Figure 1 - uploaded by Richard Bateman
Content may be subject to copyright.
A, Orchis purpurea × simia viewed northward along the Thames. B, Orchis purpurea × simia inflorescence most closely resembling O. simia, Goring. C, Orchis purpurea × simia inflorescence most closely resembling O. purpurea, Goring. D, Orchis simia inflorescence, Goring. E, Orchis purpurea inflorescence, Goring. F, Orchis militaris inflorescence, Marlow. G, Orchis simia × anthropophora inflorescence, Faversham (1985). H, Orchis anthropophora inflorescence, Faversham. All photographs by Richard Bateman.
Source publication
We report the first confirmed occurrence in Britain of Orchis ¥ angusticruris Franch. ex Rouy, a hybrid between two
closely related orchid species of anthropomorphic Orchis (O. purpurea Huds. ¥ O. simia Lam.) that hybridize
frequently in Continental Europe. Seven individual hybrids, most likely F1 plants representing a single interspecific
pollinat...
Contexts in source publication
Context 1
... the more reliably calcicolous anthropomorphic group (c. eight species) the lateral sepals are incorporated into the hood, the flowers are in most cases smaller and shorter-spurred and the four-lobed labellum is more deeply divided, conferring on the flower a strikingly anthropomorphic outline (Figs 1, 2). Molecular phylogenies based on the nuclear internal transcribed spacer (ITS) region suggest that the relatively early divergences of O. anthropophora and/or of the exclusively Mediterranean O. italica render this group paraphyletic (Bateman et al., 2003) (Fig. 3). ...
Context 2
... Britain, the anthropomorphic Orchis species are considered especially charismatic. This partly reflects their extraordinary appearance ( Fig. 1) and inferred aspects of their evolution, such as the apparent reduction to a vestigial condition of the spur in O. anthropophora that until recently led to its taxonomic segregation as a monotypic genus, Aceras ( Bateman et al., 1997;Bateman et al., 2003;Bateman, 2007). However, the increased attention paid to the group has been ...
Context 3
... thereby maximizing the risk of natural cross-pollination (Bateman & Farrington, 1987). In 1998, another Kentish site yielded two hybrid plants identified as O. purpurea ¥ O. anthropophora, although, unfortunately, issues of site confidentiality precluded more detailed scientific investigation (cf. Rose, 1998;Bateman & Farrington, 1999). (Fig. 1A). The reserve has welcomed careful visitors ever since various conservation measures successfully bulked up its once tenuous population of O. simia -the final remnant of a once extensive meta-population that stretched along much of the Thames Valley where it cuts through the chalk landscape of the Chiltern Hills (Paul, 1965;Harrap & ...
Context 4
... these occurrences prompted much speculation regarding their respective origins. Some initial identifications of the plants shown in Figure 1A-C favoured yet another anthropomorphic Orchis that is rare and heavily protected in Britain, O. militaris, which occurs in increasing numbers at two heavily conserved sites further east in the Chilterns (Figs 1F, 2). However, more careful examination suggested that these striking new arrivals represented yet another case of hybridization among the British anthropomorphic Orchis, this time between O. purpurea and O. simia (Bateman, 2006b;Raper, 2006Raper, -2008Fay et al., 2007), to generate O. ¥ angusticruris Franch. ...
Context 5
... these occurrences prompted much speculation regarding their respective origins. Some initial identifications of the plants shown in Figure 1A-C favoured yet another anthropomorphic Orchis that is rare and heavily protected in Britain, O. militaris, which occurs in increasing numbers at two heavily conserved sites further east in the Chilterns (Figs 1F, 2). However, more careful examination suggested that these striking new arrivals represented yet another case of hybridization among the British anthropomorphic Orchis, this time between O. purpurea and O. simia (Bateman, 2006b;Raper, 2006Raper, -2008Fay et al., 2007), to generate O. ¥ angusticruris Franch. ...
Context 6
... Coefficient (Gower, 1971) on unweighted data sets scaled to unit variance. This was in turn used to construct a minimum spanning tree (Gower & Ross, 1969) and subsequently to calculate principal coordinates (Gower, 1966(Gower, , 1985 -compound vectors that incorporate positively or negatively correlated (after Bateman & Farrington, 1989, fig. ...
Context 7
... for plastid microsatellites showed the weakest correlation with taxonomic assignment. Four variable regions yielded a total of 19 haplotypes (Fig. 10), although these formed two main clusters. The first cluster, epitomized by haplotype 15, is dominated by O. simia and lacks O. militaris. The second cluster, epitomized by haplotype 6, is dominated by O. militaris but contains a few individuals assigned to O. simia. Surprisingly, individuals of O. purpurea were distributed ...
Context 8
... Hill Wood in East Kent and more often, apparently at higher frequencies, in several populations from southern France. This haplotype was also inherited from these O. purpurea plants by the O. purpurea ¥ simia hybrids at Goring, where the O. simia plants reliably yielded haplotype 8 -an unusual haplotype similar to the more widespread haplotype 7 (Fig. ...
Context 9
... of the parents apparently yield extensive hybrid swarms, but, even when numbers of flowering plants of both parents are small, hybrids can result. For example, at Faversham in Kent, typically a dozen flowering plants of O. simia grew alongside an even smaller number of O. anthropophora, yet an F1 plant was generated (Bateman & Farrington, 1987) (Fig. 1G, H). Our results for AFLP analysis, and especially ITS alleles and plastid haplotypes, all suggest that hybridization progressed beyond the F1 generation and that the anthropomorphic Orchis species thereby remain evolutionarily interconnected by significant gene ...
Context 11
... WAS ORCHIS SIMIA The morphometric analyses (Figs 6, 7), AFLP study ( Fig. 8), ITS sequences ( Fig. 9) and microsatellite data (Fig. 10, Table 2) all show that the study plants from Goring were correctly identified in the field as hybrids between O. purpurea and O. simia; this is the first time that this hybrid combination has been formally recorded in Britain (cf. Stace, ...
Context 12
... purpurea and most floral characters are either intermediate between the parents or closer to O. purpurea; they more closely resemble O. simia only in the relatively long legs, inrolled (welcoming) arms, fewer, larger clusters of papillae on the chest, dark-coloured limbs and pale purple (rather than dark red) abaxial surfaces to the sepals (cf. Fig. 1B-E; Table 1). Similarly, the first axis of the multivariate ordination of hybrids plus parents places the hybrid cluster significantly closer to O. purpurea than to O. simia, although the weaker second axis has the converse effect (Fig. ...
Context 13
... inference of maternity. Plastids are typically, and perhaps universally, maternally inherited in orchids (Corriveau & Coleman, 1988; Cafasso, Widmer & ). Those of the hybrids yielded haplotypes that matched those of the two co-occurring plants of O. purpurea, which share a haplotype that is rare in populations of this species in England (Fig. 10). Predictably, the hybrids contains ITS alleles characteristic of both parental species at the Hartslock site, although the copy paternally inherited from O. simia is the less common of the two alleles found in the population (Table 2). Admittedly, the AFLP profiles of the hybrids show greater similarity to O. simia than to O. purpurea; ...
Context 14
... addition, the F1 hybrid plants are sufficiently similar in appearance to each other (Figs 1, 2) to suggest that they resulted from a single pollination event between the two mis-matched parents, most likely representing a lapse of concentration on the part of a passing bee engaged in a (fruitless) search for nectar. Genetic similarity among the hybrids is further suggested by tight clustering of their AFLP profiles (Fig. 8) and the uniformity of their ITS and plastid genotypes (Figs 9, 10). ...
Context 15
... addition, the F1 hybrid plants are sufficiently similar in appearance to each other (Figs 1, 2) to suggest that they resulted from a single pollination event between the two mis-matched parents, most likely representing a lapse of concentration on the part of a passing bee engaged in a (fruitless) search for nectar. Genetic similarity among the hybrids is further suggested by tight clustering of their AFLP profiles (Fig. 8) and the uniformity of their ITS and plastid genotypes (Figs 9, 10). Cross-fertilization most likely occurred within four years of O. purpurea first flowering at Goring in 1999, given that the closely related O. militaris can reach flowering size from seed in two years under in vitro cultivation (R. Manuel, pers. ...
Context 16
... leaf tips from each of these localities were included in the present molecular analysis. Plants from both sites proved identical and had the commonest ITS and plastid genotypes found NEW HYBRID ORCHID 703 in the species in England, but neither population showed a close genetic similarity to the two plants of O. purpurea sampled from Goring (Figs 8-10). Thus, we reject Hypothesis 2. ...
Context 17
... TO ORCHIS MILITARIS? While admitting that there was clearly considerable confusion during the 18 th and 19 th centuries, our current knowledge of the appearance of the anthropomorphic species means that few experienced field botanists would confuse O. simia with O. purpurea (cf. Fig. 1D, E). Rather, it is the partial morphological intermediacy of O. militaris (Fig. 1F) relative to the other two species that continues to cause identification problems (note that Linnaeus, and many later botanists, combined the three species under a more broadly circumscribed O. militaris L.). Indeed, this morphological intermediacy causes ...
Context 18
... While admitting that there was clearly considerable confusion during the 18 th and 19 th centuries, our current knowledge of the appearance of the anthropomorphic species means that few experienced field botanists would confuse O. simia with O. purpurea (cf. Fig. 1D, E). Rather, it is the partial morphological intermediacy of O. militaris (Fig. 1F) relative to the other two species that continues to cause identification problems (note that Linnaeus, and many later botanists, combined the three species under a more broadly circumscribed O. militaris L.). Indeed, this morphological intermediacy causes greater confusion in parts of mainland Europe, notably France and the Low ...
Context 19
... the tendency of some initial observers of the Goring hybrids to view them as O. militaris, if one compares the two taxa in sufficient detail, several significant differences emerge (cf. Fig. 1B, C vs. 1F; Table 1). The limbs of the hybrids are more incurved, longer and narrower than those of O. militaris (rendering indices 'k' and 'l' especially diagnostic), although the torso is somewhat shorter; also, the tails of the hybrids are on average twice as long. The non-labellar perianth segments are shorter and slightly narrower, ...
Context 20
... Orchis species in Britain by studying vast numbers of (mostly 19 th century) pressed specimens held in British herbaria. Many herbaria are rich in specimens of O. simia from the Thames Valley (a significant proportion labelled 'Goring') and many of those specimens are far larger and more impressive than the plants that flower there today (Fig. 11A). Aided by conservation measures, the Goring population is large and expanding at present. However, it recovered from near-extirpation during the period 1950-1970(Paul, 1965Harrap & Harrap, 2005), which undoubtedly constituted a classic genetic bottleneck Qamaruz-Zaman et al., 1998) (Table 3). This observation has led to credible ...
Context 21
... in the Goring plants relative to those from Kent is readily observed in Table 1 and Figures 6 and 7. Moreover, even brief scrutiny of herbarium labels demonstrates that O. militaris, now confined in the Chiltern Hills to two sites east of Goring, once occurred far more frequently along the Thames Valley, extending from Pangbourne in the west (Fig. 11B) to the border of Hertfordshire and Middlesex in the east. Orchis simia similarly extended further east, these wider distributions thereby offering much greater potential for hybridization between the two species. Sheets of O. simia and O. purpurea held in the herbaria at BM and K consistently appear correctly identified, whereas a ...
Context 22
... particular interest is a herbarium sheet at K that was collected from 'Hartslock Wood, Goring' in May 1831. It bears single specimens of O. simia, O. militaris and a reasonably convincing hybrid between them (Fig. 11B), suggesting that introgression between anthropomorphic orchids at Hartslock is by no means a recent phenomenon, and perhaps helping to explain the complexity of the genetic profiles shown in Britain by both O. simia and O. purpurea. One possible interpretation of the ITS data ( Fig. 9) is that most of the remaining British, and many of ...
Similar publications
3. Подсем. EPIDENDROIDEAE (примитивные трибы – Neottieae, Vanilleae, Gastrodieae, Nervilieae) Summary. The article continues serial publication of illustrated critical taxonomical survey of orchids in the flora of Vietnam. This part of the monograph includes taxonomical treatment of so-called primitive tribes (Neottieae, Vanilleae, Gastrodieae, Ner...
Species of subtribe Goodyerinae (Orchidaceae, Orchidoideae) in the flora of Vietnam have been the subject of a series of taxonomic treatments (Seidenfaden 1992, Averyanov 1994, 2008, Averyanov & Averyanova 2003, Nguyen, Averyanov & Duong 2005). Nonetheless, the inventory of this group remains incomplete due to their sporadic distribution, rarity in...
Continuing herbarium and literature studies of orchids belonging to Subtribe Goodyerinae has revealed a variety of new and noteworthy data. Aside from first records and additional synonymy the following new combinations and species are proposed, viz. Anoectochilus dewildeorum, A. falconis, Aspidogyne gigantea, Goodyera sumbawana, Odontochilus asrao...
In 2013, Caetano and colleagues published two lists of the Orchidaceae of the Municipality of Benedito Novo, Santa Catarina, totalling 99 species. Between January 2014 and September 2016, other field trips were made and new species were found in the region. Thus, in this paper an updated checklist of the Orchidaceae of Benedito Novo is presented ba...
A new species of Phragmipedium, P. andreettae, is described and illustrated from northwestern Ecuador and its affinities are discussed. The new species is closely allied to P. fischeri, from which it differs by the narrow leaves, the pale rose to white flowers, the elliptic to obovate petals with reflexed margins, the reflexed margin of the dorsal...
Citations
... 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. ...
... Characterising morphological variation in bee orchids as near-continuous and exceptionally multidimensional encourages us to compare our study of Ophrys with those conducted by us previously on other genera of European orchids [45,66,69,76,77,[79][80][81][82][83]. ...
... In addition, much of the variation tends to be encompassed by the first coordinate, creating an unusually large differential between the first coordinate and the second. Studies of genera that contain a mixture of species separated by morphological discontinuities and species possessing overlapping morphologies, such as Himantoglossum [6,82] and the anthropomorphic subgenus of Orchis [76,81,90], capture 40-50% of the total variance in the first two coordinates. But taxonomic groups in which none of the species are separated by morphological discontinuities yield plots that capture only approximately 30% of the total variation in the first two coordinates. ...
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.
... This phenomenon, which is caused by absence of strong interspecific reproductive barriers and hybrid zones, has also been found in other orchid genera, such as Orchis Tourn. ex L. (Bateman et al., 2008), Ophrys L. (Cortis et al., 2009), Epidendrum Pav. ex Lindl (Pinheiro et al., 2010), and Paphiopedilum Pfitzer (Guo et al., 2015). ...
Cymbidium, which includes approximately 80 species, is one of the most ornamental and cultivated orchid genera. However, a lack of markers and sparse sampling have posed great challenges to resolving the phylogenetic relationships within the genus. In the present study, we reconstructed the phylogenetic relationships by utilizing one nuclear DNA (nrITS) and seven plastid genes (rbcL, trnS, trnG, matK, trnL, psbA, and atpI) from 70 species (varieties) in Cymbidium. We also examined the occurrence of phylogenetic conflict between nuclear (nrITS) and plastid loci and investigated how phylogenetic conflict bears on taxonomic classification within the genus. We found that phylogenetic conflict and low support values may be explained by hybridization and a lack of informative characteristics. Our results do not support previous classification of the subgenera and sections within Cymbidium. Discordance between gene trees and network analysis indicate that reticulate evolution occurred in the genus Cymbidium. Overall, our study indicates that Cymbidium has undergone a complex evolution.
... A range of molecular markers have been used to study genetic admixture in Orchis, and few studies present evidence of hybridization beyond the first generation (Cozzolino and Widmer, 2005). Analyses of various Orchis hybrids, e.g., between O. anthropophora and O. italica (Pellegrino et al., 2009), O. purpurea and O. simia (Bateman et al., 2008) and between O. mascula and O. provincialis (Pellegrino et al., 2005) have shown that most putative hybrids belong to the F1 generation. ...
... To date, only two hybrid zones have been studied in this group of four species: one O. purpurea-O. simia hybrid zone in the UK (Bateman et al., 2008) and an O. militaris-O. purpurea hybrid zone in Belgium (Jacquemyn et al., 2012a). ...
... function was used to calculate interspecific heterozygosity for all individuals in the simulated hybrid data set and in the real hybrid zones (Gompert and Buerkle, 2010 Geometric morphometric analysis of labellum shape Three labella were sampled at random from the inflorescence of each individual and manually scaled and landmarked using tpsDig v. 2.16 (Rohlf, 2010a, b). Geometric morphometric methods have not previously been used to study petal shape in Orchis, so the set of 15 homologous landmarks used in this study ( Figure 2) was chosen to capture the main aspects of the petal shape and to complement points that have been used for measurements in traditional morphometric analyses in Orchis (Bateman and Farrington, 1987;Cozzolino and Aceto, 1994;Bateman et al., 2008). Cross-validated discriminant function analysis was conducted in MorphoJ (Klingenberg, 2011) to test the strength of taxonomic assignment statistically using labellum shape, given their taxonomic classification using molecular data. ...
Premise:
The genetic structure of hybrid zones provides insight into the potential for gene flow to occur between plant taxa. Four closely related European orchid species (Orchis anthropophora, O. militaris, O. purpurea, and O. simia) hybridize when they co-occur. We aimed to characterize patterns of hybridization in O. militaris-O. purpurea, O. purpurea-O. simia, and O. anthropophora-O. simia hybrid zones using molecular and morphological data.
Methods:
We used 11 newly isolated nuclear microsatellites to genotype 695 individuals collected from seven hybrid zones and six allopatric parental populations in France. Geometric morphometric analysis was conducted using 15 labellum landmarks to capture the main aspects of petal shape.
Results:
Backcrossing was asymmetric toward O. militaris in multiple O. militaris-O. purpurea hybrid zones. Hybrids in O. purpurea-O. simia and O. anthropophora-O. simia hybrid zones were largely limited to F1 and F2 generations, but further admixture had occurred. These patterns were reflected in labellum geometric morphometric data, which correlated strongly with nuclear microsatellite data in all three species combinations.
Conclusions:
The coexistence of parental and admixed individuals in these Orchis hybrid zones implies they are likely to be tension zones being maintained by a balance between gene flow into the hybrid zone and selection acting against admixed individuals. The pattern of admixture in the three species combinations suggests intrinsic selection acting on the hybrids is weaker in more closely related taxa.
... Values are means and standard errors (SE) across loci. Population codes correspond to those given in Table 1 Population ( species and enable the identification of admixed individuals that are morphologically similar to pure species (Bateman et al. 2008). Such methods have been used to detect cryptic hybrids in other plant groups, such as Quercus (Dodd and Afzal-Rafii 2004), Pinus (Jasińska et al. 2010), andLomatia (McIntosh et al. 2014). ...
Hybridisation is a complex process that has important evolutionary consequences. In the case of rare species, a comprehensive understanding of inter-specific hybridisation can be critical for their conservation and management. Eucalyptus tetrapleura is a rare species of ironbark that is restricted to a 40 km × 100 km area around Grafton on the North Coast of New South Wales (Australia), and is distinctive in that it has four ribs on the sides of its buds and fruits. In recent years, central populations of E. tetrapleura have been cleared to facilitate upgrades to one of the major highways in eastern Australia. This has led to increased habitat fragmentation, and there are now concerns that the species is at risk of genetic swamping by more common ironbark relatives. In this study, we investigated the population genetics and patterns of gene flow in E. tetrapleura. We
used DArTseq to genotype samples collected from across the known distribution of E. tetrapleura, as well as leaf material collected from co-occurring ironbark species. We found that while E. tetrapleura was a distinct evolutionary lineage, there was evidence of gene flow between this species and other ironbarks. Furthermore, many populations that had been identified as E. tetrapleura on the basis of morphology were of hybrid origin, thus the range of the species was much smaller than previously thought. Overall, our findings demonstrate how genomic methods can improve our understanding of admixture across closely related lineages, which can be used to inform the restoration of rare species.
... On the other hand, the morphological variation evident between genetically identical Ophrys plants (Malmgren 2008;Bateman et al. 2011), and the strong tendency of European orchid hybrids to morphologically resemble their 'mother' (ovule parent) much more closely than their 'father' (pollen parent) (e.g. Bateman et al. 2008Bateman et al. , 2017, makes such misidentification all too likely for even the most experienced experts. Claims to the contrary (e.g. ...
Zusammenfassung/Summary: Bateman, R. M. (2018): Two bees or not two bees? An overview of Ophrys systematics.-Ber. Arbeitskrs. Heim. Orchid. 35(1): 5-46. Die Diskussion darüber, ob die europäische Gattung Ophrys neun phylogenetische Arten (plus zahlreiche Unterarten) oder mindestens 350 "ethologische" Arten umfasst, dauert an. Unglücklicherweise versäu-men es die meisten derjenigen Autoren, die eine Meinung zur vermutlichen Zahl der Ophrys-Arten äu-ßern, (1) ihr eigenes Art-Konzept zu spezifizieren oder (2) darzulegen, welche Summe an grundlegenden wissenschaftlichen Daten zu sammeln sei, ehe eine bestimmte Art als mehr denn eine typologische ex kathedra-Hypothese betrachtet werden kann, welche einer weiterführenden klassisch-wissenschaftlichen Prüfung bedarf. Ich bewerte im Folgenden DNA-basierte Beiträge sowohl zur phylogenetischen Rekon-struktion als auch zur Art-Beschreibung, indem ich sie mit Ergebnissen vergleiche, die gewonnen wur-den aus der Analyse von nrITS-Sequenzen, low-copy-nuclear-gene-Sequenzen sowie umfassenderen, technisch sehr herausfordernden Matrizen, die auf neueren Sequenz-Techniken der nächsten Generation basieren. Ich identifiziere den typologischen Ansatz (unter Berücksichtigung der Tatsache, dass nur begrenzt Pflanzenproben zur Verfügung stehen und präexistente taxonomischen Konzepten vorliegen) sowie einen dadurch inadäquat verengten Blick auf die evolutionären Mechanismen als gravierendstes von diversen Hindernissen für die biologisch einwandfreie Beschreibung von Arten, ganz gleich welcher Natur die gesammelten Daten sind (sofern überhaupt welche gesammelt wurden). Ich möchte betonen, dass sich genetische Daten, auch wenn sie auf typologische Proben begrenzt sind, von allen anderen Two bees, or not two bees-that is the question: Whether 'tis nobler in the mind to suffer The slings and arrows of outrageous concepts Or to take arms against a sea of species And by opposing, end them. [with apologies to William Shakespeare] 6 Ber. Arbeitskrs. Heim. Orchid. 35 (1): 2018 Daten dadurch unterscheiden, dass sie eine abstammungsadaptierte Interpretation ermöglichen. Obwohl Untersuchungen in kleinerem Maßstab, sofern sie verschiedene analytische Ansätze integrieren, der beste Weg sind, um potentielle Artbildungsmechanismen zu identifizieren, so ermöglichen doch erst DNA-basierte Untersuchungen in größerem Maßstab (idealerweise sollten sie überall verfügbar sein) die Etablierung eines soliden Fundamentes für eine optimale Artdefinition. Im Folgenden untersuche ich das Verhältnis von Umschreibung und Identifikation und behaupte, dass die konzeptuelle und/oder methodologische Selbstgefälligkeit einiger Taxonomen und die Subjektivität anderer das Haupthindernis für eine verbesserte Identifikation und somit aussagekräftige Zuordnung von Arten sowie von Taxa un-terhalb der Art-Ebene sind.
Debate continues regarding whether the European pseudocopulatory genus Ophrys contains nine phy-logenetic species (plus numerous subspecies), or at least 350 "ethological" species (or might even be "reconciled" to simultaneously maintaining both of these contrasting species concepts). Unfortunately, most commentators who offer an opinion about how many Ophrys species exist fail to specify (1) their preferred species concept or (2) what kind and amount of underlying scientific data should be gathered before a named species can be regarded as more than merely a typological, authoritarian hypothesis in need of subsequent scientific testing. I review DNA-based contributions to both phylogeny reconstruction and species circumscription, comparing results derived from nrITS sequences, low-copy nuclear gene sequences, and much larger but more technically challenging matrices obtained through newer next-generation sequencing techniques. I identify typology (reflecting limited sampling of plants and essentially pre-scientific taxonomic concepts) as the most serious of several barriers to the circumscription of biologically sound species, irrespective of the nature of the data gathered (if any). I emphasise that, even when limited to typological sampling, genetic data differ from other kinds of data in allowing interpretation of the recent history of the lineage that they represent. Although smaller-scale studies that integrate several analytical approaches are the best way to identify potential speciation mechanisms, large-scale (ideally universally available) DNA-based surveys are the optimal approach for circumscribing plant species. I explore the relationship between circumscription and identification, arguing that the conceptual and/or methodological complacency of some taxonomists and the subjectivity of others are the primary barrier to improved identification and thereby meaningful mapping of species.
... Clearing material visualized by DIC (differential interference contrast microscopy). Bateman & Farrington, 1987;Bateman & Hollingsworth, 2004;Bateman, Smith & Fay, 2008;R Bateman, pers. comm., 2017) indicated a strong asymmetry of phenotypically expressed inheritance of orchid hybrids relative to their parent. ...
... What is interesting is that all hybrid species from the above articles resembled their seed parent. One of the possible explanations of this phenomenon could be the influence of the cytoplasm on nuclear gene expression (Bateman, Smith & Fay, 2008). Secondly, multiple introgression into one parental line may bring hybrid generations reminiscent of this one parent (e.g., Pinheiro et al., 2010;Schilling, 2016 and references cited therein). ...
The first natural hybrid in the section Irapeana of the orchid genus Cypripedium is described and illustrated based on Guatemalan material. A molecular evaluation of the discovery is provided. Specimens with intermediate flowers between C. irapeanum and C. dickinsonianum within ITS and Xdh sequences have the signal sequence of both these species. The analysis of plastid sequences indicated that the maternal line is C. irapeanum . Information about the ecology, embryology and conservation status of the novelty is given, together with a distribution map of its parental species, C. irapeanum and C. dickinsonianum . A discussion of the hybridization between Cypripedium species is presented. The potential hybrid zones between the representatives of Cypripedium section Irapeana which were estimated based on the results of ecological niche modeling analysis are located in the Maya Highlands ( C. dickinsonianum and C. irapeanum ) and the eastern part of Southern Sierra Madre ( C. molle and C. irapeanum ). Moreover, all three Cypripedium species could inhabit Cordillera Neovolcánica according to the obtained models; however, it should be noticed that this region is well-distanced from the edges of the known geographical range of C. molle .
... Clearing material visualized by DIC (differential interference contrast microscopy). Bateman & Farrington, 1987;Bateman & Hollingsworth, 2004;Bateman, Smith & Fay, 2008;R Bateman, pers. comm., 2017) indicated a strong asymmetry of phenotypically expressed inheritance of orchid hybrids relative to their parent. ...
... What is interesting is that all hybrid species from the above articles resembled their seed parent. One of the possible explanations of this phenomenon could be the influence of the cytoplasm on nuclear gene expression ( Bateman, Smith & Fay, 2008). Secondly, multiple introgression into one parental line may bring hybrid generations reminiscent of this one parent (e.g., Pinheiro et al., 2010;Schilling, 2016 and references cited therein). ...
... The spread of D. praetermissa plants is greater on both PC1 and PC3, the latter providing partial separation of the three populations. The putative hybrid would from first principles be expected to occupy the morphological discontinuity (though not necessarily to be equidistant from the two clusters: Bateman & Farrington, 1987;Bateman & Hollingsworth, 2004;Bateman et al., 2008). However, the 'hybrid' plant was actually placed well within the cluster of D. praetemissa in general and within the Sawbridgeworth (i.e. ...
... Two aspects of this morphometric result are unprecedented in our experience (cf. Bateman & Farrington, 1987;Bateman & Hollingsworth, 2004;Bateman et al., 2008;Farrington & Bateman, 1989). First, the observed level of asymmetry of inheritance from 'mother' and 'father' appeared extraordinary, even to someone who has long argued for pre-eminence of maternal inheritance in plants (e.g. ...
... First, the observed level of asymmetry of inheritance from 'mother' and 'father' appeared extraordinary, even to someone who has long argued for pre-eminence of maternal inheritance in plants (e.g. Bateman & Hollingsworth, 2004;Bateman et al., 2008). It also contradicts a rather unwise statement made by Bateman in Stace et al. (2015: 337): 'Circumstantial evidence suggests that × Dactylodenia, like hybrids within Dactylorhiza, does not exhibit the strong asymmetry in heritability that favours the phenotype of the mother over that of the father, even though this phenomenon is evident in many other genera of tribe Orchideae.' ...
We describe the first reported hybrid to occur in nature between Dactylorhiza praetermissa and Gymnadenia borealis as × Dactylodenia lacerta R.M. Bateman & Tattersall. This vigorous plant was found and digitally imaged on a roadside verge of Crousa Downs on the Lizard Peninsula of Cornwall by Barry Tattersall in June, 2016, and tentatively identified as a hybrid new to science. However, detailed morphometric analysis using 40 characters and based partly on the initial images failed to confirm its hybrid origin, suggesting instead that it was a slightly deviant plant of D. praetermissa. Comparison of the hybrid with its putative parents by DNA sequencing of nuclear (ITS) and plastid (trnL-F) regions later confirmed the F1 hybrid nature of the plant, as well as identifying D. praetermissa as the hybrid’s ovule-parent and G. borealis as its pollen-parent. This result contrasted with prevailing theory that pollinators (in this case, most likely bumblebees) should preferentially transfer pollinaria from the shorter-spurred plant to the longer-spurred plant. The Crousa Downs plant is thus best described as a cryptic hybrid that shows strong maternal dominance in inheritance patterns, perhaps reflecting the fact that its ovule-parent is allotetraploid but its pollen-parent is diploid. A brief review of × Dactylodenia hybrids in the British Isles concludes that opportunities for the origin of this particular hybrid are severely limited by the contrasting habitat preferences and geographic distributions of its parents. An ongoing future for this apparently unique plant is unlikely due to local authorities’ inappropriately-timed mowing regimes.
... When two taxa are distinguished by only subtle phenotypic differences and/or character states that are not fixed in all individuals of each taxon, it becomes impossible to reliably identify hybrids between them using morphology. Any identification based on phenotypic characters requires the presence of a clear morphological discontinuity into which any primary hybrids will fall as a result of combining numerous characteristics of both parents (e.g., Bateman & Denholm, 1983;Bateman, Smith & Fay, 2008). ...
... Although artificial crossing has been used repeatedly to estimate comparative levels of post-zygotic reproductive isolation in European orchids (e.g., Scopece et al., 2007), the phenotypes presented by the resulting plants (e.g., Fig. 21) have thus far escaped serious discussion. Such experiments actually represent a golden opportunity to explore patterns of heritability of particular phenotypic features, seeking evidence of dominance, over-expression and linkage, as well as possibly epigenetic influences-phenomena that our experiences suggest more often nudge the phenotypes of primary hybrids toward the phenotype of the ovule parent rather than that of the pollen parent (e.g., Bateman, Smith & Fay, 2008). ...
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 we use detailed morphometric data obtained from the same populations to compare genotypes with associated phenotypes, in order to better explore taxonomic circumscription and character evolution within the clade.
Methods
Between one and 12 plants found in 25 populations that encompassed the entire distribution of the Himantoglossum s.l. clade were measured in situ for 51 morphological characters. Results for 45 of those characters were subjected to detailed multivariate and univariate analyses.
Key Results
Multivariate analyses readily separate subgenus Barlia and subgenus Comperia from subgenus Himantoglossum , and also the early-divergent H. formosum from the less divergent remainder of subgenus Himantoglossum . The sequence of divergence of these four lineages is confidently resolved. Our experimental approach to morphometric character analysis demonstrates clearly that phenotypic evolution within Himantoglossum is unusually multi-dimensional.
Conclusions
Degrees of divergence between taxa shown by morphological analyses approximate those previously shown using molecular analyses. Himantoglossum s.l . is readily divisible into three subgenera. The three sections of subgenus Himantoglossum — hircinum , caprinum and formosum— are arrayed from west to east with only limited geographical overlap. At this taxonomic level, their juxtaposition combines with conflict between contrasting datasets to complicate attempts to distinguish between clinal variation and the discontinuities that by definition separate bona fide species. All taxa achieve allogamy via food deceit and have only weak pollinator specificity. Artificial crossing demonstrates that intrinsic sterility barriers are weak. Although we have found evidence of gene flow among and within the three sections of subgenus Himantoglossum , reports of natural hybrids are surprisingly rare, probably because putative parents are sufficiently similar to questionably warrant the status of species. Phenological separation and increased xeromorphy characterise the origin of subgenus Barlia . Several individual morphological characters show evidence of parallel acquisition, and loss of features is especially frequent in floral markings among members of section caprinum . Detailed patterns of gain and loss demonstrate that several different categories of flower markings are inherited independently. Along with the dimensions of labellar lobes, these pigmentation characters have been over-emphasised in previous taxonomic treatments. Increased plant vigour was a crucial element of the origin of the genus, but vegetative characters underwent remarkably little subsequent evolution. Attempts to reconstruct hypothetical ancestors at internal nodes of the phylogeny are weakened by (a) uncertain placement of Steveniella as sister to Himantoglossum s.l. and (b) uncertain relationships among subtly different putative species within section caprinum . Nonetheless, heterochronic/allometric trends, ultimately limited by functional constraints, clearly dictate transitions between contrasting flower sizes and complex labellum shapes.
... Several recent attempts to reconstruct the relationship within Rosa using amplifi ed fragment length polymorphism (AFLP) (Vos et al. 1995) have yielded encouraging results and underscores the usefulness of this technique for resolving relationships in groups with complex evolutionary history (De Cock 2008, Koopman et al. 2008, De Riek et al. 2013. AFLPs have also been successfully applied to detect hybrid ancestry in many plant taxa (Bateman et al. 2008, Schulte et al. 2010, van Hengstum et al. 2012. ...
Rosa spinosissima is an endangered species in Norway, found only within a limited area on the southwestern coast. Presumed hybrid forms between R. spinosissima and rose species within R. sect. Caninae have been recorded from this area since the late 19th century. Here, analyses of such hybrid plants and selected populations of the tentative parent species were performed using AFLP markers, nuclear DNA content, pollen viability and seed germination rates, in addition to classic morphometric analysis. It is established that the hybrid rose represents a single hybrid taxon, viz Rosa × sabinii, formed by recurrent asymmetrical hybridization events between R. spinosissima and R. mollis, with the latter being the obligate ovule donor. The evidence presented does not indicate hybridization with other co-occurring rose species, or of introgression from R. mollis into R. spinosissima through backcrossing with R. ×sabinii.
