaped and adapted to their environment over thousands of years.

We integrate functional weed ecology with crop stable carbon and nitrogen isotope analysis to assess their combined potential for inferring arable land management practices in (semi-)arid regions from archaeobotanical assemblages. Weed and GIS survey of 60 cereal and pulse fields in Morocco are combined with crop sampling for stable isotope analysis to frame assessment of agricultural labour intensity in terms of manuring, irrigation, tillage and handweeding. Under low management intensity weed variation primarily reflects geographical differences, whereas under high management intensity fields in disparate regions have similar weed flora. Manured and irrigated oasis barley fields are clearly discriminated from less intensively manured rain-fed barley terraces in southern Morocco; when fields in northern and southern Morocco are considered together, climatic differences are superimposed on the agronomic intensity gradient. Barley δ 13 C and δ 15 N values clearly distinguish among the Moroccan regimes. An integrated approach combines crop isotope values with weed ecological discrimination of low-and high-intensity regimes across multiple studies (in southern Morocco and southern Europe). Analysis of archaeobotanical samples from EBA Tell Brak, Syria suggests that this early city was sustained through extensive (low-intensity, large-scale) cereal farming. ARTICLE HISTORY

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.

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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 one of seed size. Methods Using published values of comparative seed number as the ‘golden 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 CSR 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 CSR strategies. Moreover, some species differed in life history over their geographical range. Practically, comparative seed yield per m 2 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 ‘golden standard’ predictor used. Moreover, regressions estimating comparative seed yield m -2 achieved low precision. Conclusions For the first time, estimates of comparative seed yield and number for over 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 versus indeterminate flowering (‘bet-hedging’), C-S-R methodologies, phylogeny, comparative seed yield per m 2 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.

Background The carob tree (Ceratonia siliqua L.) is one of the most iconic tree species of the Mediterranean region, with valuable economic, ecological and cultural value. Carob has been exploited around the Mediterranean region since antiquity and has been regarded as an important component of natural habitats and traditional agroecosystems. Several studies have focused on its morphological, biochemical, and genetic diversity. However, less is known about the intraspecific variation of seed traits. In this regard, and as an overall objective, we intend to evaluate the amplitude and the expression of intraspecific variations of carob seed traits at different ecological scales ranging from individual trees to different geographical landscapes. In addition, we investigated how the climate along the study area affects the extent of carob seed variability. Using image analysis techniques, we measured seven traits related to the size and the shape of 1740 seeds collected from 18 populations of spontaneous C. siliqua distributed along a latitudinal transect in Morocco under different bioclimatic conditions. Results The morphometric analysis of carob seed showed the effectiveness of adopted approach to highlight the amount and the amplitude of intraspecific variation according to geographic and climatic factors. Seed trait analysis revealed high intraspecific variability, explained by differences between and among carob populations and geographic zones. Seed area, perimeter, length, and width showed the largest variability between geographic zones. However, circularity, aspect ratio, and seed roundness showed higher variability at the tree level. Finally, our results show that seed traits vary depending on altitude and climate condition. Conclusions Revealing the amount and the structure of intraspecific traits variability of carob seed provides interesting insights to understand the mechanisms underlying trees adaptation to various environmental and ecological conditions. Therefore, intraspecific variation of seed traits should be integrated into trait-based functional ecology to assess plant species responses to environmental changes.

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