Systematics and phylogeography of the Mediterranean Helichysum pendulum complex (Compositae) inferred from nuclear and chloroplast DNA and morphometrics analyses

Multiple factors related to complex geomorphological and climatic history, in addition to biological factors such as hybridization, hinder the definition of some Mediterranean plant groups. The existence of controversial taxonomic treatments, the possible hybridization events involved, and its unknown evolutionary history, make the Helichrysum pendulum complex of H. sect. Stoechadina an ideal model to understand general processes of Mediterranean plant systematics and evolution. The mosaic range of the complex, which is distributed over several islands and continental areas in the western-central Mediterranean Basin, provides an opportunity to investigate how past connections and disconnections between landmasses may have determined the current geographic distribution of genetic variation in this area. The cpDNA region rpl32-trnL intergenic spacer and the nrDNA region ETS were sequenced for 1-8 individuals from each of the 44 populations sampled, covering all taxa and the whole geographic range of the complex. These individuals were analysed together with a broad sampling of the remaining members of sect. Stoechadina. In addition, detailed multivariate analyses of morphological characters were performed for the whole section and for the H. pendulum complex. Considering together distinctive genetic and morphological traits, our species concept is presented and discussed in a context of integrative taxonomy, and five species are recognized within the complex: H. errerae, H. melitense, H. pendulum, H. saxatile and H. valentinum. The first three species are recognizable by qualitative and quantitative morphological traits, and are genetically distinguishable from the rest as shown by the molecular markers analysed. The two last species are reported here to have a putative ancient hybrid origin and are also genetically distinguishable from the rest but morphologically recognisable only by quantitative characters. Phylogenetic relationships shown by nuclear and chloroplast markers, and an intermediate morphology between the two putative parental taxa, point to H. pendulum and H. italicum as the putative parental taxa for H. saxatile, and H. pendulum and H. stoechas as putative parental taxa for H. valentinum. In a discriminant analysis of the five species, 97.8% of all individuals were classified correctly. The high level of haplotype and ribotype diversity observed in North Africa indicates that this region is either the area of origin of the complex or a secondary contact zone. Our results suggest that the complex colonized several islands and migrated through the Gibraltar and Sicilian Straits during phases of low sea level, favoured by local dispersal events that promoted its gradual range expansion. The occurrence of the complex in the Balearic Islands, which have remained isolated even during low sea level phases, could be explained by stochastic long-distance dispersal events

Comparative phylogeography of chloroplast and nuclear DNA markers reveals ancient and present hybridization in the Mediterranean Helichrysum pendulum complex (Compositae)

The geological and climatic history of the Mediterranean basin over the last 6 million years has been determinant in shaping current geographic distribution of genetic variation in organisms. Phylogeographical approaches are considered one of most useful analysis for unraveling the evolutionary history of species. The Helichrysum pendulum complex is a group of three closely related plant species distributed in several islands and isolated continental localities of the Western-Central Mediterranean basin, providing an ideal case of study to analyze the processes involved in modelling its current genetic structure. Two cpDNA region rpl32-trnL intergenic spacer and the nrDNA region ETS were sequenced for 1-8 individuals from each of the 44 total populations sampled, covering the whole geographic range of the complex. Our results suggest that the complex originated in northern Africa and colonized several islands and continental areas of the northern Basin through the Gibraltar and Sicilian straits during phases of low sea level, favored by long distance dispersal events. While ETS data suggest a model of isolation by distance and show a main genetic barrier between populations of Western and Central Mediterranean areas, the rpl32-trnL reveals the existence of two divergent and not geographically structured haplotype groups within the complex. Ancient hybridization events among lineages of sect. Stoechadina are suggested as the most plausible cause for the haplotypes pattern observed, while several evidences of current hybridization between H. pendulum and several species of sect. Stoechadina are also detected in ETS data

New conservation viewpoints when plants are viewed at one level higher. Integration of phylogeographic structure, niche modeling and genetic diversity in conservation planning of W Mediterranean larkspurs

Protection and management of closely related endangered species and subspecies at a very narrow regional scale is the origin of multiple dysfunctional conservation decisions. These include artificially increased IUCN risk assessment categories and derived consequences: poor effectiveness in allocating public and private funds or repeat of unnecessary actions/facilities. Data provided by the revisited study of a group of W Mediterranean larkspurs (Delphinium ser. Fissa), including new data on demography, niche modeling, genetic diversity and phylogeography, contributed to a new and wider analysis of causes of threat. Although current IUCN Red List regulations did not allow for assessments at levels higher than a specific rank, scientific information suggests that in some cases this could be a better approach for sound scientifically-based biodiversity conservation planning and action

The Korean Baekdudaegan Mountains: A Glacial Refugium and a Biodiversity Hotspot That Needs to Be Conserved

The Baekdudaegan (BDDG; Figure 1) is a mountain range relatively unknown outside Korea. From recent times, however, the BDDG is known outside Korea because it shelters the small county of Pyeongchang, the venue of the 2018 Winter Olympic Games.This research was supported by Korea Research Foundation grants; KRF-2013R1A1A2063524 to MYC and NRF-2011-0017236, NRF-2013R1A1A3010892, and NRF-2017R1A2B4012215 to MGC and was carried out as part of the Infrastructure for the Conservation and Restoration of Rare and Endemic Plants in Korea National Arboretum that supported to MGC from 2015 to 2018.Peer reviewe

Effects of climate change on the potential distribution of the threatened relict Dipentodon sinicus of subtropical forests in East Asia: Recommendations for management and conservation

Dipentodon sinicus Dunn. (Dipentodonaceae) is a rare and threatened relict plant species usually found co-dominating with other relict plants in subtropical forest patches in highly fragmented habitats of southwestern China, northern Vietnam and northeastern Myanmar of East Asia. To date, its management and conservation strategies in the light of climate change have not been explored. We evaluated effects of climate change on the distribution of climatically suitable areas of D. sinicus as found prevailing during the last glacial maximum (LGM), the mid-Holocene and the present time, and predicted the distribution of climatically suitable habitats in 2070 throughout East Asia. The results as derived from ecological niche modeling (ENM) show the current distribution to be limited to the prehistoric (the mid-Holocene and LGM) refugia, and to indicate decreasing probability of presence and a reducing range of distribution for 2070. In addition, the suitable areas predicted with high probability (0.5–1) only account for on average 9.8% of the total area of potential habitats (threshold‒1) among the models for the year 2070, thereby indicating that D. sinicus is highly vulnerable. Under all the future scenarios for the year 2070, 69–74.2% of potential habitats in China would be outside protected areas. We assess and propose priorities for protected areas, and provide suggestions for conservation management strategies.This study received financial support from Science and Technology Department of Yunnan University, China (2019YNU002), the Ministry of Science and Technology of China (2015FY210200-15), Ajuts a Grups de Recerca Consolidats” (grants nos. 2014-SGR514-GREB and 2017-SGR1116) from the Generalitat de Catalunya (Spain), Applied Basic Research Foundation of Yunnan Province, China (Grant No. 2019FB058), the Environment Research and Technology Development Fund (JPMEERF15S11407) of the Environmental Restoration and Conservation Agency of Japan, and the Kakenhi Grant Number 15H02833.Highlights Abstract Keywords 1. Introduction 2. Material and methods 2.1. Species 2.2. Occurrence data and ecological niche modeling 3. Results 3.1. Model performance and present potential distribution 3.2. Projected distribution during the mid-Holocene (ca. 6000 yr BP) and LGM (ca. 21,000 yr BP) 3.3. Projected distribution under future climate (2070) 4. Discussion 4.1. Effects of climate change on spatial distribution patterns of D. sinicus 5. Recommendations for future conservation efforts and management Declaration of competing interest Acknowledgements Appendix A. Supplementary data Reference

The recent neophyte Opuntia aurantiaca Lindl.: distribution and potential invasion in Spain

Trabajo presentado en el XVI Optima Meeting (Organization for the Phyto-Taxonomic Investigation of the Mediterranean Area), celebrado en Atenas (Grecia), del 2 al 5 de octubre de 2019The Cactaceae, and especially its most emblematic genus, Opuntia, have become one of the groups of plants with greater invasion potential in Spain. In fact, this area is considered one of the three global hotspots of invasion of cacti, together with South Africa and Australia. In our country, one of the most recently detected species is Opuntia aurantiaca Lindl, a small cactus with an enormous capacity of dispersion by means of cladodes and (sterile) fruits, which are easily detached and adhere to animals, clothes, and footwear. Probably native to the Southern Cone (Argentina and Uruguay), it behaves as a very aggressive invader in Australia and South Africa. In Europe, it only occurs on the Mediterranean coast of the Iberian Peninsula (Catalonia and Valencian Community). In this study, the geographic range of the species is updated at the peninsular level. Firstly detected at the beginning of the last decade in Navajas (Castellón), it has been subsequently observed in other places of the province of Castellón, but also in Valencia, Tarragona and Barcelona. With all occurrence data, its potential distribution is estimated using the algorithm of maximum entropy implemented in the program MaxEnt. Distribution models are built for the current climatic conditions as well as for different scenarios of global warming. Due to its possible invasive nature, it is necessary to eradicate the existing populations at this initial stage. Legislation should consider the prohibition of trade and transport of this plant to avoid new introductions

El gènere "Kalanchoe" (Crassulaceae) a Catalunya : situació i distribució potencial del tàxon invasor "K. ×houghtoni"

Cinc tàxons del gènere Kalanchoe (K. daigremontiana, K. tubiflora, K. fedtschenkoi, K. sexangularis i K. ×houghtonii) han estat citats de diverses localitats de Catalunya. Tanmateix, recentment hem pogut comprovar que les citacions a Catalunya de K. daigremontiana que s'han pogut revisar corresponen en realitat a l'híbrid K. ×houghtonii, d'una capacitat invasora molt superior a la de les espècies parentals. A partir d'una revisió de la literatura actual i de noves citacions aportades en aquest treball, s'ha actualitzat la distribució a Catalunya dels tàxons del gènere presents al territori i s'ha avaluat la distribució potencial de K. ×houghtonii utilitzant tècniques de modelització de nínxol ecològic. També aportem una clau per identificar les espècies presents o que s'han citat a Catalunya.Up to five taxa of Kalanchoe (K. daigremontiana, K. tubiflora, K. fedtschenkoi, K. sexangularis and K. ×houghtonii) have been reported from several localities of Catalonia. However, we have recently noted that records of K. daigremontiana from Catalonia that have been revised were misidentifications and actually belong to the hybrid K. ×houghtonii, which its invasive potential is higher than those of the parental species. Based on a review of the current literature but also on new citations that are provided in the present work, we update the distribution of the taxa occurring in Catalonia, and we estimate the potential distribution of K. ×houghtonii using an ecological niche modelling approach. Finally, an identification key of Kalanchoe taxa occurring or cited in Catalonia is provided.Cinco taxones del género Kalanchoe (K. daigremontiana, K. tubiflora, K. fedtschenkoi, K. sexangularis y K. ×houghtonii) han sido citados de diversas localidades de Cataluña. Sin embargo, recientemente hemos podido comprobar que muchas de las citas en Cataluña de K. daigremontiana que se han podido revisar corresponden en realidad al híbrido K. ×houghtonii, con una capacidad invasora muy superior a la de las especies parentales. A partir de una revisión de la literatura actual y de nuevas citas aportadas en el presente trabajo, se ha actualizado la distribución en Cataluña de los taxones del género presentes en el territorio y se ha evaluado la distribución potencial de K. ×houghtonii utilizando técnicas de modelización de nicho ecológico. También aportamos una clave para identificar las especies presentes o que se han citado en Cataluña

Phylogeography of the endangered orchids Cypripedium japonicum and Cypripedium formosanum in East Asia: Deep divergence at infra- and interspecific levels

To date, little is known about the past evolutionary trajectories of rare and endangered orchids native to mainland China, Japan, and Korea (the CJK region). In this study, we focus on two endangered orchids, Cypripedium japonicum (present in the three countries) and C. formosanum (endemic to Taiwan), to understand the divergence/speciation models that would have been operating in this group, including genetic diversity, geographic structure, and colonization pathways across the region. Using a combination of five cpDNA regions, we reconstructed phylogenetic trees and investigated the genetic diversity/structure of 20 populations. Ecological niche modeling was used to gain insight into the paleodistribution and dispersal corridors at the Last Glacial Maximum and to survey climatic niche differences. Populations from mainland China + Korea, Japan, and Taiwan formed three distinct monophyletic lineages and were placed into separate genetic clusters, agreeing with geographic barriers and species boundaries. Populations of C. japonicum in mainland China harbored the highest diversity, suggesting the presence of multiple glacial refugia. The Korean populations would have originated from either western/central or eastern China, probably using a dispersal corridor across the East China Sea shelf. The divergence of C. formosanum is proposed under an allopatric speciation model, also highly influenced by a climate niche shift. In the context of previous studies, a deep divergence in cpDNA sequences between Chinese + Korean and Japanese populations of C. japonicum may be taken as an example of the speciation events of the CJK flora since the late Neogene that have led to its current species richness.This study was supported by the Biodiversity Survey, Observation and Assessment Program of the Ministry of Ecology and Environment of China to HZT and by Basic Science Program through the National Research Foundation of Korea (NRF-2017R1A2B4012215) to MGC, and funded by the Ministry of Science and ICT of the Republic of Korea (NRF-2020R1I1A3074635) to MYC.INTRODUCTION MATERIALS AND METHODS Study species Population sampling DNA extraction cpDNA-PCR optimum primer selection cpDNA sequence alignment and assembly Haplotype distribution, phylogenetic analyses, and genetic diversity Genetic differentiation and structure Mismatch distribution analysis, neutrality detection, and demographic history ENM and population connectivity Niche comparisons in E-space RESULTS Haplotype distribution and phylogeny Genetic diversity Genetic differentiation and structure Mismatch distribution analysis, neutrality detection, and demographic history ENM and population connectivity Niche comparisons in E-space DISCUSSION Deep genetic and climatic divergence of Cypripedium sect. Flabellinervia in the CJK region: taxonomic considerations Haplotype and nucleotide diversity in Cypripedium sect. Flabellinervia: inference of glacial refugia and demographic history Origin of Korean populations of Cypripedium japonicum Origin of Cypripedium formosanum CONCLUSIONS AUTHOR CONTRIBUTIONS ACKNOWLEDGEMENTS Appendix 1: The cpDNA sequence information of Cypripedium sect. Flabellinervia deposited in the GenBank databas

Generic boundaries in subtribe Saussureinae (Compositae: Cardueae): Insights from Hyb-Seq data

The subtribe Saussureinae is a highly speciose group with more than 600 species distributed in the Northern Hemisphere and is particularly species-rich at the high mountains of central and eastern Asia. Saussurea and Jurinea are the two main genera described within the subtribe. However, up to 15 satellite genera are recognized in some recent taxonomic treatments with an analytical viewpoint. For the first time, we carried out a complete sampling to clarify generic boundaries based on a well-resolved phylogeny of Saussureinae. We employed a Hyb-Seq technique that targets 1061 nuclear conserved ortholog loci designed for Compositae. After a filtering of potential paralogs, 588 loci were retained to infer phylogenetic trees under concatenation and coalescence approaches. High branch support resolution was recovered at the generic level, but a non-monophyletic pattern was detected for most of the genera as they are currently circumscribed. Accordingly, we propose a new generic delimitation based on the three main clades recovered in the backbone tree, which are also in agreement with morphological evidence: Dolomiaea, Saussurea, and Jurinea. Following this classification into three genera, 18 new combinations are proposed. This new genus delineation will be used as a basis for future evolutionary studies in the Saussureinae.Financial support from the Ministerio de Ciencia e Innovación (Project CGL2015-66703-P MINECO/FEDER, UE and Ph.D. grant to Sonia Herrando-Moraira) and the Catalan government (“Ajuts a grups consolidats” 2017-SGR1116) is also greatly acknowledged. This study has been performed under the Ph.D. program “Plant Biology and Biotechnology” of the Autonomous University of Barcelona (UAB)INTRODUCTION MATERIALS AND METHODS RESULTS DISCUSSION TAXONOMIC IMPLICATIONS NOTE ADDED IN PROOF AUTHOR CONTRIBUTIONS ACKNOWLEDGMENT

The recent neophyte Opuntia aurantiaca (Cactaceae): distribution and potential invasion in the Iberian Peninsula

The Cactaceae, and especially its most emblematic genus, Opuntia, is one of the groups of plants with greater invasion potential in the Iberian Peninsula. One of the most recently detected species is Opuntia aurantiaca, a small cactus with an enormous capacity of dispersion. Probably native to Argentina and Uruguay, it behaves as a very aggressive invader in Australia and South Africa. In Europe, it only occurs on the Mediterranean coast of the Iberian Peninsula (Catalonia and Valencian Community). In this study, the geographic range of the species is accurately delineated at the peninsular level. Detected firstly at the beginning of the last decade in Navajas (Castelló Province), it has been subsequently observed in other places of Castelló, but also in Valencia, Tarragona and Barcelona, and since 2017 in Girona. With all gathered occurrence data, the potential distribution of O. aurantiaca is estimated (for the current climatic conditions as well as for different scenarios of global warming). Despite the fact that the species seems to be spreading, maps of potential distribution do not forecast large expansions to other areas of the Iberian Peninsula, both for the present and for the year 2070.This work received financial support from the “Proyecto Intramural Especial, PIE” (grant no. 201630I024) from the CSIC (Spain) and from the “Ajuts a Grups de Recerca Consolidats” (grants nos. 2014-SGR514-GREB and 2017-SGR1116) from the Generalitat de Catalunya (Spain).Abstract Introduction Materials & methods The study species Search for occurrences Ecological niche modelling Results and Discussion Current distribution range of Opuntia aurantiaca in the Iberian Peninsula Present and future potential distribution of Opuntia aurantiaca Acknowledgement