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Aesculus hippocastanum L. (European Horse-chestnut) constitutes a biogeographical relict species of the Balkan Peninsula, occurring in isolated and topographically distinct localities in Albania, Bulgaria, Greece and North Macedonia. Despite its great botanical, ornamental and pharmaceutical value, a thorough investigation of Ae. hippocastanum habitat diversity in its native distribution range has not been conducted yet. The present study aims at the syntaxonomic classification and ecological features of plant communities dominated by this species across its overall native distribution range. On the basis of 55 phytosociological relevés, five ecologically, floristically, and spatially well differentiated clusters were identified, with the main revealed gradients of differentiation being geographic location (longitude, latitude), altitude, annual precipitation and precipitation seasonality. The distinct microhabitats with a special refugial character where these plant communities occur meet the species' requirement for relatively high air and soil humidity. They have allowed the preservation of Ae. hippocastanum through time highlighting their great conservational value. The last one could be useful for the implementation of some appropriate measures for effective conservation of these communities. [ABSTRACT FROM AUTHOR]

Pinus heldreichii Christ. (Bosnian pine), a Tertiary relict and Balkan sub-endemic, has not been comprehensively studied for its essential oil (EO) profile and bioactivity of its different plant parts. This study aimed to determine the EO yield, composition and antimicrobial activity from different parts of P. heldreichii at three different populations (mountains) in Bulgaria. Furthermore, the study assessed the antioxidant activities of plant tissue, including leaves (needles), twigs wood, male and female cones. The EOs yield from different plant parts ranged from 0.09 % (leaves) to 0.74 % (wood of twigs), with monoterpenes being the predominant class. Limonene, α-pinene, β-caryophyllene, germacrene D, β-pinene, and β-myrcene were detected in the EO extracted from all analyzed trees. However, these compounds were not found in the EO extracted from all plant parts of the same trees. Four chemical groups (chemotypes) were identified for EO from twigs, and three chemotypes were identified for EO from leaves. The chemotypes were based on the percent ratio of the main EO constituents (>5 %). Leaves tissue showed the highest values in terms of polyphenols and flavonoids, as well as higher ABTS radical scavenging activity, while the highest antimicrobial activity against Staphylococcus aureus subsp. aureus was seen in the EOs obtained from twigs. This is the first study to identify several chemotypes based on leaf and twigs EO of P. heldreichii distributed in Bulgarian flora. Furthermore, the EO of twigs tips (TT), male cones (MC), and wood of one-two-year-old twigs (WT) of the same trees were reported for the first time. The total polyphenol, flavonoid content, and radical scavenging activity of tissues of annual twigs wood and biennial twigs wood, leaf tissue, MC tissue, and the twigs tips tissue is also reported for the first time in the accessible literature. These findings highlight the potential of P. heldreichii to provide EOs with varying compositions and bioactivities, making them suitable for nutraceutical, pharmacological, and potentially food additive applications. Furthermore, the identification of chemotype accessions in this study suggests their selection for the development of new forest crop as a source for natural products with desirable composition and bioactivity.

Waldsteinia is a small plant genus inhabiting the temperate regions of the Northern Hemisphere. According to the latest revisions, Waldsteinia is included in Geum. We have obtained a phylogenetic reconstruction based on the nuclear (ITS) and plastid (trnL-trnF) DNA to understand the phylogenetic structure of Waldsteinia and its relationships with other taxa of Geum s.l. Phylogenetic analysis based on the joint ITS + trnL-trnF dataset demonstrated Waldsteinia monophyly. The phylogenetic relationships of Waldsteinia species were better explained by their geographical distribution than their morphology. Hence, Euro-Siberian, Northeast Asian, and North American phylogeographic groups were distinguished, with East Asia having been suggested as the place of Waldsteinia origin. Considering the incongruence in W. geoides (a type species) position on the plastid and nuclear DNA trees, together with the discrepancy between the species morphology and its location on the plastid DNA tree, a hybrid origin was suggested for this species. Despite the fact that the position of W. maximowicziana is still not fully resolved, we support the point of view that claims it should be separated from the W. ternata aggregate (traditionally including W. trifolia, W. ternata s.str., and W. maximowicziana) and considered a separate species. The American W. doniana, W. fragarioides, and W. lobata belong to a single maternal lineage, but the observed genetic differences are too small to serve as a convincing argument for species segregation, so their relationships still remain unresolved. [ABSTRACT FROM AUTHOR]

Fritillaria Tourn. ex L. is a genus of Liliaceae including a little more than 150 species occurring in the temperate Holarctic. Fritillaria sonnikovae Shaulo & Erst is the most recently described Siberian species in the genus. In the affinity section of the F. sonnikovae diagnosis, only F. dagana Turcz. and F. roylei Hook. are mentioned. Our study is an original attempt to shed light on the F. sonnikovae origin and its evolutionary relationships with other Fritillaria using nuclear (ITS) and plastid (matK + rps16 + trnH-psbA) DNA markers. Our results showed that F. sonnikovae together with F. dagana and F. maximowiczii Freyn belongs to the North Asian lineage of the Liliorhiza subgenus and produced no evidence supporting relationship between F. sonnikovae and F. roylei. Monophyly of Fritillaria sonnikovae was not reliably confirmed in our study since its close affinity with F. maximowiczii was demonstrated by phylogenetic analysis and morphology. Fritillaria dagana was shown to be a sister to the F. maximowiczii + F. sonnikovae group. Most authors of the present study suggest considering F. sonnikovae as a synonym for F. maximowiczii. In this view, F. sonnikovae may be considered a narrow endemic and one of the light-perianth morphs of F. maximowiczii, which has emerged in the Western Sayan and remained there as a tertiary relict. [ABSTRACT FROM AUTHOR]

Introduction: East Asia (EA), which falls within the region of the Asian monsoon that is composed of the East Asia monsoon (EAM) and the Indian monsoon (IM), is known for its high species diversity and endemism. This has been attributed to extreme physiographical heterogeneity in conjunction with climate and sealevel changes during the Pleistocene, this hypothesis has been widely proven by phylogeographic studies. Recently, dated phylogenies have indicated that the origins (stem age) of the flora occurred after the Oligocene–Miocene boundary and are related to the establishment of the EAM. Methods: Hence, this study further examined whether the strengthening of the monsoons triggered floral evolution via a meta-analysis of the tempo-spatial pattern of evolutionary radiation dates (crown ages) of 101 endemic seed plant genera. Results: Taxonomic diversification began during the late Eocene, whereas the accumulated number of diversifications did not significantly accelerate until the late Miocene. The distribution of the weighted mean and the average divergence times in the EAM, IM, or transitional regions all fall within the midlate Miocene. Fossils of the Tertiary relict genera are mostly and widely distributed outside EA and only half of the earliest fossils in the EA region are not older than Miocene, while their divergence times are mostly after the late Miocene. The pattern of divergence time of monotypic and polytypic taxa suggest the climatic changes after the late Pliocene exert more influence on monotypic taxa. Discussion: The two key stages of floral evolution coincide with the intensifications of the EAM and IM, especially the summer monsoon which brings a humid climate. An integrated review of previous studies concerning flora, genus, and species levels further supports our suggestion that monsoon intensification in EA triggered the evolution of its flora. [ABSTRACT FROM AUTHOR]

Eranthis Salisb. (Ranunculaceae) is a herbaceous plant genus, including few species disjunctively distributed throughout the temperate zone from Southeastern Europe to Eastern Asia. Until recently, only Eranthis sibirica DC. was known in South Siberia, being considered endemic and tertiary relict. Not long ago, Eranthis tanhoensis Erst was also described in Siberia. We report here a reconstruction of the phylogenetic relationships between the Siberian Eranthis species based on nuclear (ITS) and plastid (trnL + trnL-trnF + trnH-psbA) DNA. The phylogeographic structure of Siberian Eranthis is distinguished by the presence of the two "eastern" and "western" supergroups, which most likely formed as a result of disjunction caused by active mountain uplifts during the late Neogene–early Quaternary and subsequent progressive Pleistocene cooling. The eastern supergroup combines lineage I, containing populations from the eastern Khamar-Daban Ridge, the Eastern Sayan Mountains, and the Tannu-Ola Ridge, and lineage II containing western Khamar-Daban populations. The western supergroup includes only lineage III, containing Western Sayan populations. Our data clearly show that E. tanhoensis is nested in the E. sibirica clade, thereby indicating that its description as a separate species is unjustified, as it compromises the monophyletic status of E. sibirica. Therefore, we suggest here to consider E. tanhoensis as a synonym of E. sibirica. [ABSTRACT FROM AUTHOR]

Eranthis Salisb. (Ranunculaceae) is a herbaceous plant genus, including few species disjunctively distributed throughout the temperate zone from Southeastern Europe to Eastern Asia. Until recently, only Eranthis sibirica DC. was known in South Siberia, being considered endemic and tertiary relict. Not long ago, Eranthis tanhoensis Erst was also described in Siberia. We report here a reconstruction of the phylogenetic relationships between the Siberian Eranthis species based on nuclear (ITS) and plastid (trnL + trnL-trnF + trnH-psbA) DNA. The phylogeographic structure of Siberian Eranthis is distinguished by the presence of the two “eastern” and “western” supergroups, which most likely formed as a result of disjunction caused by active mountain uplifts during the late Neogene–early Quaternary and subsequent progressive Pleistocene cooling. The eastern supergroup combines lineage I, containing populations from the eastern Khamar-Daban Ridge, the Eastern Sayan Mountains, and the Tannu-Ola Ridge, and lineage II containing western Khamar-Daban populations. The western supergroup includes only lineage III, containing Western Sayan populations. Our data clearly show that E. tanhoensis is nested in the E. sibirica clade, thereby indicating that its description as a separate species is unjustified, as it compromises the monophyletic status of E. sibirica. Therefore, we suggest here to consider E. tanhoensis as a synonym of E. sibirica.

The Balkan Peninsula is renowned as a refugium of forest species, which were more widespread in the Tertiary. We here investigated three species from one of the largest genera of flowering plants, Euphorbia, which survived the Pleistocene glaciations in the Balkan Peninsula, but responded differently to Holocene warming. Using ITS sequences, multivariate morphometrics and relative genome size (RGS) measurements, we explored relationships among E. amygdaloides, E. heldreichii and E. orjeni, for which different taxonomic treatments have been proposed. The ITS-inferred phylogenies indicate that all three species form independent evolutionary lineages, which have similar RGS, but differ morphologically, and should thus be treated as independent species. The most enigmatic of them, E. orjeni, was found for the first time after 100 years at its type locality in Montenegro, and, based on herbarium revision, was discovered also in the surroundings of Belgrade. Euphorbia heldreichii is more widespread, distributed in the southern Balkan Peninsula, whereas E. amygdaloides occurs in forests throughout Europe, North Africa and Western Asia. Our study emphasises the importance of further botanical exploration of the Balkan Peninsula, a hotspot of European biotic diversity. [ABSTRACT FROM AUTHOR]

The article discusses the relationships between Tertiary relict and circumboreal woodland floras in Chimaphila (Ericaceae). Topics discussed include generation of phylogenies of Chimaphila on the basis of cpDNA and nuclear ITS; the use of multiple DNA markers in improving phylogenetic resolution at low taxonomic levels and direction of movement of Pyroloideae lineages.

• Dracaena draco is able to germinate from 12 °C to 33 °C. • Dracaena draco requires moist and dark conditions to germinate. • Native Dracaena draco occurrences are in areas with temperatures below optimum. The Canary Islands dragon tree (Dracaena draco (L.) L. subsp. draco) is an endangered Tertiary relict species endemic to the Canary and Madeira archipelagos. Based on an ex-situ experiment under controlled conditions, we investigated its germination response to both constant and alternating temperatures, moisture levels (water potential), exposure to light, and desiccation. We found that suitable temperatures for germination (> 5 % of germinated seeds) range from 12.1 °C to 32.8 °C (thermal germination niche limits) and optimal temperatures (> 95 % of germinated seeds) range from 16.6 °C to 28.3 °C. Light exposure and water potential below 0 MPa drastically decreased germination performance while seed desiccation pre-treatments and alternating temperatures had slightly negative effects. Comparing the spatial projection of the thermal germination niche with actual occurrences of D. draco subsp. draco revealed that native populations are mainly distributed in areas where temperatures are colder than the optimum for germination. For native populations, germination rate is predicted to reach 78 % on average based on temperature requirements. Our results suggest that niche conservatism and adaptation are involved in shaping the germination requirements of the dragon tree. [ABSTRACT FROM AUTHOR]

Pinus heldreichii Christ. (Bosnian pine), a Tertiary relict and Balkan sub-endemic, has not been comprehensively studied for its essential oil (EO) profile and bioactivity of its different plant parts. This study aimed to determine the EO yield, composition and antimicrobial activity from different parts of P. heldreichii at three different populations (mountains) in Bulgaria. Furthermore, the study assessed the antioxidant activities of plant tissue, including leaves (needles), twigs wood, male and female cones. The EOs yield from different plant parts ranged from 0.09 % (leaves) to 0.74 % (wood of twigs), with monoterpenes being the predominant class. Limonene, α -pinene, β- caryophyllene, germacrene D, β- pinene, and β -myrcene were detected in the EO extracted from all analyzed trees. However, these compounds were not found in the EO extracted from all plant parts of the same trees. Four chemical groups (chemotypes) were identified for EO from twigs, and three chemotypes were identified for EO from leaves. The chemotypes were based on the percent ratio of the main EO constituents (>5 %). Leaves tissue showed the highest values in terms of polyphenols and flavonoids, as well as higher ABTS radical scavenging activity, while the highest antimicrobial activity against Staphylococcus aureus subsp. aureus was seen in the EOs obtained from twigs. This is the first study to identify several chemotypes based on leaf and twigs EO of P. heldreichii distributed in Bulgarian flora. Furthermore, the EO of twigs tips (TT), male cones (MC), and wood of one-two-year-old twigs (WT) of the same trees were reported for the first time. The total polyphenol, flavonoid content, and radical scavenging activity of tissues of annual twigs wood and biennial twigs wood, leaf tissue, MC tissue, and the twigs tips tissue is also reported for the first time in the accessible literature. These findings highlight the potential of P. heldreichii to provide EOs with varying compositions and bioactivities, making them suitable for nutraceutical, pharmacological, and potentially food additive applications. Furthermore, the identification of chemotype accessions in this study suggests their selection for the development of new forest crop as a source for natural products with desirable composition and bioactivity., Competing Interests: The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (© 2023 The Authors.)

Tertiary relict plant species of Europe have had a large distribution range before the Pleistocene but today are confined to small refugial areas. Syringa josikaea of the largely East Asian genus Syringa is a shrub of temperate forests in the Carpathians, restricted to altogether 25 small populations in two disjunct areas, the Apuseni Mountains ( Romania) and the Ukrainian Carpathians. Miocene and Pleistocene fossil remains indicate the long-term presence of the species in Central Europe; hence S. josikaea has been considered a Tertiary relict. We aimed at clarifying the historical biogeography of S. josikaea by estimating the divergence time between S. josikaea and its Asian relatives, and by analysing intraspecific variation using multiple DNA sequences as well as microsatellites. The estimated divergence time between S. josikaea and its closest relatives based on nuclear ribosomal DNA sequences is 1.88 Mya [0.30-4.04 Mya highest posterior density], suggesting a relatively recent disruption of a formerly continuous distribution area. This time corresponds to the period of Early Pleistocene extinctions, when many Tertiary plant taxa went extinct in Europe. Sequence variation was found to be very low within S. josikaea. Complete identity among all samples in cp DNA may imply a bottleneck. Four different ribotypes found showed no geographic differentiation between the Apuseni Mountains and the Ukrainian Carpathians, and differentiation between these two areas was weak when considering microsatellite variation. Together these observations may imply that the extant disjunct distribution of the species arose only recently, either through colonization from one glacial refugial area or from disruption of one such area. © 2015 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, ●●, ●●-●●. [ABSTRACT FROM AUTHOR]

Premise of research. Even though the Balkan Peninsula is a biodiversity hot spot in southeastern Europe harboring many endemic plants, very little is known about the temporal extent of the phylogeographic history and the contemporary genetic diversity of the endemics there. Haberlea rhodopensis is one of the European Gesneriaceae species occurring in this region and represents appropriate study material to address these questions. Methodology. We generated fossil-dated molecular phylogenies (atpB-rbcL, trnH-psbA, trnLF, ITS) across the Lamiales to determine the origin and age of H. rhodopensis and conducted phylogeographic (trnH-psbA, ITS) and population genetic (ISSRs) analyses on 17 populations from Bulgaria and Greece, covering the entire distribution range of the species, to investigate their biogeographic history, present-day genetic diversity, and differentiation levels. Pivotal results. The European Gesneriaceae genera have a Tertiary origin in the early Oligocene, while the Haberlea lineage emerged in the late Oligocene. Extinctions appear to have marked the history of the genus for a long period of time, and the extant populations diverged in the late Pleistocene. A significant differentiation was apparent between populations from Bulgaria, that is, the Balkan Mountains in the north and the Bulgarian side of the Rhodopi massif in the south, but there was an even stronger differentiation between the latter and populations from Greece. This might be explained by the Rhodopi massif representing a barrier to gene flow, enforced during the Last Glacial Maximum, during which populations descended on opposing north- and south-facing mountain slopes. Conclusions. Haberlea represents an ancient lineage with recent diversification. The extant populations are of recent origin and indicate glacial refugial areas in Bulgaria and Greece. This study sheds light on historic and current phylogenetic and phylogeographic events that shaped the flora of the Balkan Peninsula, an area that has long been recognized for its species diversity and richness. Our data suggest that it may have acted as a persistent refugial area in southeast Europe since the mid-Tertiary. [ABSTRACT FROM AUTHOR]

Background and Aims In the Northern Hemisphere, Tertiary relict disjunctions involve older groups of warm affinity and wide disjunctions, whereas circumboreal distributions in Arctic-Alpine taxa tend to be younger. Arctic-Alpine species are occasionally derived from Tertiary relict groups, but Pyrola species, in particular, are exceptional and they might have occurred multiple times. The aim of this study was to reconstruct the biogeographic history of Pyrola based on a clear phylogenetic analysis and to explore how the genus attained its circumboreal distribution. Methods Estimates of divergence times and ancestral geographical distributions based on neutrally evolving DNA sequence variation were used to develop a spatio-temporal model of colonization patterns for Pyrola. Key Results Pyrola originated and most diversification occurred in Asia; North America was reached first by series Scotophyllae in the late Miocene, then by sub-clades of series Pyrola and Ellipticae around the Pliocene. The three circumboreal taxa, P. minor, P. chlorantha and the P. rotundifolia complex, originated independently of one another, with the last two originating in Asia. Conclusions Three circumboreal Pyrola lineages have arisen independently and at least two of these appear to have originated in Asia. The cool, high-altitude habitats of many Pyrola species and the fact that diversification in the genus coincided with global cooling from the late Miocene onwards fits a hypothesis of pre-adaptation to become circumboreal within this group. [ABSTRACT FROM PUBLISHER]

Introduction: East Asia (EA), which falls within the region of the Asian monsoon that is composed of the East Asia monsoon (EAM) and the Indian monsoon (IM), is known for its high species diversity and endemism. This has been attributed to extreme physiographical heterogeneity in conjunction with climate and sea-level changes during the Pleistocene, this hypothesis has been widely proven by phylogeographic studies. Recently, dated phylogenies have indicated that the origins (stem age) of the flora occurred after the Oligocene-Miocene boundary and are related to the establishment of the EAM., Methods: Hence, this study further examined whether the strengthening of the monsoons triggered floral evolution via a meta-analysis of the tempo-spatial pattern of evolutionary radiation dates (crown ages) of 101 endemic seed plant genera., Results: Taxonomic diversification began during the late Eocene, whereas the accumulated number of diversifications did not significantly accelerate until the late Miocene. The distribution of the weighted mean and the average divergence times in the EAM, IM, or transitional regions all fall within the mid-late Miocene. Fossils of the Tertiary relict genera are mostly and widely distributed outside EA and only half of the earliest fossils in the EA region are not older than Miocene, while their divergence times are mostly after the late Miocene. The pattern of divergence time of monotypic and polytypic taxa suggest the climatic changes after the late Pliocene exert more influence on monotypic taxa., Discussion: The two key stages of floral evolution coincide with the intensifications of the EAM and IM, especially the summer monsoon which brings a humid climate. An integrated review of previous studies concerning flora, genus, and species levels further supports our suggestion that monsoon intensification in EA triggered the evolution of its flora., Competing Interests: The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest., (Copyright © 2022 Ye, Tian and Li.)

The persistence of the Tertiary relict tree Tetracentron sinense Oliv. on the eastern slope of the Ailao Mountains, Yunnan, SW China, was here studied in terms of population structure (size, age) and regeneration patterns. T. sinense occurred in unstable habitats by stream banks, on steep slopes, on scree slopes, or on roadsides near streams in narrow valleys, all places subject to frequent natural disturbances, whereas none were found on stable gentle slopes free of major disturbances at similar altitudes. Further, no established saplings of T. sinense were found in forests having high bamboo ( Yushania crassicollis Yi) coverage in their understory. The size and age structure of T. sinense were multimodal. The reproduction of the tree was either by means of abundant minute wind-dispersed seeds or by resprouts in unstable habitats. These populations depended on disturbance or gap regeneration to survive. T. sinense, along with other tree life-forms including evergreen broad-leaved species and conifers, dominated in the forest canopy layer, even reaching the emergent layer in places. Results of the study provide insight into the ecological characteristics and survival mechanisms of this East Asian paleoendemic tree species. The study will provide a scientific basis for recommendations for the conservation of this species and for other Tertiary relict plants having similar regeneration dynamics. [ABSTRACT FROM AUTHOR]

Leaf fossils of Rosa lignitum Heer have often been documented as an accessory element in mixed mesophytic forests from the Oligocene and Miocene of central Europe. Its relationship to extant rose species is not yet firmly understood because leaf morphology contributes only marginally to the current taxonomy of the genus Rosa L. In this study, we investigate which extant rose species have structural leaf characters most similar to those of R. lignitum and whether the distribution and ecology of these extant species fit with the conditions hypothesized for the paleovegetation containing R. lignitum. Therefore, we examined the leaf characters of 32 extant rose species for their diagnostic value in taxonomy and compared these data with characters observed in R. lignitum from five Paleogene floras in central Europe. Most remarkably, we detected semicraspedodromous venation in R. lignitum. This venation pattern was found only in a few species of Rosa distributed in Southeast Asia, whereas the majority of extant rose species developed strictly craspedodromous venation. The distribution of extant species with semicraspedodromous venation in mixed broad-leaved deciduous and evergreen forests in subtropical China clearly corresponds to the hypothesized mixed mesophytic forests in the European paleovegetation. Thus, this study of detailed leaf morphology of Rosa again supports the previous assumption that the zonal vegetation of the central European Oligocene is partly comparable to extant forest types in Southeast Asia. [ABSTRACT FROM AUTHOR]