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Typical flowers of taxa of Himantoglossum analysed in the present study, 2: H. adriaticum, H. hircinum, H. calcaratum calcaratum and H. calcaratum jankae (formerly H. jankae and H. caprinum s.n.). Images: Attila Molnár V.
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Background and Aims
The charismatic Himantoglossum s.l. clade of Eurasian orchids contains an unusually large proportion of taxa that are of controversial circumscriptions and considerable conservation concern. Whereas our previously published study addressed the molecular phylogenetics and phylogeography of every named taxon within the clade, here...
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... observation, as the major phenological divergence between these two species meant that it was necessary to freeze the pollinaria of H. robertianum for approximately three months until they could be defrosted and applied to the stigma of the captive H. jankae 'mother' plant. The fact that the resulting F1 plants both grew and flowered vigorously (Fig. 22) suggests that intrinsic sterility barriers are at best weak within the group, even when the parents of the primary hybrids are sampled from within different ...
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... et al., 2012). As already noted, in members of subgenus Himantoglossum-at least, in those individuals that bear labellum markings-the epidermal cells that contain the anthocyanins and so delimit the markings occur within a central region of the labellar epidermis that expands outward to form a continuous mat of prominent, densely packed papillae (Fig. 22B). This papillose mat may offer footholds to visiting insects, but more significantly, it may also be an osmophore that is responsible for much of the volatile organic compounds emitted by the plant (see also Vöth, ...
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... is tempting to view changes in the length and/or width of the two lateral petals and especially of the three sepals as direct functional consequences of the changes documented in the shape and/or size of the third, median petal-the labellum-which must be wholly enclosed throughout the development of the bud (Fig. 22A). These perianth segments became elongated in H. comperianum to accommodate the filiform limbs of its distinctive labellum and broadened in the remaining species to encompass their more robust labella. The entire flower became more compact during the origin of section hircinum, the column also being shortened. Both the column and ...
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... them). With regard to markings on the sepals of Himantoglossum, interior spots arose only after the divergence of H. comperianum, the marginal stripe only after the divergence of the robertianum group, and the interior dashes only after the separation of H. formosum, when the labellum markings also became raised on a central papillate region (Figs. 22C and 23) ). All markings other than sepal marginal stripes were subsequently lost independently from H. formosum and from H. caprinum (which also lost the papillae that are reliably correlated with labellum markings in the hircinum and jankae ...
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... in both the potential for phenotypic diversification but also the developmental and structural constraints that ultimately limit that diversification. For example, the complexity of the labellum is ultimately constrained by the (admittedly remarkably sophisticated and efficient) manner in which the labellum is packaged within the developing bud (Fig. 22A). The relative sizes and shapes of contrasting regions of the labellum offer a marvellous case-study in allometry and especially heterochrony within a single organ (most studies of heterochrony focus on evolutionary changes in the relative timing of developmental events between organs rather than within them). The fluctuations in the ...
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Citations
... (Dirwimmer et al. 2016 Greuter], but that genus is in fact nested in Himantoglossum, see e.g. Bateman et al. (2017). It is now included in the key and a full account is provided. ...
... (Kreutz & Steinfeld 2013) but its taxonomic value requires confirmation. Such parapatric populations may have been the result of polyploidization (Bateman et al. 2017 • D. incarnata (L.) Soó: the infraspecific variability of this species in the Flora area is poorly understood. Rather numerous infraspecific taxa have been reported lately, especially by Kreutz (2019), but the taxonomic value of most of these taxa needs to be confirmed by molecular studies. ...
Following the publication of the seventh edition of the Nouvelle Flore, in 2023, this contribution provides an overview of nomenclatural and taxonomic changes compared to the previous edition of the Flora, which was published in 2012.
... Before drawing specific conclusions regarding phenotypic variation across the genus Ophrys, we will first critically appraise the morphometric approach that we have employed here, in order to establish limits to the strength of the conclusions that can reasonably be drawn from this study. We should begin by noting that this is the 22nd such empirical study of European native orchids that one of us (R.B.) has published using this morphometric approach, together spanning seven genera: Dactylorhiza (6 papers [66,77]), Gymnadenia (2 papers [69,78]), Platanthera (4 papers [79,80]), Pseudorchis (1 paper), Orchis (4 papers [76,81]), Himantoglossum (3 papers [82]) and Ophrys (1 paper [45]). The strengths and weaknesses of our approach have therefore been amply demonstrated empirically. ...
... Admittedly, as demonstrated here, valuable information is rarely obtained from coordinates of lower order than the third. Superimposing a minimum spanning network onto a principal coordinates plot can, in some cases, usefully assist interpretation, though this approach becomes impractical to draft for presentation in ordinations involving large numbers of points; it is most effective when applied to plots ordinating mean values for populations [69,77,82], rather than to plots based on raw data for individual plants, such as those presented herein in Figures 5-16. ...
... The amount of research time invested in such a study therefore depends on the number of characters measured and relative numbers of putative species, populations per putative species and plants per population that are scored, based on a carefully planned and geographically extensive sampling strategy [29,67,68]. Typical figures for these parameters in our previous morphometric studies have been 2-10 putative species, 5-20 populations per species and 10(-20) plants per population [66,69,82,83]. Unfortunately, it was impractical to bring an equivalent level of sampling rigour to the present study, given that the primary objective was to characterise the broader morphological trends across a genus considered by some specialists to contain approximately 400 (micro)species. ...
Simple Summary: Our frequently deployed approach to optimally circumscribing species requires large-scale field sampling within and between populations for large numbers of morphometric characters, followed by multivariate ordinations to objectively seek discontinuities (or, failing that, zones of limited overlap) among sets of populations considered to represent species. Corresponding boundaries are sought in DNA-based outputs, either phylogenies or preferably ordinations based on population genetic data. Herein, we analyse within a molecular phylogenetic framework detailed morphometric data for the charismatic bee orchids (Ophrys), seeking a 'mesospecies' species concept that might provide a compromise between the nine 'macrospecies' recognised primarily through DNA barcoding and the several hundred 'microspecies' recognised primarily through perceived pollinator specificity. Our analyses failed to find robust groupings that could be regarded as credible mesospecies, instead implying that each macrospecies constitutes a morphological continuum. This problematic result encouraged us to reappraise both our morphometric approach and the relative merits of the contrasting macrospecies and microspecies concepts, and to reiterate the key role played by genetics in species circumscription. Abstract: Despite (or perhaps because of) intensive multidisciplinary research, opinions on the optimal number of species recognised within the Eurasian orchid genus Ophrys range from nine to at least 400. The lower figure of nine macrospecies is based primarily on seeking small but reliable discontinuities in DNA 'barcode' regions, an approach subsequently reinforced and finessed via high-throughput sequencing studies. The upper figure of ca. 400 microspecies reflects the morphological authoritarianism of traditional taxonomy combined with belief in extreme pollinator specificity caused by reliance on pollination through pseudo-copulation, enacted by bees and wasps. Groupings of microspecies that are less inclusive than macrospecies are termed mesospecies. Herein, we present multivariate morphometric analyses based on 51 characters scored for 457 individual plants that together span the full morphological and molecular diversity within the genus Ophrys, encompassing 113 named microspecies that collectively represent all 29 mesospecies and all nine macrospecies. We critique our preferred morphometric approach of accumulating heterogeneous data and analysing them primarily using principal coordinates, noting that our conclusions would have been strengthened by even greater sampling and the inclusion of data describing pseudo-pheromone cocktails. Morphological variation within Ophrys proved to be exceptionally multidimensional, lacking strong directional trends. Multivariate clustering of plants according to prior taxonomy was typically weak, irrespective of whether it was assessed at the level of macrospecies, mesospecies or microspecies; considerable morphological overlap was evident even between subsets of the molecularly differentiable macrospecies. Characters supporting genuine taxonomic distinctions were often sufficiently subtle that they were masked by greater and more positively correlated variation that reflected strong contrasts in flower size, tepal colour or, less often, plant size. Individual macrospecies appear to represent morphological continua, within which taxonomic divisions are likely to prove arbitrary if based exclusively on morphological criteria and adequately sampled across their geographic range. It remains unclear how much of the mosaic of subtle character variation among the microspecies reflects genetic versus epigenetic or non-genetic influences and what proportion of any contrasts observed in gene frequencies can be attributed to the adaptive microevolution that is widely considered to dictate speciation in the genus. Moreover, supplementing Biology 2023, 12, 136. https://doi.org/10.3390/biology12010136 https://www.mdpi.com/journal/biology Biology 2023, 12, 136 2 of 52 weak morphological criteria with extrinsic criteria, typically by imposing constraints on geographic location and/or supposed pollinator preference, assumes rather than demonstrates the presence of even the weakest of species boundaries. Overall, it is clear that entities in Ophrys below the level of macrospecies have insufficiently structured variation, either phenotypic or genotypic, to be resolved into discrete, self-circumscribing ("natural") entities that can legitimately be equated with species as delimited within other less specialised plant genera. Our search for a non-arbitrary (meso)species concept competent to circumscribe an intermediate number of species has so far proven unsuccessful.
... The vulnerability of the Iranian H. affine populations has been confirmed by lower intra-population genetic diversity and gene flow [11][12][13][14]. In addition to the effect of overexploitation, global climate change is one of the main reasons for the broad extent of fragmentation observed in H. affine populations on a global scale [15,16]. We decided to investigate H. affine as domestication and commercial Salep production using species with big tubers could be more economical and could help alleviate exploitation pressure on species with small tubers. ...
Himantoglossum affine is a threatened terrestrial orchid. We aimed to optimize asymbiotic seed germination and direct embryogenesis and to analyze the phytochemical profile and physico-biochemical analysis of leaf and tuber. The individual use of organic nitrogen compounds resulted in higher germination efficiencies, while the shortest times to germination were observed using coconut water plus casein hydrolysate. Plantlets grown on media supplemented with pineapple juice and peptone had the highest plantlet length and weight. For embryogenesis, the highest regeneration rate (44%) and embryo number/explant (10.12 ± 2.08) were observed in young protocorm-like body (PLB) explants with 0.5 mg/L naphthalene acetic acid (NAA) and 1 mg/L thidiazuron (TDZ). During the acclimatization process, the scattered vascular tubes converted to fully developed vascular tissues, ensuring maximum sap flux. Gas chromatography–mass spectrometry analysis identified 1,2,3-propanetriol, monoacetate, 4H-pyran-4-one, 2,3-dihydro-3,5-dihydroxy-6-methyl, and 2-butenedioic acid, 2-methyl-, (E)- as the most prevalent compounds. We reported higher contents of total phenolics and flavonoids and antioxidant activity compared to other terrestrial orchids. The glucomannan content (36.96%) was also higher than starch content (31.31%), comparable to those reported in other tuberous orchids. Based on the fragmentation of H. affine populations in the Middle East and Euro-Mediterranean countries due to over-harvesting, climate change, and/or human impact, our procedure offers a tool for the re-introduction of in vitro-raised plants to threatened areas.
... Traditional morphometric analyses, which are defined as the quantitative evaluation of biological forms, have been widely used in various disciplines. In botany, they have been a very useful tool in the field of systematics of various plant families (Benca et al. 2014, Atria et al. 2017, Bateman et al. 2017, including palms (Henderson 2006). This tool is not only useful in the field of taxonomy, but also offers a first indication of the identity and importance of evolutionary forces that may promote or limit population differentiation (Domínguez et al. 1998, Gutiérrez-Ortega et al. 2020. ...
Palms (Arecaceae) represent a challenge for studies of morphological variation for several various reasons. For instance, the height of many species makes them difficult to collect, and in some cases, the size of the leaves or the presence of spines make them hard to process and preserve in herbaria. Arecaceae is one of the most diverse families of monocots (ca 2600 species), and many of its species remain poorly studied. The genus Gaussia has five recognized species. Their distributions are restricted to Central America and the Caribbean islands, and for most of them, their taxonomic descriptions are the only information available. Two species – Gaussia maya and G. gomez‐pompae – are exclusively distributed in Mesoamerica. They occur allopatrically and their populations are isolated from each other. The aim of this work was to evaluate the levels of intra‐ and inter‐population morphological variation in both species and determine the possible factors that influence such a variation. We examined the morphological variation of 13 vegetative structures of 155 adult individuals from nine populations of G. maya and G. gomez‐pompae. The differentiation among populations was significant for almost all the variables and was consistent with an isolation‐by‐distance pattern. A linear discriminant analysis, Mahalanobis distances and UPGMA cluster analysis based on Euclidean distances among populations showed that three morphological groups could be distinguished. One group contained only individuals of G. maya, the second only G. gomez‐pompae, while the third group represents a yet unknown taxon. Pairwise differentiation tests clarified the morphological distinction among the three groups. This work highlights the importance of quantifying morphological variation between populations of little‐studied palms.
... [7] [93][114] [229] ...
Középiskolás koromban határoztam el, hogy a hazai növényvilág ritka és szépséges fajait szeretném fényképezni és vizsgálni. Érdeklődésemet legelőször a kosborfélék keltették fel, és ez a „szerelem” a mai napig megmaradt. Ugyanakkor a kezdetektől kiemelt figyelmet szenteltem a veszélyeztetett fajoknak, amelyek kutatását hivatásomul választottam. Ennek annál is inkább szükségét éreztem, mert a flóra napjainkban olyan gyors ütemben alakul át, hogy ez már egy emberöltő alatt is érzékelhető. Az elmúlt évtizedekben (a rendszerváltás és különösen hazánknak az Európai Unióhoz történt csatlakozása óta) a tájhasználatban és a művelés intenzitásában korábban nem látott mértékű és sebességű változások történtek. Az országhatárokon belül zajló (de a nemzetközi trendekhez illeszkedő) folyamatok mellett évről évre jobban érezhetők az éghajlatváltozásnak az élővilágra gyakorolt hatásai, és érzékelhetőek a növényvilágnak az ezekre adott válaszai. Ez a könyv azzal a céllal született, hogy a színes fényképek révén egyrészt némi betekintést nyújtson növényvilágunk lenyűgöző gazdagságába, másrészt magyar nyelven számoljon be a bemutatott növényekkel kapcsolatos tudományos munkánk eredményeiről. Kutatásaim általában hazai növényritkaságok és mediterrán orchideák elterjedésének, életmódjának, rokonsági viszonyainak jobb megismerésére irányultak. Ám ahhoz, hogy megismerhessük mondjuk a magyar látonya vagy az adriai sallangvirág rendszertani helyzetét, rokonsági-leszármazási viszonyait, szükség volt az adott nemzetség lehetőleg minden fajának eredeti élőhelyén történő vizsgálatára. E növények nyomában jutottam el a Kaukázustól a Kanári-szigetekig számos területre, és ennek köszönhetően szerepelnek ebben a könyvben például kilenc látonya- és kilenc sallangvirágfaj fényképei.
... In addition to H. jankae, the endemic H. calcaratum with a significantly longer spur also occurs in the Western Balkans, a taxon mentioned under the name H. hircinum β calcaratum in the Flora of BiH (Beck-Mannagetta, 1904). Based on morphometric and molecular analysis of all taxa of the genus Himantoglos sum (Bateman et al., 2017) . According to Bateman et al. (2017), H. hircinum does not grow east of Italy, Austria and Germany (see Fig. 5 in Bateman et al., 2017). ...
... Based on morphometric and molecular analysis of all taxa of the genus Himantoglos sum (Bateman et al., 2017) . According to Bateman et al. (2017), H. hircinum does not grow east of Italy, Austria and Germany (see Fig. 5 in Bateman et al., 2017). ...
... Based on morphometric and molecular analysis of all taxa of the genus Himantoglos sum (Bateman et al., 2017) . According to Bateman et al. (2017), H. hircinum does not grow east of Italy, Austria and Germany (see Fig. 5 in Bateman et al., 2017). ...
Albeit rich in a number of taxa, flora of Orchidaceae in Bosnia and Herzegovina has not been sufficiently studied. The paper presents distribution of ten rare and threatened orchid taxa in the territory of Bosnia and Herzegovina. Serapias bergonii and Himantoglossum calcaratum subsp. rumelicum are new for Bosnia and Herzegovina, while for other studied taxa (Serapias parviflora, Serapias vomeracea, Himantoglossum adriaticum, H. calcaratum subsp. calcaratum, Gymnadenia frivaldii, Pseudorchis albida, Orchis quadripunctata, Cypripedium calceolus) in addition to field investigation, checking and revision of herbarium material, as well as literature sources were also used for supplementing distribution data. The distribution maps of these taxa in Bosnia and Herzegovina are created on a 10 km × 10 km MGRS grid system. Based on the IUCN Red List criteria, the threat status in Bosnia and Herzegovina was assessed for all studied taxa. Data concerning their habitat preferences and population size are also provided.
... In recent years, Himantoglossum s.l. has included other taxa of considerable interest and conservation [1,2]. Currently, the expanded genus Himantoglossum is composed of the subgenus Himantoglossum including all the species of the former genus Himantoglossum, the subgenus Barlia, consisting of the two species of the genus Barlia, and the subgenus Comperia consisting only of the former species Comperia comperiana ( Table 1). ...
Citation: Mecca, M.; Racioppi, R.; Romano, V.A.; Viggiani, L.; Lorenz, R.; D'Auria, M. The Scent of Himantoglossum Species Found in Basilicata (Southern Italy). Compounds 2021, 1, 164-173. https://doi. Abstract: The SPME (Solid Phase Microextraction) analysis of the scent of H. hircinum showed the presence of elemicin in the presence of a relevant amount of eugenol. The scent of the sample of H. adriaticum collected in Abruzzo showed the presence 4-amino-5-(4-morpholinylmethyl)-2-oxazolidinone, β-ocimene, decyl decanoate, and 9-tricosene as main components. The sample of H. adriaticum collected at Marsico Nuovo has an aroma where the main component was pentadecyl hexanoate, 9-tricosene, methyleugenol, tetradecane, pentadecane, and elemicin. The samples of H. adriaticum collected at Viggianello showed some similarities in the scent: the main components were 9-tricosene and methyleugenol.
... Ströhlein and Sundermann (1972) reported 2n = 2x = 30 in Himantoglossum comperianum (Steven, 1829) P. Delforge, 1999, andBernardos et al. (2006) reported 2n = 2x = 36 for Himantoglossum metlesicsianum (W.P. Teschner, 1982) P. Delforge, 1999. The chromosomal numbers of the other species of the genus such as H. formosum (Steven, 1813) K. Koch, 1849, H. calcaratum (Beck, 1887) Schlechter, 1927, H. caprinum Sprengel, 1826and H. montis-tauri Kreutz et W. Lüders, 1997(Bateman et al. 2017) are unknown. Cases of aneuploidy with 2n = 36+1B are known in both H. hircinum and H. adriaticum (Capineri and Rossi 1987;D'Emerico et al. 1993). ...
... It is interesting to note that the World Checklist of selected Plant Families (WCSP) reports Barlia robertiana (Loiseleur-Deslongchamps, 1807) Greuter, 1967 as a synonym for Himantoglossum robertianum (Loiseleur-Deslongchamps, 1807) P. Delforge, 1999. Furthermore, in the new classification based on morphological and molecular data (Sramkó et al. 2014), Bateman et al. (2017 Comparative investigations of Himantoglossum hircinum and Barlia robertiana show similar karyotype morphologies, with mainly metacentric chromosomes, low asymmetry and little constitutive heterochromatin. H. hircinum was found to have four nucleoli in interphase nuclei. ...
In our study, FISH mapping using 18S-5.8S-25S rDNA and 5S rDNA sequences was performed for the first time on Ophrys tenthredinifera Willdenow, 1805, Serapias vomeracea (Burman f., 1770) Briquet, 1910 and Himantoglossum hircinum (Linnaeus, 1753) Sprengel, 1826. A detailed study was also performed on O. tenthredinifera using Giemsa-staining, silver-staining, CMA fluorescence banding and fluorescence in situ hybridisation (FISH) with rDNA probes. We analysed two subspecies, i.e. O. tenthredinifera subsp. neglecta (Parlatore, 1860) E.G. Camus, 1908 and O. tenthredinifera subsp. grandiflora (Tenore, 1819) Kreutz, 2004 by the traditional Feulgen method and constructed the karyotype. The cytotaxonomic implications for both taxa are also discussed. In Himantoglossum hircinum , FISH and silver staining highlighted differences in the number of two rDNA families (35S and 5S) with respect to Barlia robertiana (Loiseleur-Deslongchamps, 1807) Greuter, 1967. In addition, fluorescence in situ hybridisation was also applied to diploid (2 n = 2 x = 36) and triploid (2 n = 3 x = 54) Anacamptis morio (Linnaeus, 1753) R.M. Bateman, Pridgeon et M.W. Chase, 1997. As far as we are aware, this is the first case of autotriploidy observed in A. morio .
... During the 180 years after its description [28], almost nothing was known about the species [29], and it was re-discovered in 1994 [30]. According to recent studies, this is an ancient, phylogenetically isolated [31] Plants 2021, 10, 2779 2 of 10 and morphologically well separated [32] bona fide species. It is listed as Vulnerable (Rare) in the IUCN Red List of Threatened Plants [33]. ...
... Plants 2021, 10, x FOR PEER REVIEW 2 of 10 cies [29], and it was re-discovered in 1994 [30]. According to recent studies, this is an ancient, phylogenetically isolated [31] and morphologically well separated [32] bona fide species. It is listed as Vulnerable (Rare) in the IUCN Red List of Threatened Plants [33]. ...
In order to explore their orchid flora, we performed surveys of 96 Azerbaijani burial places in 2018 and 2019. Altogether, 28 orchid taxa were found in 37 visited cemeteries. In the orchid diversity a remarkable pattern was observed: geographic latitude was significantly and positively related to the number of taxa and number of individuals. The most widespread and abundant orchids in Azerbaijani graveyards were Anacamptis pyramidalis and A. papilionacea (found in 23 and 8 cemeteries, respectively). Azerbaijani cemeteries can be important refuges for rare and threatened orchids, e.g., Himantoglossum formosum (three cemeteries), Ophrys sphegodes subsp. mammosa (eight), Orchis adenocheila (two), O. punctulata (three), O. stevenii (one) and Steveniella satyrioides (one). Epipactis turcica, detected in a single locality, was previously unknown to the flora of Azerbaijan. Additionally, we documented orchid tuber (salep) collection in two cemeteries.
... Flower parts descriptive terminology was adopted according to Bateman et al. [43] while we followed Akbulut et al. [44] for the description of papillae. ...
Hybridization can often lead to the formation of novel taxa which can have traits that resemble either or both parental species. Determining the similarity of hybrid traits to parental taxa is particularly important in plant conservation, as hybrids that form between rare and common taxa may more closely resemble a rare parental species, thereby putting the rare parental taxon at further risk of extinction via increased backcrossing and introgression. We investigated the floral (morpho-logical and chemical) traits and orchid mycorrhizal (OrM) fungal associations of the endangered orchid Orchis patens, its more common sister species O. provincialis, and their natural hybrid O. × fallax in natural sympatric populations. We found that both morphological and chemical floral traits of O. × fallax are shared by the parents but are more similar to O. patens than O. provincialis. OrM fungi were shared among all three taxa, indicating that the availability of OrM fungi should not represent a barrier to establishment of individuals of any of these taxa. These results suggest that O. × fallax may be able to expand its distribution within a similar niche to O. patens. This highlights the importance of quantifying differences between hybrids and parental taxon in species conservation planning.
