Aim Gymnosperms are often described as a marginal and threatened group, members of which tend to be out-competed by angiosperms and which therefore preferentially persist at higher latitudes and elevations. The aim of our synthesis was to test these statements by investigating the global latitudinal and elevational distribution of gymnosperms, as well as their conservation status, using all extant gymnosperm groups (cycads, gnetophytes, ginkgophytes and conifers).Location Worldwide.Methods We developed a database of 1014 species of gymnosperms containing latitudinal and elevational distribution data, as well as their global conservation status, as described in the literature. The 1014 species comprised 305 cycads, 101 gnetophytes, the only living representative of ginkgophytes, and 607 conifers. Generalized additive models, frequency histograms, kernel density estimations and distribution maps based on Takhtajan's floristic regions were used.Results Although the diversity of gymnosperms decreases at equatorial latitudes, approximately 50% of the extant species occur primarily between the tropics. More than 43% of gymnosperms can occur at very low elevations (≤ 200 m a.s.l.). Gymnosperms, considering all species together as well as their main taxonomic groups separately, do not exhibit a latitudinal diversity gradient as commonly observed for many other taxa. Gymnosperms, and especially conifers, are on average less threatened at higher and equatorial latitudes.Main conclusions Gymnosperms display an unusual latitudinal diversity gradient, which we suggest cannot fully be accounted for by angiosperm dominance and competitive superiority. We hypothesize that other factors explain their present distribution, such as the development of centres of endemism in several regions and the adaptation of certain taxa to cold and arid climates.

Aim To elucidate the phylogeographical patterns in three Cenozoic relict species: Zelkova sicula, Z. abelicea and Z. carpinifolia (Ulmaceae).Location Sicily, Crete and Transcaucasia.Methods Two chloroplast loci (trnH-psbA and trnL) and the nuclear ribosomal markers ITS1 and ITS2 were sequenced for 154 samples collected from 14 populations of Z. abelicea, 16 populations of Z. carpinifolia, and the two known populations of Z. sicula. We obtained georeferenced data, calculated median joining networks and carried out diversity analyses. A few ex situ samples collected from botanical gardens, of the East Asian species Zelkova serrata, Z. schneideriana and Z. sinica, were also analysed for comparative purposes.Results High levels of variability were found in the chloroplast markers within Z. carpinifolia (15 haplotypes) and Z. abelicea (33 haplotypes), in association with strong phylogeographical structure. Conversely, Z. sicula was characterized by low diversity, with each population exhibiting a single haplotype. Lower variability was found for ITS1 and ITS2 within Z. carpinifolia and Z. abelicea (13 and 7 ribotypes per species, respectively), with Z. carpinifolia showing a high proportion of populations with no intragenomic polymorphism. In the triploid and clonal Z. sicula, all individuals displayed intragenomic polymorphism and seven ribotypes were identified.Main conclusions The chloroplast diversity of Z. abelicea and Z. carpinifolia suggests a very ancient history of diversification and structuring, with footprints of past expansions and more recent bottlenecks. Zelkova sicula has had a history of severe isolation and is likely to be of hybrid origin.

Aim To establish a complete database of Juglandaceae at a spatiotemporal scale and develop a phylogeographic framework with which to elucidate the distributional patterns, diversity patterns, origins, evolution, and conservation priority regions of this family. Location Worldwide. Taxon Walnut family (Juglandaceae). Methods Data on the distribution of all the extant and fossil species of Juglandaceae were collected, followed by analyses of its latitudinal distribution, elevational distribution, and species and generic diversity. Furthermore, based on all genera and 87% of the species, we reconstructed phylogenetic relationships, estimated divergence times, calculated phylogenetic diversity and inferred ancestral distributions. Results Extant Juglandaceae (10 genera and 60 species) are mainly distributed in eastern Asia and North America (principally between 20 and 40°N). Tropical Juglandaceae mainly inhabit mountainous areas higher than 1,000 m, especially in the New World. Southwest China and northern Vietnam are characterized by high species, generic and phylogenetic diversity. The United States of America has only high species diversity. The area of origin of Juglandaceae was North America and Europe in the early Eocene, and its widespread dispersal mainly occurred between 13 and 26 Ma. Main conclusions The members of Juglandaceae inhabit areas with temperate climatic conditions. The diversification centre has shifted intercontinentally from North America and Europe to Southwest China and northern Vietnam, which are identified as conservation priority regions. The high‐latitude cooling during the Oligocene followed by a long‐term stable warmer climate in the early and middle Miocene drove the southward translocation of the family.

Relict species provide a unique opportunity to understand past and recent biogeographical and evolutionary processes. Zelkova abelicea (Ulmaceae), which is endemic to the island of Crete (Greece), is one of the most prominent Tertiary relict trees of the Mediterranean region. We collected distribution, threat and population structure data by reviewing literature and herbaria and through field surveys at 14 study plots throughout the range of the species. The present distribution of Z. abelicea is extremely fragmented. Although the total estimated number of individuals is relatively high, the populations are dominated by dwarf, severely browsed, non-flowering individuals. The population structure is asymmetric. At most, 5% of a plot's trees are large and fruit-bearing. The asymmetric structure is particularly pronounced in isolated and small populations. Based on its limited geographical range, the fragmented spatial pattern, and data on distribution and population structure, our study confirms that Z. abelicea is a threatened species (IUCN category Endangered). Our research aim is to promote the development of new approaches for the improvement of conservation strategies for Tertiary relict trees characterized by major local disjunctions.

Field studies and conceptual work on hybridization-mediated extinction risk in climate relicts are extremely rare. Nuphar pumila (Nymphaeaceae) is one of the most emblematic climate relicts in Europe with few isolated populations in the Alpine arc. The extent of introgression with related lowland and generalist species Nuphar lutea has never been studied using molecular methods. All biogeographical regions where N. pumila naturally occurs in the neighbourhood of the Alpine arc were sampled and studied using nuclear microsatellite markers. Furthermore, we used forward-in-time simulations and Approximate Bayesian Computation to check whether an introgression scenario fits with the observed admixture patterns and estimated the demographic parameters associated with this process. Our study confirms ongoing hybridization between N. pumila and N. lutea and validates it by the use of population models. More than 40 % of investigated N. pumila individuals were admixed and hybrids were found in over 60 % of studied populations. The introgression is bidirectional and is most likely a result of very recent gene flow. Our work provides strong evidence for rapid extinction risk and demographic swamping between specialized climatic relicts and closely related generalists. The remaining pure populations of N. pumila are rare in the Alpine arc and deserve high conservation priority.