Tropical mountains are hot spots of biodiversity and endemism, but the evolutionary origins of their unique biotas are poorly understood. In varying degrees, local and regional extinction, long-distance colonization, and local recruitment may all contribute to the exceptional character of these communities. Also, it is debated whether mountain endemics mostly originate from local lowland taxa, or from lineages that reach the mountain by long-range dispersal from cool localities elsewhere. Here we investigate the evolutionary routes to endemism by sampling an entire tropical mountain biota on the 4,095-metre-high Mount Kinabalu in Sabah, East Malaysia. We discover that most of its unique biodiversity is younger than the mountain itself (6 million years), and comprises a mix of immigrant pre-adapted lineages and descendants from local lowland ancestors, although substantial shifts from lower to higher vegetation zones in this latter group were rare. These insights could improve forecasts of the likelihood of extinction and 'evolutionary rescue' in montane biodiversity hot spots under climate change scenarios.

Background and Aims The breadfruit genus (Artocarpus, Moraceae) includes valuable underutilized fruit tree crops with a centre of diversity in Southeast Asia. It belongs to the monophyletic tribe Artocarpeae, whose only other members include two small neotropical genera. This study aimed to reconstruct the phylogeny, estimate divergence dates and infer ancestral ranges of Artocarpeae, especially Artocarpus, to better understand spatial and temporal evolutionary relationships and dispersal patterns in a geologically complex region.Methods To investigate the phylogeny and biogeography of Artocarpeae, this study used Bayesian and maximum likelihood approaches to analyze DNA sequences from six plastid and two nuclear regions from 75% of Artocarpus species, both neotropical Artocarpeae genera, and members of all other Moraceae tribes. Six fossil-based calibrations within the Moraceae family were used to infer divergence times. Ancestral areas and estimated dispersal events were also inferred.Key Results Artocarpeae, Artocarpus and four monophyletic Artocarpus subgenera were well supported. A late Cretaceous origin of the Artocarpeae tribe in the Americas is inferred, followed by Eocene radiation of Artocarpus in Asia, with the greatest diversification occurring during the Miocene. Borneo is reconstructed as the ancestral range of Artocarpus, with dozens of independent in situ diversification events inferred there, as well as dispersal events to other regions of Southeast Asia. Dispersal pathways of Artocarpus and its ancestors are proposed.Conclusions Borneo was central in the diversification of the genus Artocarpus and probably served as the centre from which species dispersed and diversified in several directions. The greatest amount of diversification is inferred to have occurred during the Miocene, when sea levels fluctuated and land connections frequently existed between Borneo, mainland Asia, Sumatra and Java. Many species found in these areas have extant overlapping ranges, suggesting that sympatric speciation may have occurred. By contrast, Artocarpus diversity east of Borneo (where many of the islands have no historical connections to the landmasses of the Sunda and Sahul shelves) is unique and probably the product of over water long-distance dispersal events and subsequent diversification in allopatry. This work represents the most comprehensive Artocarpus phylogeny and biogeography study to date and supports Borneo as an evolutionary biodiversity hotspot.

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We present a 517-gene phylogenetic framework for the breadfruit genus Artocarpus (ca. 70 spp., Moraceae), making use of silica-dried leaves from recent fieldwork and herbarium specimens (some up to 106 years old) to achieve 96% taxon sampling. We explore issues relating to assembly, paralogous loci, partitions, and analysis method to reconstruct a phylogeny that is robust to variation in data and available tools. Although codon partitioning did not result in any substantial topological differences, the inclusion of flanking noncoding sequence in analyses significantly increased the resolution of gene trees. We also found that increasing the size of data sets increased convergence between analysis methods but did not reduce gene-tree conflict. We optimized the HybPiper targeted-enrichment sequence assembly pipeline for short sequences derived from degraded DNA extracted from museum specimens. Although the subgenera of Artocarpus were monophyletic, revision is required at finer scales, particularly with respect to widespread species. We expect our results to provide a basis for further studies in Artocarpus and provide guidelines for future analyses of data sets based on target enrichment data, particularly those using sequences from both fresh and museum material, counseling careful attention to the potential of off-target sequences to improve resolution. [Artocarpus; Moraceae; noncoding sequences; phylogenomics; target enrichment.]

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Assessment of the progress of the Aichi Biodiversity Targets set by the Convention on Biological Diversity (CBD) and the safeguarding of ecosystems from the perverse negative impacts caused by Reducing Emissions from Deforestation and Forest Degradation Plus (REDD+) requires the development of spatiotemporally robust and sensitive indicators of biodiversity and ecosystem health. Recently, it has been proposed that tree-community composition based on count-plot surveys could serve as a robust, sensitive, and cost-effective indicator for forest intactness in Bornean logged-over rain forests. In this study, we developed an algorithm to map tree-community composition across the entire landscape based on Landsat imagery. We targeted six forest management units (FMUs), each of which ranged from 50,000 to 100,000 ha in area, covering a broad geographic range spanning the most area of Borneo. Approximately fifty 20 m-radius circular plots were established in each FMU, and the differences in tree-community composition at a genus level among plots were examined for trees with diameter at breast height ≥10 cm using an ordination with non-metric multidimensional scaling (nMDS). Subsequently, we developed a linear regression model based on Landsat metrics (e.g., reflectance value, vegetation indices and textures) to explain the nMDS axis-1 scores of the plots, and extrapolated the model to the landscape to establish a tree-community composition map in each FMU. The adjusted R 2 values based on a cross-validation approach between the predicted and observed nMDS axis-1 scores indicated a close correlation, ranging from 0.54 to 0.69. Histograms of the frequency distributions of extrapolated nMDS axis-1 scores were derived from each map and used to quantitatively diagnose the forest intactness of the FMUs. Our study indicated that tree-community composition, which was reported as a robust indicator of forest intactness, could be mapped at a landscape level to quantitatively assess the spatial patterns of intactness in Bornean rain forests. Our approach can be used for large-scale assessments of tree diversity and forest intactness to monitor both the progress of Aichi Biodiversity Targets and the effectiveness of REDD+ biodiversity safeguards in production forests in the tropics.