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Growth habits of different taxa from the Pinus mugo s. lato complex or aggregate (P. mugo agg.); compiled from drawings of Kindel (1995)
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The Pinus mugo complex (P. mugo Turra sensu lato, and/or species aggregate or group - P. mugo agg., including its internal as well as external aggregate hybrids) is an important fragment of European dendroflora. For its high variability this complicated group has not been solved in taxonomical terms satisfactorily yet. Most problems are mainly caus...
Citations
... The dwarf mountain pine (Pinus mugo Turra) mainly grows as a shrub adapted to various rocky habitats in the high-altitude mountains, including the Alps, Pyrenees, Carpathians, and Balkans [1]. It shows a high capacity for vegetative reproduction and is characterized by a large morphological diversity [2,3]. A related species found in the Pyrenees and Western Alps is distinguished as a separate taxon, P. uncinata, which develops a 12-20 m high tree form [2,4]. ...
... m a.s.l. [3,5]. ...
Pinus mugo plays a significant ecological role in the natural environment at high altitudes in the mountains including the Alps, Pyrenees, Carpathians, and Balkans. In such severe conditions, it is subjected to the harmful effects of various abiotic and biotic factors. In one of the areas of its natural occurrence in Tatra Mts. (southern Poland), for the last few years, a significant intensification of needle disease has been observed. Symptoms similar to those recorded in Tatra Mts. also occur on other Pinus species in Europe and North America, where they are caused by fungi belonging to the genus Elytroderma, Lophodermella, Lophophacidium or Ploioderma (Rhytismataceae). The current paper presents the results of research which was mainly aimed at characterization of disease symptoms observed for the first time in Poland on P. mugo needles, and identification of the main causal agent with use of the morphological and molecular technique. Based on the analyses performed at different times of the year (2015–2020), it was found that dieback symptoms initially appeared only on first-year needles, a few weeks after their development. Symptoms occur on one or both needles in the bundle. The distal parts of the needles died, while the basal parts remained green. In the following year, mainly in June and July, on the previous year’s needles attached to the shoots, mature ascomata can be seen. The fungus Lophodermella sulcigena has been identified as the cause of these symptoms. So far, the related species L. conjuncta has not been found. The morphological features of the pathogen microstructure produced on P. mugo needles are presented. Attention was drawn to certain features that may make its identification difficult, especially in terms of shapes and sizes of ascospores. The phylogenetic position of the identified causal agent in relation to closely related other species was determined. The current results confirmed that L. sulcigena shows great phylogenetic similarity to L. montivaga, which is found in North America. Nine rDNA barcode sequences of L. sulcigena obtained in this work will enrich the NCBI GenBank database. The obtained results, indicating the presence of other fungi in L. sulcigena ascomata, which may limit the spread of its ascospores, were also discussed.
... Botanical nomenclature follows The Plant List (TPL) and the following specific criteria particularly referred to Pinus fragments. Anatomically, the identification of the species belonging to the European mountain pine complex (Hamerník and Musil, 2007;Sokołowska et al., 2021) is difficult as diagnostic traits are very similar and hybridization is frequent; in anthracology this difficulty is even greater given the poorly preserved small fragments available for taxonomic identification (Schweingruber, 1990;Allué et al., 2018;Figueiral and Carcaillet 2005). In the present study, the distinction between the different mountain pine species followed the criteria proposed by Euba (2008) (e.g., width of growth rings, position of the resiniferous channels etc.) but above all the local biogeographical setting. ...
... Norway spruce is considered the most abundant treeline-forming tree species (Treml and Banaš 2008) and dominates montane forests up to the timberline, forming clonal groups or solitary trees above the limit of the closed forest in Central Europe (Šenfeldr and Maděra 2011). Prostrate dwarf pine is limited to alpine areas from the Pyrenees to the Balkan Peninsula and is common in the altitudinal belt above the upper limit of closed forests in the eastern Alps, Sudetes, and Carpathians (Hamerník and Musil 2007). However, it expands into surrounding communities in many areas of the Carpathians (Švajda et al. 2011), Apennines (Calabrese et al. 2018), Alps (Dullinger et al. 2003), and Sudetes (Treml et al. 2010), which has a strong impact on biodiversity (Zeidler et al. 2021). ...
The advancement of upper forest limits is driven by environmental conditions, but our current understanding overlooks the attributes of habitats and germination ability of woody species. Habitats, through plant litter, impact the competitive relation of germination and seedling growth. The aims of this study were to identify the selective effect of six litter leachates on the germination of indigenous Norway spruce (Picea abies) and nonindigenous dwarf pine (Pinus mugo) and compare the germination rates of the species. We collected plant litter and seeds from the (sub)alpine belt of the Hrubý Jeseník Mts. (the Eastern Sudetes Mts.; the Czech Republic). We evaluated the effect of plant litter leachates from alpine heathlands, wind-swept alpine grasslands, subalpine tall-herb plants, Pinus mugo scrub, subalpine Vaccinium vegetation, and Norway spruce clonal groups on germination process under standard light and temperature conditions. The germination of Norway spruce was inhibited by the litter leachate from subalpine tall-herb vegetation mainly dominated by Calamagrostis villosa, whereas that of dwarf pine was not. The other five litter leachates had no significant effect on the both. Under standard conditions, the germination time of dwarf pine is on average one day faster. These results suggest that most of the litter leachates examined may have small impacts on the germinability and germination time of Norway spruce and dwarf pine, but litter from the subalpine tall-herb vegetation can act as a filter that influences the seedling composition of the woody species.
... Гибриды сосны обыкновенной (Р. sylvestris) с сосной горной встречаются по всему ареалу последнего вида (Christensen, 1987;Hamerník, Musil, 2007), гибриды кедра сибирского (P. sibirica) с кедровым стлаником -по всей области наложения ареалов (Goroshkevich et al., 2008;Горошкевич и др., 2010). ...
For the first time, a comparative analysis of crown morphogenesis in Siberian stone pine (Pinus sibirica), Siberian dwarf pine (P. pumila), their hybrids and backcrosses was carried out using the example of 10-year-old seed offspring from the Northern Baikal natural population. The goal is to “decompose” the life form phenomenon into elementary traits, to trace their formation at the beginning of ontogenesis on two contrasting examples (an upright tree vs. prostrate tree) and inheritance after interspecific hybridization. The parent species differed fundamentally in (1) stem inclination and (2) apical dominance at two levels (1-year-old shoots and perennial branches). The phototropic slope of the Siberian dwarf pine stem “triggered” the mechanism of prostrate tree formation. Apical dominance in the system of perennial branches was the main factor in the crown formation. It, in turn, consisted of two key characteristics: strength and selectivity. In Siberian stone pine, apical dominance was strong and nonselective. In Siberian dwarf pine, apical dominance was weak and selective: some of the largest branches did not obey him. All key traits were inherited by hybrids and backcrosses strictly intermediate in accordance with the proportion of species in the offspring genome.
... shrubby habit (multiple trunks, up to a few meters high), which inhabit central and eastern massifs-Alps, Sudetes, Carpathians, and Balkan mountain chains (Hamerník & Musil, 2007). Pyrenean pine is a typical coniferous tree (grows straight and may reach up to 25 m), which occurs from the Alps westward-in the Massif Central, Jura and Vosges, Pyrenees, and a few remote populations inside the Iberian Peninsula (Jalas & Suominen, 1973). ...
... Pyrenean pine is a typical coniferous tree (grows straight and may reach up to 25 m), which occurs from the Alps westward-in the Massif Central, Jura and Vosges, Pyrenees, and a few remote populations inside the Iberian Peninsula (Jalas & Suominen, 1973). Although they are distinguished by a number of other, more subtle traits, for example, phenology, cone size and shape, needle characteristics, and composition of volatiles and allozymes (please, see Table S1; Lewandowski et al., 2000;Monteleone et al., 2006;Boratyńska & Boratyński, 2007;Boratyńska et al., 2015;Wachowiak et al., 2018;Adams & Tashev, 2019), they are often aggregated under the P. mugo complex together with a few less defined groups (Christensen, 1987;Hamerník & Musil, 2007). For simplicity reasons, and following Businský & Kirschner (2010), we call them here separate species. ...
The environment is a powerful selective pressure for sessile organisms, such as plants, and adaptation to the environment is particularly important for long‐lived species, like trees. Despite the importance of adaptive trait variation to the survival and success of trees, the molecular basis of adaptation is still poorly understood. Gene expression patterns in three closely related, but phenotypically and ecologically divergent, pine species were analyzed to detect differentiation that may be associated with their adaptation to distinct environments. Total RNA of Pinus mugo, P. uncinata and P. sylvestris samples grown under common garden conditions was used for de novo transcriptome assembly, providing a new reference dataset that includes species from the taxonomically challenging Pinus mugo complex. Gene expression profiles were found to be very similar with only 121 genes significantly diverged in any of the pairwise species comparisons. Functional annotation of these genes revealed major categories of distinctly expressed transcripts including: wood trait properties, oxidative stress response, and response to abiotic factors such as salinity, drought and temperature. We discuss putative associations between gene expression profiles and adaptation to different environments, for example: upregulation of genes involved in lignin biosynthesis in the species which have adapted to mountainous regions characterized by strong winds and thick snow cover. Our study provides valid candidates for verification of the importance of the gene expression role, in addition to evidence for selection within genomic regions, in the process of ecological divergence and adaptation to higher altitudes in pine taxa. This article is protected by copyright. All rights reserved.
... sylvestris L.) and dwarf mountain pine (P. mugo Turra s. str.) dominate ecologically in Central and Southern Europe [4,5]. As typical pioneer woody plants, their parental species occupy sunny to partially shaded, nutrient-poor sites from lowlands and foothills (P. ...
The genetic structures of the four putative hybrid swarms of Pinus sylvestris × P. mugo in Slovakia were analyzed in terms of individual admixture proportions calculated via inter-primer binding site (iPBS) marker loci. This work aimed to reevaluate the hybrid swarms’ differentiation status as postulated in the previous studies at both population and genomic levels. The study confirmed intermediate divergence of each of the swarms examined. Based on 80-loci hybrid index scores, we have revealed the presence of introgressive and intermediate hybrids with frequencies corresponding to differentiation estimates. Surprisingly, irrespective of individual phenotypes, the most frequent intermediates were found in Sucha Hora (29.5%) and Obsivanka (28.6%) populations, which resemble rather pure P. mugo and were previously considered as bimodal hybrid zones with a negligible amount of hybrids. The remaining hybrid zone population Zuberec seems to be highly introgressed to P. sylvestris, while Tisovnica is clearly inclined to P. mugo. The results and different methodologies are discussed.
... The Pinus mugo complex is a large and polymorphic complex of closely related pines native to the main mountains of Europe, including the Pyrenees, the Alps and the Carpathians [1,2]. Some researchers indicate that in this group there may be even more than a hundred endemic forms classified into various taxonomic ranks, i.e., species, subspecies or varieties [1]. ...
The Pinus mugo complex is a large group of closely related mountain pines, which are an important component of the ecosystems of the most important mountain ranges, such as the Alps, Carpathians and Pyrenees. The phylogenetic relationships between taxa in this complex have been under discussion for many years. Despite the use of many different approaches, they still need to be clarified and supplemented with new data, especially those obtained with high-throughput methods. Therefore, in this study, the complete sequences of the chloroplast genomes of the three most recognized members of the Pinus mugo complex, i.e., Pinus mugo, Pinus rotundata and Pinus uncinata, were sequenced and analyzed to gain new insight into their phylogenetic relationships. Comparative analysis of their complete chloroplast genome sequences revealed several mutational hotspots potentially useful for the genetic identification of taxa from the Pinus mugo complex. Phylogenetic inference based on sixteen complete chloroplast genomes of different coniferous representatives showed that pines from the Pinus mugo complex form one distinct monophyletic group. The results obtained in this study provide new and valuable omics data for further research within the European mountain pine complex. They also indicate which regions may be useful in the search for diagnostic DNA markers for the members of Pinus mugo complex and set the baseline in the conservation of genetic resources of its endangered taxa.
... Studies of the adaptive variation of these species are especially important due to environmental changes and ongoing habitat loss in mountainous areas. Although these taxa face rather similar environmental pressures and share a common genetic background [25,31], they differ significantly in a number of characteristics and are widely recognized to be two separate taxa [38]. For evolutionary assessments we used a novel SNP array and the reference Scots pine to, first, determine the discrimination power of this new tool in the studied system, and to elucidate the genetic differentiation and divergence of these species. ...
... Our study provides a new example [40][41][42] of a successful application of a genomewide scan for delineation of phylogenetic relationships among taxonomically challenging plant groups which, in this study, were from the P. mugo complex [28,38]. The two sister taxa appeared separated in multivariate analysis plots, having disjointed 95% confidence intervals in PCA tests. ...
Mountain plants, challenged by vegetation time contractions and dynamic changes in environmental conditions, developed adaptations that help them to balance their growth, reproduction , survival, and regeneration. However, knowledge regarding the genetic basis of species adaptation to higher altitudes remain scarce for most plant species. Here, we attempted to identify such corresponding genomic regions of high evolutionary importance in two closely related European pines, Pinus mugo and P. uncinata, contrasting them with a reference lowland relative-P. sylvestris. We genotyped 438 samples at thousands of single nucleotide polymorphism (SNP) markers, tested their genetic differentiation and population structure followed by outlier detection and gene ontol-ogy annotations. Markers clearly differentiated the species and uncovered patterns of population structure in two of them. In P. uncinata three Pyrenean sites were grouped together, while two outlying populations constituted a separate cluster. In P. sylvestris, Spanish population appeared distinct from the remaining four European sites. Between mountain pines and the reference species, 35 candidate genes for altitude-dependent selection were identified, including such encoding proteins responsible for photosynthesis, photorespiration and cell redox homeostasis, regulation of transcription , and mRNA processing. In comparison between two mountain pines, 75 outlier SNPs were found in proteins involved mainly in the gene expression and metabolism.
... The most known and the best studied are Pinus mugo Turra, Pinus uncinata Ramond, and Pinus uliginosa Neumann ex Wimmer. The taxonomic and evolutionary status in this complex is still not resolved, which is founded by the following: common usage of synonymous names describing probably the same taxa, sympatric occurrence of the taxa accompanied by the presence of hybrid individuals, and ongoing hybridization processes [26][27][28], which altogether led to difficulties in the discrimination of species with a similar morphology, or inaccurate assigning of individuals to a particular taxa. Taxonomic and evolutionary relationships among closely related taxa in the Pinus mugo complex have been studied in detail for many years using various tools and methods, i.e., serological [29], allozymatic [30,31], RAPD markers [32], molecular cytogenetics, and flow cytometry [33,34], as well as the DNA barcoding approach [1]. ...
... In other reports, Pinus × rhaetica and Pinus uliginosa are considered to be hybrid individuals, representing two separate taxa [34]. Even more puzzling, the postulated parental species for the two above-mentioned taxa are exactly the same, i.e., monocormic (Pinus sylvestris) and polycormic (Pinus mugo) [27]. The macroscopically observed variation between the STP profiles was confirmed using digital image processing tools. ...
... In the present study, one taxon (Pinus × rhaetica) did not have any species specific protein band, despite the presence of 24 bands in the protein profile, which is a relatively high number. In the scientific literature, Pinus × rhaetica is sometimes used as a synonymous name of Pinus uliginosa, suggesting the existence of only one taxon [27]. In other reports, Pinus × rhaetica and Pinus uliginosa are considered to be hybrid individuals, representing two separate taxa [34]. ...
The Pinus mugo complex includes several dozen closely related European mountain pines. The discrimination of specific taxa within this complex is still extremely challenging, although numerous methodologies have been used to solve this problem, including morphological and anatomical analyses, cytological studies, allozyme variability, and DNA barcoding, etc. In this study, we used the seed total protein (STP) patterns to search for taxonomically interesting differences among three closely-related pine taxa from the Pinus mugo complex and five more distant species from the Pinaceae family. It was postulated that STP profiling can serve as the backup methodology for modern taxonomic research, in which more sophisticated analyses, i.e., based on the DNA barcoding approach, have been found to be useless. A quantitative analysis of the STP profiles revealed characteristic electrophoretic patterns for all the analyzed taxa from Pinaceae. STP profiling enabled the discrimination of closely-related pine taxa, even of those previously indistinguishable by chloroplast DNA barcodes. The results obtained in this study indicate that STP profiling can be very useful for solving complex taxonomic puzzles.
... The Pinus mugo complex (P. mugo Turra sensu lato or P. mugo aggregate) represents a highly polymorphic species group distributed in the major mountain massifs of central and southern Europe (Hamern ık and Musil 2008). Due to its prominent phenotypic plasticity, geographic and ecological differentiation of populations and their possible hybridization with P. sylvestris L., this complicated group is presently still unsatisfactorily resolved in taxonomical terms (Hamern ık and Musil 2008). ...
... mugo Turra sensu lato or P. mugo aggregate) represents a highly polymorphic species group distributed in the major mountain massifs of central and southern Europe (Hamern ık and Musil 2008). Due to its prominent phenotypic plasticity, geographic and ecological differentiation of populations and their possible hybridization with P. sylvestris L., this complicated group is presently still unsatisfactorily resolved in taxonomical terms (Hamern ık and Musil 2008). Most of the studies agree that the P. mugo complex includes three major entities, i.e., P. mugo Turra sensu stricto (dwarf mountain pine), P. uncinata Ramond ex DC. (mountain pine), and P. rotundata Link (bog pine). ...
Pinus mugo Turra sensu stricto (dwarf mountain pine) is a species native to central and southern Europe, which performs a key role in subalpine areas by preventing avalanches and soil erosion. In this study, headspace extraction coupled with GC-MS/FID analyses was used to analyze diversity of needle volatiles in 10 natural populations of P. mugo s. str. growing in the Julian Alps, southern Carpathians and Balkan Peninsula. The multivariate statistical analyses revealed clear differentiation between Julian Alpine and southern Carpathian populations, supporting the hypothesis about existence of vicariant gene pools in these two European mountain chains. Although several earlier studies have pointed out heterogeneity of the Balkan populations, none of the investigated Balkan populations have shown the volatile pattern similar to the Carpathian group, as they all formed a common group with the Alpine population. AHC of bioclimatic data as well as regression analysis suggested that general volatile differentiation in P. mugo s. str. essentially does not represent an adaptive response to bioclimatic, orographic or geological conditions. Therefore, the described population groups might be provisionally considered as two P. mugo s. str. chemotypes. Our findings regarding volatile characters complement those obtained from morpho-anatomical, phytochemical and molecular data reported so far.
