Recruitment facilitation in expanding forests of Mediterranean juniper is sex-biased

Despite noticeable concern about the deforestation rate worldwide, the forest surface in Europe has considerably expanded over the past centuries as a consequence of the rural exodus and abandonment of agrarian practices. Tree recruitment associated with forest regrowth is a multi-stage process influenced by several biotic and abiotic factors. Yet, it is uncertain whether their influence on recruitment patterns and dynamics varies along a gradient of forest expansion. Similarly, for dioceious species, the influence of tree sex in recruitment is not entirely understood. Here, we aim to elucidate what drives Spanish juniper recruitment in expanding forests. Specifically, we hypothesized that facilitation by conspecifics and heterospecific woody species would occur at the expanding front, where environmental conditions are harsher and that recruitment would be preferably associated to female trees because of the likelihood of mature cones produced by them germinating in the nearby area. The study was conducted in Mediterranean forests of Juniperus thurifera in central Spain. A total of 17 plots were delimited along a gradient of forest expansion including three stages: i) old forests, ii) an intermediate zone and iii) novel forests at the expanding front. Within each plot all J. thurifera individuals (saplings and adults) were mapped. We also recorded bio-volumetric characteristics and tree sex for all adult trees and estimated the percentage of cover of heterospecific woody species within the area of influence of each adult individual. We analysed the spatial pattern of J. thurifera individuals for each stand (plot). Using a novel spatial approach, we evaluated how conspecific (female and male tree sizes) and heterospecific (woody cover) vegetation influenced sapling density along a forest expansion gradient. We also studied the effects of the stage of the forest expansion gradient and the sex of adult trees on the spatial association between adults and saplings. Our results showed that sapling recruitment was negatively influenced by conspecific adult size along the whole gradient, while the effect of heterospecific woody vegetation was always positive. Conspecific facilitation of recruitment in J. thurifera forests occurred at their expanding front where saplings were associated to male adult trees. Despite having been overlooked in conservation policies, recently colonised areas in extreme environments are key targets to implement management measures aimed at achieving forest restoration, which aligns with the Aichi targets and the biodiversity policies of the European Union.This work was made possible by the International Laboratory on Global Change LINCGlobal (www.lincg.uc-csic.es) and funding from the grants SPONFOREST (BiodivERsA3-2015-58, PCIN-2016-055, financed by the Spanish Research Agency (AEI) and the Spanish Ministry of Economy, Industry and Competitiveness (MINECO)), COMEDIAS (MINECO, CGL2017-83170-R) and REMEDINAL TE (Ref. TE-CM. S2018/EMT-4338, 2019-2023-Comunidad de Madrid). Coauthor CCB was supported by a postdoctoral fellowship from the Ramon Areces Foundation.Peer reviewe