Richard Bateman

Royal Botanic Gardens, Kew · Jodrell Laboratory

Background and aims: Bee orchids (Ophrys) have become the most popular model system for studying reproduction via insect-mediated pseudo-copulation and for exploring the consequent, putatively adaptive, evolutionary radiations. However, despite intensive past research, both the phylogenetic structure and species diversity within the genus remain h...

The developmental morphology and genetics of the orchid flower is described in order to explore the evolutionary 'no man's land' that separates the Extended Synthesis from the Modern Synthesis. The gynostemium, ubiquitous among orchids and developed through congenital fusion (and dorsal suppression) of fertile reproductive organs, is an unbreakable...

Several recent palaeobotanical studies claim to have found and described pre-Cretaceous angiosperm macrofossils. With rare exceptions, these papers fail to define a flower, do not acknowledge that fossils require character-based rather than group-based classification, do not explicitly state which morphological features would unambiguously identify...

Plastid sequences have long dominated phylogeny reconstruction at all time depths, predicated on a usually untested assumption that they accurately represent the evolutionary histories of phenotypically circumscribed species. We combined detailed in situ morphometrics (124 plants) and whole-plastome sequencing through genome skimming (71 plants) in...

A large English population of the temperate tuberous Greater Butterfly-orchid, Platanthera chlorantha, was monitored through a 16-year period. Each June the number of flowering plants was counted and 60 flowering plants were measured in situ for four morphological traits, selected for both ease of measurement and their contrasting contributions to...

Between 1959 and 1988, three populations of purple-flowered terrestrial orchids attributable to Dacty-lorhiza subgenus Dactylorhiza were discovered in Canada. The populations at Timmins, Ontario, and St John's, New-foundland were strongly marked on both flowers and leaves, in contrast with the anthocyanin-deficient population at Tilt Cove, Newfound...

The intensively studied Eurasian orchid genus Dactylorhiza has become a model system for exploring allopolyploid evolution, yet determining the optimal circumscriptions of, and most appropriate ranks for, its constituent taxa remain highly controversial topics. Here, novel allozyme data and detailed morphometric data for 16 Scottish marsh-orchid po...

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 consider...

Cryptic species are organisms which look identical, but which represent distinct evolutionary lineages. They are an emerging trend in organismal biology across all groups, from flatworms, insects, amphibians, primates, to vascular plants. This book critically evaluates the phenomenon of cryptic species and demonstrates how they can play a valuable...

Rhizomorphic lycopsids constitute the most derived lycophyte clade and some of the world’s best-known plant fossils. Arboreous taxa within the clade evolved independently of other nonlycophyte trees. Their rootstocks (rhizomorphs) have distinctive morphologies and anatomies and unusually canalized ontogenies, leading to debates regarding homology a...

Prompted by concurrent completion of the latest plant atlas for Britain and Ireland, the orchid flora of these islands is reviewed in detail, focusing on 21st century progress in both systematics research and formal conservation categorisation under IUCN criteria. DNA-based phylogenies consistently circumscribe monophyletic groups that constitute o...

The first study of the taxonomically critical European orchid genus Dactylorhiza to use next-generation DNA sequencing generated the statistically best-supported reconstruction of its phylogeny to date. However, the two competing topologies obtained within the monophyletic Section Maculatae differed radically in the placement of the D. maculata s.s...

Although conceptual whole-plants are occasionally reconstructed, disarticulation means that palaeobotanical taxo-nomy remains primarily the preserve of fragmented organ-species, each bearing a Linnaean binomial. Thus, a "natural" taxon that would in any neobotanical classification bear a single valid binomial (any other epithets automatically being...

Despite limited exposures, detailed field mapping and sedimentological analyses – particle size, clast petrography and fine-sand mineralogy (plus limited coarse silt and clay mineralogy) – have elucidated the ages, provenance and depositional environments of the complex and controversial Quaternary deposits covering the Ayot Paleogene Outlier. Most...

Recent phylogenetic trees of the phenotypically diverse Eurasian orchid genus Gymnadenia s.l., based on Sanger sequencing of nrITS and next-generation RAD-seq data, largely agree on species boundaries but disagree radically regarding relationships among those species. Moreover, both contrasting topologies receive support from a recent transcriptome...

Circumscriptions of both the genus Gymnadenia and the 11–27 species that it contains are highly controversial. These Eurasian terrestrial orchids are nectar-rewarding and pollinated primarily by Lepidoptera. Opinions expressed on the number of species occurring in the British Isles range from one to four, though there exists broad agreement that at...

A much-debated population of Epipactis known since 1992 to occur at Whiteleaf in the Chiltern Hills of southern England was recently formally described as Epipactis neglecta var. collina by Kreutz et al. (2020). This taxonomic decision was based on a combination of traditional morphological descrip- tion plus the nuclear DNA-based analysis previous...

Paralycopodites Morey & Morey, a Carboniferous‐age arboreous lycopsid that grew in the tropical wetlands of Pangea, is the phylogenetically basalmost member of the Carboniferous stigmarian lycopsids to be conceptually reconstructed. We update its description through reciprocal illumination between anatomy (coal‐balls) and gross morphology (adpressi...

Recent application of next-generation sequencing technology to Eurasian taxa of the primitive epidendroid orchid clade Epipactis section Epipactis has further advanced our knowledge of what has become a model system for studying the origins of numerous autogamous taxa from within arguably only one allogamous ancestral species, E. helleborine s.s. C...

Poor morphological and molecular differentiation in recently diversified lineages is a widespread phenomenon in plants. Phylogenetic relationships within such species complexes are often difficult to resolve because of the low variability in traditional molecular loci. Furthermore, biological phenomena responsible for topological incongruence such...

Disentangling phylogenetic relationships proves challenging for groups that have evolved recently, especially if there is ongoing reticulation. Although they are in most cases immediately isolated from diploid relatives, sets of sibling allopolyploids often hybridize with each other, thereby increasing the complexity of an already challenging situa...

Background and aims: The terrestrial orchid genus Epipactis has become a model system for the study of speciation via transitions from allogamy to autogamy, but close phylogenetic relationships have proven difficult to resolve through Sanger sequencing. Methods: We analysed with RAD-seq 108 plants representing 29 named taxa that together span th...

Premise of research. The evolutionary origin of the seed habit coincided with profound physiological and structural changes associated with underlying developmental patterns. A coenocyte is formed during megagametophyte development in many vascular plants, including some lycophytes and all spermatophytes; this structure compares closely with simila...

Himantoglossum adriaticum H. Baumann is a long-lived perennial orchid with an adriato-mediterranean distribution. The species-level separation of this species from the more geographically widespread H. hircinum has only recently been confirmed via a combination of molecular and morphometric techniques, which are further developed here. To provide a...

Pulsatilla (Anemoneae, Ranunculaceae) is sister to Anemone s.s. and contains ca 40 perennial species of considerable horticultural and medical importance. We sequenced 31 of those species, plus nine subspecies, two cultivars and six outgroups, for two nuclear regions (high-copy nrITS and low-copy MLH1) and three plastid regions (rbcL, accD–psaI, tr...

Stomata play a critical ecological role as an interface between the plant and its environment. Although the guard‐cell pair is highly conserved in land plants, the development and patterning of surrounding epidermal cells follow predictable pathways in different taxa that are increasingly well understood following recent advances in the development...

The digitate-tubered clade (Dactylorhiza s.l. plus Gymnadenia s.l.) within subtribe Orchidinae is an important element of the North-temperate orchid flora and has become a model system for studying the genetic and epigenetic consequences of organism-wide ploidy change. Here, we integrate morphological phylogenetics with Sanger sequencing of nrITS a...

Poor morphological and molecular differentiation in recently diversified lineages is a widespread phenomenon in plants. Phylogenetic relationships within such species complexes are often difficult to resolve because of the low variability in traditional molecular loci, as well as various other biological phenomena responsible for topological incong...

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 ver...

Background and aims: Recent tissue-level observations made indirectly via flow cytometry suggest that endoreplication (duplication of the nuclear genome within the nuclear envelope in the absence of subsequent cell division) is widespread within the plant kingdom. Here, we also directly observe ploidy variation among cells within individual petals...

Two decades have passed since DNA evidence first demonstrated an intimate relationship between the circumboreal species 'Coeloglossum' viride and the temperate Eurasian genus Dactylorhiza s.s. Most subsequent molecular phylogenies showed 'C.' viride to diverge after the D. incarnata group. The law of monophyly therefore dictated inclusion in Dactyl...

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 identifi...

Although the majority of taxonomic studies of European orchids treat Pseudorchis as a monotypic genus, some observers have argued that the apparently circumboreal segregate P. straminea should also be treated as a full species. Here, we compare detailed in vivo measurements of 55 plants from nine populations of P. albida in Britain with ten plants...

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...

Premise of the study: Noeggerathiales are an extinct group of heterosporous shrubs and trees that were widespread and diverse during the Pennsylvanian-Permian Epochs (323-252 Ma) but are of controversial taxonomic affinity. Groups proposed as close relatives include leptosporangiate ferns, sphenopsids, progymnosperms, or the extant eusporangiate f...

Over the past decade, the various potential practical benefits offered by DNA ‘barcoding’ have been rehearsed ad nauseam, typically with at least modest exag- geration. However, definitions of barcoding, most of which remain rooted in the already passé holy grail of identifying the perfect genic region to characterize (and thereby identify) all lif...

Background and Aims. The largely Mississippian strata of the Kilpatrick Hills, located at the western end of the Scottish Midland Valley, enclose several macrofossil floras that together contain ca 21 organ-species of permineralised plants and ca 44 organ-species of compressed plants, here estimated to represent 25 whole-plant species (Glenarbuck =...

In most seed plants, cotyledons formed within the seed act as haustorial organs, as well as playing a key role in releasing the shoot apex from the seed coat on germination. Emergence of the shoot apex often results from asymmetric intercalary growth of the cotyledon bases. This process avoids the principal spatial constraint on germination in seed...

• Triassic and Jurassic fossils record structural changes in conifer seed cones through time, provide the earliest evidence for crown-group conifer clades, and further clarify sister-group relationships of modern conifer families. A new and distinct seed-cone from the Isle of Skye in western Scotland provides the oldest detailed evidence for the an...

Hybridization is a fundamental process in biology and can lead to new evolutionary lineages. However, if the parental taxa involved are rare, difficult decisi- ons may have to be made regarding the conservation of the biological process versus the conservation of the parental taxa. The genus Orchis in Europe is a good example of a group of species...

Background: Tribe Orchideae dominates the orchid flora of the temperate Northern Hemisphere but its representatives in East Asia had been subject to less intensive phylogenetic study than those in Eurasia and North America. Although this situation was improved recently by the molecular phylogenetic study of Jin et al., comparatively few species wer...

Background and Aims. This paper concludes our series of publications comparing island and mainland speciation in European butterfly-orchids, by studying the morphology, phylogenetics and reproductive biology of the controversial circum-arctic species Platanthera (Limnorchis) hyperborea—the most frequent of seven Icelandic orchids. We draw particula...

Opportunistic observations of the longest-standing British population of Orchis militaris L. suggest that bumblebees (Bombus spp.) may have become the primary pollinators of the orchid at the site. We document pollination by (female) workers of B. pratorum and males of the brood parasite B. vestalis. Placement of the pollinaria on the bodies of the...

Background and Aims Lizard orchids of the genus Himantoglossum include many of Eurasia's most spectacular orchids, producing substantial spikes of showy flowers. However, until recently the genus had received only limited, and entirely traditional, systematic study. The aim of the current work was to provide a more robust molecular phylogeny in ord...

• Premise of the study: Most orchid species native to the Macaronesian islands reflect immigration from western Europe or North Africa followed by anagenesis. The only putative exception is the butterfly orchids ( Platanthera ) of the Azores, where three species apparently reflect at least one cladogenetic speciation event. This multidisciplinary s...

Background and Aims. The Macaronesian islands represent an excellent crucible for exploring speciation. This dominantly phenotypic study complements a separate genotypic study, together designed to identify and circumscribe Platanthera species (butterfly-orchids) on the Azores, and to determine their geographic origin(s) and underlying speciation m...

The strong positive relationship evident between cell and genome size in both animals and plants forms the basis of using the size of stomatal guard cells as a proxy to track changes in plant genome size through geological time. We report for the first time a taxonomic fine‐scale investigation into changes in stomatal guard‐cell length and use thes...

• Premise of the study: Rhizomorphic lycopsids evolved the tree habit independently of all other land plants. Newly discovered specimens allow radical revision of our understanding of the growth architectures of the extinct Paleozoic sister-genera Synchysidendron and Diaphorodendron. • Methods: Detailed descriptions of six remarkable adpression spe...

Himantoglossum hircinum is one of the rarer and more charismatic orchids in the British flora. Morphometric comparison of the two largest and best-known populations in southern England – the coastal dune population at Sandwich and the chalk grassland population at Newmarket – using 46 characters showed that they differ only subtly, the Sandwich pla...

I. II. III. IV. V. VI. VII. VIII. References SUMMARY: We evaluate stomatal development in terms of its primary morphogenetic factors and place it in a phylogenetic context, including clarification of the contrasting specialist terms that are used by different sets of researchers. The genetic and structural bases for stomatal development are well co...

Background and Aims. The presence of novel structures in orchid flowers, including auricles, rostellum and bursicles on the gynostemium and a lobed labellum, has prompted long-standing homology disputes, fuelled by conflicting evidence from a wide range of sources. Re-assessment of this debate using an improved model is timely, following recent phy...

A new name, Himantoglossum jankae, is given to the widely recognised lizard orchid species that is distributed primarily in the Balkan Peninsula and the northwestern region of Asia Minor and has been erroneously named H. caprinum in most previous literature. The new species differs from its closest relatives in having the combination of relatively...

We consider the conceptual relationship between pollinator specfificity, genetic isolation and species delimitation, critiquing Darwin’s (1877) hypothesis that differential placement of pollinia on the probosces and eyes of sphingid moths explained the divergence of the Eurasian orchids Platanthera bifolia and P. chlorantha — species that have simi...

The Mediterranean orchid genus O phrys is remarkable for its pseudocopulatory pollination mechanism; naïve male pollinators are attracted to the flowers by olfactory, visual and tactile cues. The most striking visual cue is a highly reflective, blue speculum region at the centre of the labellum, which mimics the corresponding female insect and reac...

We present the first ultrastructural study of stomatal development in the “living fossil” gymnosperm species Ginkgo biloba. The Ginkgo lineage occupies a pivotal position in seed-plant phylogenies, but as most representatives are known only from fossils, the developmental pattern in G. biloba is critical for inferring patterns of stomatal evolution...

BACKGROUND AND AIMS: Patterns of ploidy variation among and within populations can provide valuable insights into the evolutionary mechanisms shaping the dynamics of plant systems showing ploidy diversity. Whereas data on majority ploidies are, by definition, often sufficiently extensive, much less is known about the incidence and evolutionary role...

There are two morphologically similar and closely related eastern species of lizard orchids of the Eurasian Himantoglossum Sprengel (1826: 694) that, according to current literature (Sundermann 1980, Buttler 1996, Kreutz 1998, Baumann et al. 2006, Delforge 2006), can easily be distinguished by anthocyanin markings on the labella (‘lips’) of their f...

The KNOTTED1-like homeobox (KNOX) genes are best known for maintaining a pluripotent stem-cell population in the shoot apical meristem that underlies indeterminate vegetative growth, allowing plants to adapt their development to suit the prevailing environmental conditions. More recently, the function of the KNOX gene family has been expanded to in...

Spasmodic progress in understanding the morphology of European allotetraploid marsh-orchids of the genus Dactylorhiza achieved in the second half of the twentieth century has in the twenty-first century been largely superseded by rapid progress in the molecular systematics of the group. However, a grand synthesis is still needed to integrate the re...

Effective species circumscription requires sampling as widely as possible in the eld, maximising numbers of individuals, populations, taxa and characters to be measured. Subsequent data analysis should be based on morphological similarity plus either molecularly-delimited monophyly or reproductive isolation, or prefer- ably both; congruence between...

Modern approaches to circumscribing genera and other higher taxa rely heavily on the reconstruction of phylogenies (evolutionary trees) from matrices of quantitative data. Although the number of characters included in DNA matrices is expanding rapidly as sequencing technology advances, increasing the number of plants analysed is at least as importa...

Detailed morphometric comparison of the three extant English populations of Orchis militaris with a population from the Vercors region of southern France uses both univariate and multivariate approaches. These analyses reveal only subtle distinctions of similar magnitude between the four populations, and slightly greater variation within the French...

Genetic and molecular studies have recently come to dominate botanical research at the expense of more traditional morphological approaches. This broad introduction to modern flower systematics demonstrates the great potential that floral morphology has to complement molecular data in phylogenetic and evolutionary investigations. Contributions from...

Relationships among allotetraploid marsh orchids collectively assignable to Dactylorhiza majalis s.l. are exceptionally complex because of multiple origins, secondary hybridization with each other, and with their parental lineages, local adaptation, and genomic reorganization processes, all extending through time and space. We sampled 34 population...

Spurs are tubular outgrowths of perianth organs that have evolved iteratively among angiosperms. They typically contain nectar and often strongly influence pollinator specificity, potentially mediating reproductive isolation. The identification of Antirrhinum majus mutants with ectopic petal spurs suggested that petal-spur development is dependent...

Orchid of Hungary. Morphology, ecology, distribution, reproduction and threat.

This subjective and highly personal commentary critically reviews the last three decades of research on British and Irish tetraploid marsh-orchids, with particular reference to narrow-leaved marsh-orchids of the D. traunsteineri aggregate. I emphasise how and why interpretations have shifted (sometimes radically) as sample sizes have increased and...

Climate change has shaped life in the past and will continue to do so in the future. Understanding the interactions between climate and biodiversity is a complex challenge to science. With contributions from 60 key researchers, this book examines the ongoing impact of climate change on the ecology and diversity of life on earth. It discusses the la...

The cDNA-AFLP data matrix and input file for SAM [58]. Excel file with one worksheet comprising the cDNA-AFLP binary data for diploid and polyploid individuals analysed here. A second worksheet includes the input for SAM [58].

Hybridization and polyploidy are potent forces that have regularly stimulated plant evolution and adaptation. Dactylorhiza majalis s.s., D. traunsteineri s.l. and D. ebudensis are three allopolyploid species of a polyploid complex formed through unidirectional (and, in the first two cases, recurrent) hybridization between the widespread diploids D....

We are surprised that our recent comparative sWe are surprised that our recent comparative study of the micromorphology of all species of the European orchid genus Ophrys L. (Bradshaw et al., 2010) has attracted wide-ranging criticism (Vereecken et al., 2011) from a well-established group of researchers who, for the sake of brevity, we will term ‘e...

An exceptionally wide range of systematic data has been accumulated on subtribe Orchidinae, which dominates the European orchid flora. Despite (or perhaps because of) this wealth of information, delimitation of genera within the subtribe continues to prompt extensive debate among taxonomists. Here, we use the controversial genus Orchis s.l. as a ca...

An exceptionally wide range of systematic data has been accumulated on subtribe Orchidinae, which dominates the European orchid flora. Despite (or perhaps because of) this wealth of information, delimitation of genera within the subtribe continues to prompt extensive debate among taxonomists. Here, we use the controversial genus Orchis s.l. as a ca...

Conifer cones are reproductive structures that are typically of restricted growth and either exclusively pollen-bearing (male) or exclusively ovule-bearing (female). Here, we review two common spontaneous developmental abnormalities of conifer cones: proliferated cones, in which the apex grows vegetatively, and bisexual cones, which possess both ma...

Terrestrial Orchids: From Seed to Mycotrophic Plant. RasmussenHanne N.. Cambridge: Cambridge University Press. 1995. xii + 444pp. ISBN 0 521 45165 5. £45.00 (US$64.95) (hardback). - Volume 54 Issue 3 - R. M. Bateman

Epigenetic information includes heritable signals that modulate gene expression but are not encoded in the primary nucleotide sequence. We have studied natural epigenetic variation in three allotetraploid sibling orchid species (Dactylorhiza majalis s.str, D. traunsteineri s.l., and D. ebudensis) that differ radically in geography/ecology. The epig...