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This article examines the relationship between John Leigh (1812–1888) and Lydia Becker (1827–1890). Leigh was a prominent figure in the scientific circles of Manchester in the mid-nineteenth century and the city's Medical Officer for Health. Becker was a botanist and Leigh's second cousin. She corresponded with Charles Darwin and became a pioneer in the women's suffrage movement. Previous studies have argued that Leigh patronized and discouraged Becker's botanical interests. However, newly-discovered correspondence shows that Leigh respected her abilities and encouraged her development as a botanist, including attendance at the British Association for the Advancement of Science meetings where she presented one of the first scientific papers by a female botanist in Britain. While social and institutional norms in the Victorian era discouraged women from entering science, these norms could be transgressed in interactions involving specific individuals.

Burnout, mental health disorders and suicide are more common among doctors than the general population. Burnt-out doctors self-report increased rates of medical errors and the provision of suboptimal patient care. Surgeons in training are particularly at risk of burnout and are also less likely to seek professional support. Female surgical trainees have especially high rates of attrition, potentially because of issues surrounding childcare and motherhood. Several strategies to reduce burnout and promote resilience have been trialled among doctors. Schwartz rounds and mindfulness training have been shown to be effective, but only in those motivated to participate. A reduction in working hours has conflicting results, particularly among surgical trainees, which may be linked to the subsequent reduction in training opportunities, such as operative time and the ability to complete assessments. Early identification and targeted support of at-risk individuals is a potentially effective strategy that requires further research.

Polyploidy is widely considered as a major process in the evolution of plants but the accumulation of polyploid species diversity is still controversial. Some recent studies proposed increased extinction risk in neopolyploids compared with their diploid ancestors. The high proportion of polyploid ferns is expected to be formed mainly by neopolyploids, whereas paleopolyploid species are predicted to be clustered in clades founded by whole genome duplications. Here, we test this prediction by exploring the evolution of polyploidy in the derived fern family Aspleniaceae. The family has a global distribution and shows the highest frequency of polyploid taxa among all ferns. To test the hypothesis, we obtained a comprehensive phylogeny using chloroplast DNA sequences of 883 specimens representing 292 species. All published chromosome counts were mapped onto this phylogenetic framework in order to explore the evolution of polyploids. We recovered evidence for several whole genome duplications in the history of Aspleniaceae. Phylogenetic relationships of polyploids exceeding the tetraploid level suggest that tetraploid Asplenium species may have replaced their diploid ancestors as the main evolutionary players in some clades of this family.

The genome evolution of ferns has been considered to be relatively static compared with angiosperms. In this study, we analyse genome size data and chromosome numbers in a phylogenetic framework to explore three hypotheses: the correlation of genome size and chromosome number, the origin of modern ferns from ancestors with high chromosome numbers, and the occurrence of several whole-genome duplications during the evolution of ferns. To achieve this, we generated new genome size data, increasing the percentage of fern species with genome sizes estimated to 2.8% of extant diversity, and ensuring a comprehensive phylogenetic coverage including at least three species from each fern order. Genome size was correlated with chromosome number across all ferns despite some substantial variation in both traits. We observed a trend towards conservation of the amount of DNA per chromosome, although Osmundaceae and Psilotaceae have substantially larger chromosomes. Reconstruction of the ancestral genome traits suggested that the earliest ferns were already characterized by possessing high chromosome numbers and that the earliest divergences in ferns were correlated with substantial karyological changes. Evidence for repeated whole-genome duplications was found across the phylogeny. Fern genomes tend to evolve slowly, albeit genome rearrangements occur in some clades.

The phylogeny of 58 Pelargonium species was estimated using five plastid markers (rbcL, matK, ndhF, rpoC1, trnL-F) and one mitochondrial gene (nad5). The results confirmed the monophyly of three major clades and four subclades within Pelargonium but also indicate the need to revise some sectional classifications. This phylogeny was used to examine karyotype evolution in the genus: plotting chromosome sizes, numbers and 2C-values indicates that genome size is significantly correlated with chromosome size but not number. Accelerated rates of nucleotide substitution have been previously detected in both plastid and mitochondrial genes in Pelargonium, but sparse taxon sampling did not enable identification of the phylogenetic distribution of these elevated rates. Using the multigene phylogeny as a constraint, we investigated lineage- and locus-specific heterogeneity of substitution rates in Pelargonium for an expanded number of taxa and demonstrated that both plastid and mitochondrial genes have had accelerated substitution rates but with markedly disparate patterns. In the plastid, the exons of rpoC1 have significantly accelerated substitution rates compared to its intron and the acceleration was mainly due to nonsynonymous substitutions. In contrast, the mitochondrial gene, nad5, experienced substantial acceleration of synonymous substitution rates in three internal branches of Pelargonium, but this acceleration ceased in all terminal branches. Several lineages also have dN/dS ratios significantly greater than one for rpoC1, indicating that positive selection is acting on this gene, whereas the accelerated synonymous substitutions in the mitochondrial gene are the result of elevated mutation rates.

The need for action on the global environment is now well understood and governments, agencies, non-governmental organizations and botanic gardens have all been working in their various ways to promote environmental sustainability and reduce species and habitat loss for at least 10–20 years. The Global Strategy for Plant Conservation (GSPC) has been widely adopted, particularly by the botanic garden community, and has resulted in many successes despite failing to achieve its ultimate goal of halting the loss of plant biodiversity. The objectives and targets for Phase 2 of the GSPC, running from 2010 to 2020, mirror those of Phase 1 and had been largely agreed prior to their formal adoption at the Conference of the Parties to the Convention on Biological Diversity in Nagoya in October 2010. However, to be successful, the scientific contribution of botanic gardens needs to be strengthened, as does government policy and commitment. Botanic garden research to underpin conservation action, including the role of botanic garden horticulture, training and international capacity building, has a major part to play and needs to be better understood and better coordinated. We provide examples based on the experience of the Royal Botanic Garden Edinburgh in the UK and overseas. Government policy, at national and international levels, needs to reflect the fundamental importance of plant diversity in maintaining the biosphere and supporting humanity. The commitment of significant new resources is an essential prerequisite for success, but this needs to be well coordinated, inclusive of all stakeholders and carefully targeted. A further challenge is the need to integrate better the plant diversity-related activities of what are currently diverse and disconnected sectors, including agriculture, forestry, protected area management and botanic gardens.

We used morphological and geographical data of 128 herbarium specimens of the Taxus wallichiana complex in eastern Asia to investigate their utilization in discriminating and identifying taxa included in the complex. One bud scale and 26 leaf characters were used to separate T. fuana, T. wallichiana var. wallichiana, T. wallichiana var. mairei, T. wallichiana var. chinensis and T. sumatrana by K-means clustering and dendrograms using Ward's distance. Out of the 27 characters examined 21 were found to be well correlated with geographical patterns. T. fuana was morphologically the most distant taxon, while T. sumatrana clustered among the T. wallichiana varieties. After correcting misidentifications of the specimens in T. wallichiana, its varieties occupied discrete geographical ranges, except for some limited sympatry of varieties mairei and chinensis east of the Tanaka Kaiyong line and the Sichuan Basin, China. Our analysis demonstrates the importance of consistency in character selection and definition in the identification of morphologically difficult taxa and the power of combining morphometric and geographical data in clarifying their spatial distribution. (C) 2007 The Linnean Society of London, Botanical Journal of the Linnean Society, 2007, 155, 307-335.

We studied the phylogeography of Chinese yew (Taxus wallichiana), a tree species distributed over most of southern China and adjacent regions. A total of 1235 individuals from 50 populations from China and North Vietnam were analysed for chloroplast DNA variation using polymerase chain reaction-restriction fragment length polymorphism of the trnL-F intron-spacer region. A total of 19 different haplotypes were distinguished. We found a very high level of population differentiation and a strong phylogeographic pattern, suggesting low levels of recurrent gene flow among populations. Haplotype differentiation was most marked along the boundary between the Sino-Himalayan and Sino-Japanese Forest floristic subkingdoms, with only one haplotype being shared among these two subkingdoms. The Malesian and Sino-Himalayan Forest subkingdoms had five and 10 haplotypes, respectively, while the relatively large Sino-Japanese Forest subkingdom had only eight. The strong geography-haplotype correlation persisted at the regional floristic level, with most regions possessing a unique set of haplotypes, except for the central China region. Strong landscape effects were observed in the Hengduan and Dabashan mountains, where steep mountains and valleys might have been natural dispersal barriers. The molecular phylogenetic data, together with the geographic distribution of the haplotypes, suggest the existence of several localized refugia during the last glaciation from which the present-day distribution may be derived. The pattern of haplotype distribution across China and North Vietnam corresponded well with the current taxonomic delineation of the three intraspecific varieties of T. wallichiana.

The cladistic relationships of endemic Commidendrum (four species) and Melanodendron (one species) from St Helena were inferred from sequences of ITS1 and ITS2 of nuclear ribosomal DNA. Despite showing a range of morphological and ecological variation, the four species of Commidendrum, C. spurium, C. robustum, C. rotundifolium, and C. rugosum, form a closely related monophyletic group with percentage sequence divergence between 0.2 and 0.9%. Melanodendron integrifolium is sister to Commidendrum indicating that the two genera may have evolved from a common ancestor that arrived in St Helena via a single dispersal event. The closest relatives of Commidendrum and Melanodendron appear to be South African, in the predominantly shrubby genus Felicia, although further sampling of South African Astereae is required. We discuss the evolution and adaptive radiation of these rare and threatened species with particular reference to the possible role of heterochrony. © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society, 2004, 144, 69–83.

An RFLP analysis of chloroplast and nuclear DNA was followed in this taxonomic appraisal of section Ciconium. The section appears to be a monophyletic one, which divides into two groups, or sub-sections. The Zonal ‘Geranium’, P. × hortorum is derived from section Ciconium, and the species most likely to be its ancestors are shown to be P. inquinans and P. zonale. Maternal inheritance of chloroplasts was demonstrated by comparing chloroplast DNA from interspecific hybrids with their parents. These data also suggest that P. inquinans was probably the maternal parent of P. × hortorum in the original crosses. The ‘Ivy-leaved’ cultivars appear to be derived from P. peltatum, with some contribution from P. zonale and/or P. × hortorum.

Summary Although Scotland's vascular plant flora is depauperate in comparison with that of Mediterranean countries, it has a particularly rich bryophyte and lichen flora of international importance. It is a country with great habitat diversity including unique habitats like the woodlands and mountains of the west coast. Conservation of plant species and habitats in Scotland has been directed through targeted action plans since 1995. Analyses have been made of their effectiveness and of the research needs arising from these targets. Monitoring and survey work are essential for effective target setting, and for reviewing progress. Important cross-cutting issues include the science for conservation and genetic diversity, ecosystem function, the impact of non-native species and climate change. The Biodiversity Action Plan process has proved to be an effective tool. However, we also need to consider a broader spectrum of the flora, to include other key habitats or species with Scottish populations of internati...

Summary The Killarney Fern (Trichomanes speciosum) was considered amongst the rarest and most vulnerable of Europe's plants. Research which fed into the Biodiversity Action Plan and developed alongside it, has forced us to reconsider the level of threat to this species. A hitherto overlooked stage of the life-cycle, the gametophyte, has been shown to be comparatively abundant, genetically diverse, largely threat-free and with the potential over time to regenerate the familiar sporophyte stage. This research also promoted wider public interest which itself generated additional records, such that even the sporophyte is now known at more British sites than at any time previously and the species is accordingly considered ‘recovered’. However, continued legal protection and the need for further research are still recommended.

Phylogenetic studies using DNA sequences of two chloroplast regions, rbc L and trn L-F, demonstrate that the proposed genus Ceterach is a small clade within the large genus Asplenium , and sister to the Phyllitis clade. The Ceterach clade is characterised by irregular anastomosing veins and often densely scaled leaf blades. Its taxonomic status as a group nested within Asplenium is confirmed, and it is accepted here as a subgenus with seven species. The Ceterach clade comprises four lineages that correspond to disjunct polyploid complexes: the A. aureum clade forming a polyploid complex (4×, 6×, 8×) in Macaronesia, the A. ceterach clade forming a polyploid complex (2×, 4×, 6×) in the Mediterranean Basin, the A. paucivenosum clade (4×, 6×) in central Asia, and the A. dalhousiae clade (2×) with a disjunct distribution in the Himalaya, Yemen and Eritrea, and southwestern North America. Asplenium paucivenosum is sister to all other members of the Ceterach clade, whereas A. dalhousiae is sister to the A. aureum clade that includes tetraploid A. aureum , hexaploid A. lolegnamense , and octoploid A. parvifolium . Asplenium ceterach and its variations – including the hexaploid A. ceterach subsp. mediterraneum subsp. nov. first described below – form a monophyletic unit, sister to a clade consisting of A. aureum and A. dalhousiae. Asplenium cordatum from Africa and A. haugthonii from the isolated atlantic island of St. Helena are not members of the Ceterach clade, which suggests that leaf blades with dense indumenta have evolved at least twice within asplenioid ferns. The allotetraploid species A. hybridum has the chloroplast DNA from A. ceterach , and therefore the latter species is the maternal ancestor of the former. The other parent of this hybrid species is A. sagittatum that is nested within the sister clade of Ceterach , the Phyllitis clade comprising A. sagittatum and A. scolopendrium . The findings suggest that the current distribution of Ceterach is either the result of long-distance dispersal or represents fragmented relicts of a previously more widely distributed species.

A simple procedure for the amplification of chloroplast DNA from pollen is described. This allows the potential for biparental or paternal inheritance of chloroplasts in angiosperms to be established rapidly and reliably. Such information is important when chloroplast DNA, which is generally assumed to be inherited maternally, is used to study gene flow or reconstruct phylogenies. The technique could also prove valuable in assessing the risks of transgene escape via pollen of genetically modified plants, in which the chloroplast genome rather than the nucleus has been transformed.

Overall phylogenetic relationships within the genus Pelargonium (Geraniaceae) were inferred based on DNA sequences from mitochondrial(mt)-encoded nad1 b/c exons and from chloroplast(cp)-encoded trnL (UAA) 5 9 exon‐trnF (GAA) exon regions using two species of Geranium and Sarcocaulon vanderetiaeas outgroups. The group II intron between nad1 exons b and c was found to be absent from the Pelargonium, Geranium,and Sarcocaulon sequences presented here as well as from Erodium, which is the first recorded loss of this intron in angiosperms. Separate phylogenetic analyses of the mtDNA and cpDNA data sets produced largely congruent topologies, indicating linkage between mitochondrial and chloroplast genome inheritance. Simultaneous analysis of the combined data sets yielded a well-resolved topology with high clade support exhibiting a basic split into small and large chromosome species, the first group containing two lineages and the latter three. One large chromosome lineage (x 5 11) comprises species from sections Myrrhidium and Chorisma and is sister to a lineage comprising P. mutans (x 5 11) and species from section Jenkinsonia (x 5 9). Sister to these two lineages is a lineage comprising species from sections Ciconium (x 5 9) and Subsucculentia (x 5 10). Cladistic evaluation of this pattern suggests that x 5 11 is the ancestral basic chromosome number for the genus.

In a leaf phenolic survey of 58 Pelargonium taxa, quercetin was almost universally present. Tannins were major components, namely proanthocyanidins and ellagitannins (in 53% of taxa) and free ellagic acid (in 50%). Myricetin (in 38%), flavone C -glycosides (in 36%) and luteolin (in 49% of taxa) were other regular constituents. A correlation was found between the presence of myricetin and proanthocyanidins. Although no individual compound or group of compounds appears to be a useful taxonomic marker for sectional classification, the data do indicate which sections are homogeneous, e.g. Otidia and Jenkinsonia . The chemistry supports the removal of some taxa from section Ligularia to the new sections Chorisma and Subsucculentia but does not distinguish between the latter two groups and indicates that section Ligularia is still an unnatural assemblage. Similarly, the data support the separation of P. reniforme and related species from section Cortusina into the new section Reniformia but P. odoratissimum appears misplaced in the latter group. The data suggest the placement of P. rodneyanum in section Peristera with the other Australian species. From a phylogenetic point of view the presence of ellagitannins or absence of tannins may be considered a more advanced state than the presence of proanthocyanidins and myricetin in the genus Pelargonium . For example, the most primitive state was found in P. crispum (section Pelargonium ) and the most advanced in P. multibracteatum (section Ciconium ). The chemistry also lends support for the division of the genus into two cytological groups based on chromosome size, in that most ellagitannin-producing species have large chromosomes while most proanthocyanidin-producing species have small chromosomes. There is agreement with the cytological and molecular data in grouping P. exstipulatum with P. album, P. reniforme and P. sidoides .

The macromorphology, palynology, karyology, cpDNA trnL-F sequences and phenolic components of P appendiculatum (L f) Willd., P crassipes Harv., P fulgidum (L) L’ Herit, P. hirtum (Burm f) Jacq, P. hystrix Harv., P oreophilum Schltr., P pulchellum Sims, P sericifolium J.J.A v.d. Walt, P.stipulaceum (L f.) Willd and P. torulosum E M Marais were studied It is concluded that these 10 species are related and that they should be grouped in a single section

Patterns of substitution in chloroplast encoded trnL_F regions were compared between species of Actaea (Ranunculales), Digitalis (Scrophulariales), Drosera (Caryophyllales), Panicoideae (Poales), the small chromosome species clade of Pelargonium (Geraniales), each representing a different order of flowering plants, and Huperzia (Lycopodiales). In total, the study included 265 taxa, each with900-bp sequences, totaling 0.24 Mb. Both pairwise and phylogeny-based comparisons were used to assess nucleotide substitution patterns. In all six groups, we found that transition/transversion ratios, as estimated by maximum likelihood on most-parsimonious trees, ranged between 0.8 and 1.0 for ingroups. These values occurred both at low sequence divergences, where substitutional saturation, i.e., multiple substitutions having occurred at the same (homologous) nucleotide position, was not expected, and at higher levels of divergence. This suggests that the angiosperm trnL-F regions evolve in a pattern different from that generally observed for nuclear and animal mtDNA (transitional/transversion ratioor = 2). Transition/transversion ratios in the intron and the spacer region differed in all alignments compared, yet base compositions between the regions were highly similar in all six groups. A-T and G-C transversions were significantly less frequent than the other four substitution types. This correlates with results from studies on fidelity mechanisms in DNA replication that predict A-T and G-C transversions to be least likely to occur. It therefore strengthens confidence in the link between mutation bias at the polymerase level and the actual fixation of substitutions as recorded on evolutionary trees, and concomitantly, in the neutrality of nucleotide substitutions as phylogenetic markers.

The subsection Polyactium of the genus Pelargonium (section Polyactium ) is treated taxonomically in view of currently available information. All species are geophytes with massive, naked underground tubers. Seven species are recognized, which have almost identical flowers but differ greatly in respect of the morphology, texture, and indumentum of the leaves. All share a basic chromosome number of x = 11; but polyploidy of up to hexaploid level occurs, with each species tending to have a characteristic level of ploidy. Six of the species are restricted to the winter rainfall area, while the seventh occurs in the winter rainfall area but also extends quite far into the summer rainfall region.

Genetic diversity in the Killarney fern, Trichomanes speciosum Willd. has been investigated in south-western Scotland, the northern-most limit of the distribution of the sporophyte. T. speciosum is unique amongst European pteridophytes in that both phases of the life cycle are perennial and capable of vegetative propagation. Within sites no variation was revealed by allozyme electrophoresis, even where both generations were growing together. In contrast, diversity was observed among sites, with seven different multilocus phenotypes (MLPs) present in the area. Two of these MLPs covered large areas while the others were restricted to one, or few localities. Asexual reproduction of the gametophyte via gemmae is assumed to be the main means of dispersal in recent times, allowing single clones to become widespread, while the overall genetic variability may be attributed to sexual reproduction and spore dispersal in historic times under more favourable climatic conditions. We suggest that it is not inbreeding, nor lack of genetic variation that limits sporophyte production, but rather the prevailing climatic conditions. The sporophyte is extremely rare and vulnerable. However, when the gametophyte is considered, the species is neither threatened with extinction, nor does it appear to face the danger of marked genetic erosion, because the long-lived gametophyte stage contains all of the genetic variability present in the area and can be regarded as a valuable «seed-bank».

The Killarney fern Trichomanes speciosum Willd. (Hymenophyllaceae) is unique in possessing both extensive sexual (sporophyte and gametophyte generations present) and asexual (gametophyte only) ranges. It was first discovered in central Europe in 1993 and is represented in this area only by its perennial, vegetatively propagating gametophyte generation. Genetic variation has been investigated at 35 sites. Allozyme diversity is partitioned primarily between, not within, sites. Although genetic variation exists at a fine scale (lt 5 m) within some populations, the results suggest that clones were not intimately associated in these cases. The majority of sites support unique multilocus phenotypes. Where phenotypes were present at more than one site they tended to recur at the next closest site. However, similar phenotypes link eastern and western Pfalzerwald sites up to c. 70 km apart. This pattern of diversity suggests that colonisation was not solely of a “stepping stone” or “leading edge” type. We suggest that during a climatically favourable period, probably the Atlantic hypsithermal, there may have been an explosive colonisation by long-distance dispersal from refugial areas. This was followed by a short period during which sporophyte production, sexual reproduction and local spread were possible. With climatic change, reduction in the available habitat and the loss of the sporophyte generation, different individual genets became fixed within small, favourable, but scattered, sites. The possibility that some central European sites north of the Alps acted as periglacial refugia cannot be discounted, but would appear less likely than (re-) colonisation from the Atlantic fringe.

Length polymorphism in a non-coding spacer (trnLUAA-trnFGAA) in the chloroplast DNA was used in the investigation of the origin of the most common and conspicuous European fern hybrid, Asplenium x alternifolium (Aspleniaceae, Pteridophyta). The origins of A. x alternifolium, the hybrid between A. trichomanes s.l. and A. septentrionale s.l. was studied at three ploidy levels, diploid, triploid and tetraploid. The cpDNA technique allowed us to investigate the mode of hybrid formation between sexual species for the first time over a wide geographic range and with a large sample size. Morphological variation in this hybrid has previously been attributed to different reciprocal parental combinations, and to the influence of chloroplast genes on morphogenesis. Our results demonstrate that one parent, A. septentrionale s.l., acts predominantly as the female parent in these hybrids, with only one population of A. x alternifolium showing reciprocal hybridity. The discovery of predominantly unidirectional hybrid formation in this hybrid may be explained by the different breeding systems of the parental taxa. The role of gametophyte ecology is also assessed.

Phylogenetic analysis of nrDNA ITS and tmL (UAA) 5' exon-tmF (GAA) chloroplast DNA sequences from 17 species of Pelargonium sect. Peristera, together with nine putative outgroups, suggests paraphyly for the section and a close relationship between the highly disjurmt South African and Australian species of sect. Peristera. Representatives from Pelargonium sectt. Reniformia, Ligularia s. 1. and Isopetalum (the St. Helena endemic P. cotyledonis) appear to be nested within the Peristera clade. The dose relationship between the South African and Australian Peristera is interpreted as being caused by long-range dispersal to Australia, probably as recent as the late Pliocene

Chromosome numbers of 65 species of sect.Hoarea have been determined. These show three basic chromosome numbers, x = 11, 10 and 9. Only a few species are tetraploid. In five species both diploid and tetraploid cytotypes are reported. Several cases of deviations in chromosome numbers and cytological abnormalities were found, most of these being related to the presence of B chromosomes that occur in eight species. Evidence is presented to suggest that the basic chromosome numbers of x = 10 and x = 9 are derived from x = 11 by centric fusion. Although variation in basic chromosome number withinPelargonium has been the subject of detailed study, this is the first time that evidence has been found for a mechanism of change in basic number, that of centric fusion by Robertsonian translocation. For the species of sect.Hoarea with x = 9, where the evidence for Robertsonian translocation is greatest, this process has probably taken place quite recently. In contrast to results from other sections of the genusPelargonium, the three different basic numbers of sect.Hoarea do not contradict its delimitation as a natural taxon.

The presumed relationship between P. grandicalcaratum Knuth, P. karooicum Compton & Barnes, P. otaviense Knuth and P. spinosum Willd., based on karyological characteristics, is confirmed by macromorphological, palynological, rbc L gene sequence studies and an analysis of phenolic compounds. These four species form the new section Subsucculentia J.J.A. v.d. Walt. Taxonomic treatments of the section and species are presented.

The need for action on the global environment is now well understood and governments, agencies, non-governmental organizations and botanic gardens have all been working in their various ways to promote environmental sustainability and reduce species and habitat loss for at least 10–20 years. The Global Strategy for Plant Conservation (GSPC) has been widely adopted, particularly by the botanic garden community, and has resulted in many successes despite failing to achieve its ultimate goal of halting the loss of plant biodiversity. The objectives and targets for Phase 2 of the GSPC, running from 2010 to 2020, mirror those of Phase 1 and had been largely agreed prior to their formal adoption at the Conference of the Parties to the Convention on Biological Diversity in Nagoya in October 2010. However, to be successful, the scientific contribution of botanic gardens needs to be strengthened, as does government policy and commitment. Botanic garden research to underpin conservation action, including the role of botanic garden horticulture, training and international capacity building, has a major part to play and needs to be better understood and better coordinated. We provide examples based on the experience of the Royal Botanic Garden Edinburgh in the UK and overseas. Government policy, at national and international levels, needs to reflect the fundamental importance of plant diversity in maintaining the biosphere and supporting humanity. The commitment of significant new resources is an essential prerequisite for success, but this needs to be well coordinated, inclusive of all stakeholders and carefully targeted. A further challenge is the need to integrate better the plant diversity-related activities of what are currently diverse and disconnected sectors, including agriculture, forestry, protected area management and botanic gardens.

The pollen grains from a selection of species representing each of the recognised sections of Pelaronium and covering the geographical range of the genus were studied with a view to compiling keys and descriptions for their identification in the scanning electron microscope and compound light microscope. The pollen was examined and separated into pollen types representing the lowest recognisable taxonomic units on the basis of pollen morphological characteristics, principally exine structure and ornamentation. The greater majority of species were found to fall into a general striate/reticulate ornamentation type within which it was not possible to distinguish between species, although a number of other pollen types and subgroups could be identified by differences in their reticulum and ornamentation. The palynological results are discussed with respect to current systematic treatments of Pelargonium .

A study of 33 species ofPelargonium sect.Ligularia reveals four basic chromosome numbers, x = 8, 9, 10, and 11, and variation in chromosome size. From evidence of karyology and hybridization attempts, proposals are made to divide the section into smaller groups and to transfer some species to other sections.

Pelargonium otaviense Knuth andP. spinosum Willd. are excluded from sect.Glaucophyllum, whileP. grandiflorum (Andr.)Willd.,P. patulum Jacq. andP. tabulare (Burm. f.)L'Herit. of sect.Eumorpha are included. Sect.Glaucophyllum is characterized by green to glaucous vegetative organs and zygomorphic white to pink corolla with five narrow petals. All the species have an identical pollen and chromosome morphology, the same basic chromosome number (x = 11) and similar flavonoid patterns. A close relationship between sect.Glaucophyllum and sect.Pelargonium is indicated by the occurrence of natural hybrids and concordant characters. Isorhamnetin and luteolin have been detected in the genus for the first time.

Pelargonium sect.Ciconium and sect.Dibrachya have a basic chromosome number of x = 9, whereas sect.Jenkinsonia has x = 9, 11, and 17.

Morphology-based delimitation of genera in the Cheilanthoid ferns has proved to be problematic and understanding of the phylogeny and relationships amongst Cheilanthoid ferns based on morphological characters has posed even further difficulties, owing perhaps in large part to adaptation by many taxa to xeric habitats, as well as convergent evolution. It is only now with the application of DNA sequence data that relationships of species and genera are becoming clear. Here, we present results of cpDNA sequence data from species that have been traditionally placed in the genus Doryopteris and, based on both these results, and morphological and distribution data, this study helps clarify the concept of the genus Doryopteris its position within the Cheilanthoid ferns and the status of Lytoneuron. As a result, three genera are redefined: Doryopteris, Lytoneuron and Ormopteris.

Little is known about the mode of transmission of chloroplasts in ferns, despite the importance of such knowledge for molecular phylogenetic and biosystematic studies. Andersson-Kotto (1930, 1931) inferred from crossing experiments that chloroplasts are inherited biparentally in Asplenium scolopendrium L. Here we present evidence from artificial hybridisation that demonstrates maternal inheritance of chloroplast DNA in the genus Asplenium (Aspleniaceae, Pteridophyta) using length variation in a non-coding spacer (trnLUAA-trnFGAA) in the chloroplast DNA.

Allozyme polymorphism was used 1) to investigate the relationships of three threatened species of Elaphoglossum from St Helena, E. nervosum, E. bifurcatum and E. dimorphum, and 2) to estimate levels of genetic diversity and its partitioning among populations. Despite showing morphological and ecological variation, the three species are closely related with high genetic identities. Evidence from one enzyme locus (Mdh-1) suggests that E. dimorphum is of hybrid origin involving E. nervosum and E. bifurcatum. Levels of genetic diversity were low in the three species, but comparable with other insular endemic angiosperms. Populations of E. nervosum and E. bifurcatum showed significant genetic differentiation, which should be taken into account in any conservation programme.

The endemic elaphoglossoid ferns, Elaphoglossum dimorphum, E. nervosum and Microstaphyla furcata of St Helena, form a closely related group within section Lepidoglossa when analysed phylogenetically using sequences from the chloroplast trnL intron (partial) and trnL-F intergenic spacer. Microstaphyla furcata, traditionally placed in its own genus, is clearly shown to belong to Elaphoglossum confirming the previous transfer of this species to Elaphoglossum as E. bifurcatum. There is hardly any trnL-F sequence divergence between the species, in fact sequences of E. nervosum and E.dimorphum are identical. These results are consistent with the possible origin of E. dimorphum as a hybrid between E. bifurcatum and E. nervosum or with the view that the three species are the result of a recent radiation. The potential conflict between phylogenetic and morphological distinctness in determining species conservation priorities is discussed.

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Chloroplast DNA sequences were obtained from 331 Asplenium ceterach plants representing 143 populations from throughout the range of the complex in Europe, plus outlying sites in North Africa and the near East. We identified nine distinct haplotypes from a 900 bp fragment of trnL-trnF gene. Tetraploid populations were encountered throughout Europe and further afield, whereas diploid populations were scarcer and predominated in the Pannonian-Balkan region. Hexaploids were encountered only in southern Mediterranean populations. Four haplotypes were found among diploid populations of the Pannonian-Balkans indicating that this region formed a northern Pleistocene refugium. A separate polyploid complex centred on Greece, comprises diploid, tetraploid and hexaploid populations with two endemic haplotypes and suggests long-term persistence of populations in the southern Mediterranean. Three chloroplast DNA (cpDNA) haplotypes were common among tetraploids in Spain and Italy, with diversity reducing northwards suggesting expansion from the south after the Pleistocene. Our cpDNA and ploidy data indicate at least six independent origins of polyploids.

Phylogenetic analysis was performed of 921 positions of trnL (UAA) 5′ exon — trnF (GAA) exon chloroplast DNA regions from 68 representatives ofPelargonium sectt.Campylia, Cortusina, Glaucophyllum, Hoarea, Isopetalum, Ligularia, Otidia, Pelargonium, Peristera, Polyactium, andReniformia, together with five putative outgroup species from sectionsCiconium, Chorisma andJenkinsonia. The total data set therefore comprised 67.2 kb of DNA sequence. Two main ingroup clades were identified: one clade contains sectionsPeristera, Reniformia, andIsopetalum, the other contains sectionsCampylia, Cortusina, Glaucophyllum, Hoarea, Ligularia, Otidia, Pelargonium, Polyactium and two species currently grouped in sect.Peristera. Branching order among five main clades within the latter clade was not resolved. The trnL-F sequence data support monophyly only for sectionsReniformia andHoarea, the remainder of the currently recognized sections ofPelargonium being either paraphyletic or polyphyletic. The data further suggest that sect.Polyactium is diphyletic and that sect.Glaucophyllum is nested within sect.Pelargonium. One relatively derived clade, which represents half of the genus, contains predominantly geophytic and succulent species, occurring in the geographically restricted winter rainfall region of the South African Cape. This pattern is interpreted as reflecting explosive radiation, possibly as an adaptive response to recent aridification in the western Cape.

Phylogenetic hypotheses for the largely South African genus Pelargonium L'Hér. (Geraniaceae) were derived based on DNA sequence data from nuclear, chloroplast and mitochondrial encoded regions. The datasets were unequally represented and comprised cpDNA trnL-F sequences for 152 taxa, nrDNA ITS sequences for 55 taxa, and mtDNA nad1 b/c exons for 51 taxa. Phylogenetic hypotheses derived from the separate three datasets were overall congruent. A single hypothesis synthesising the information in the three datasets was constructed following a total evidence approach and implementing dataset specific stepmatrices in order to correct for substitution biases. Pelargonium was found to consist of five main clades, some with contrasting evolutionary patterns with respect to biogeographic distributions, dispersal capacity, pollination biology and karyological diversification. The five main clades are structured in two (subgeneric) clades that correlate with chromosome size. One of these clades includes a "winter rainfall clade" containing more than 70% of all currently described Pelargonium species, and all restricted to the South African Cape winter rainfall region. Apart from (woody) shrubs and small herbaceous rosette subshrubs, this clade comprises a large "xerophytic" clade including geophytes, stem and leaf succulents, harbouring in total almost half of the genus. This clade is considered to be the result of in situ proliferation, possibly in response to late-Miocene and Pliocene aridification events. Nested within it is a radiation comprising c. 80 species from the geophytic Pelargonium section Hoarea, all characterised by the possession of (a series of) tunicate tubers.

The gametophyte of most ferns is a small, thalloid, heart-shaped, short-lived structure. In some species of Trichomanes (Hymenophyllaceae), the gametophytes are filamentous and consist of a weft of inter-twining branched filaments. Such gametophytes are perennial and can reproduce asexually by means of gemmae. Independent colonies of gametophytes have been reported from North America (Farrar, 1967, 1992), Britain (Rumsey et al., 1990) and Central Europe (Vogel et al., 1993, Jerome and Rasbach, 1994). These colonies of filamentous gametophytes can exist as very small patches (

From studies in the field, botanic garden and herbarium we concluded that the following four species of Dryopteris grow on the Canary Islands: D. aemula (n= 41) so far found only on Gomera; D. pseudomas (n= 82, apogamous) similarly recorded only for Gomera; D. oligodonta (n= 41) the most common representative of the genus in the Islands is possibly an old Canary endemic related to some African species; D. guanchica a new tetraploid species (n= 82) hitherto confused with D. austriaca (D. dilatata). On four islands of the Azores the following were found: D. aemula (n= 41); D. azorica (n= 41); D. pseudomas (n= 82, apogamous); and D. crispifolia a new tetraploid species (n= 82) which is very distinct when living but in the dried pressed state is difficult to differentiate from D. austriaca and D. guanchica.

In an attempt to establish the origin of the North American tetraploid Dryopteris campyloptera, hybrids have been synthesized and the chromosome pairing at meiosis investigated. It seems probable that it is derived from two diploid species, D. assimilis and D. intermedia. Dryopteris austriaca, a tetraploid species from Europe, is derived from the same diploid species but it may well have had an independent origin.

The chromosome numbers of seven species ofPelargonium sect.Eumorpha have been determined from material of known wild origin, and karyotypic comparisons have been made. Within the section there is variation in basic chromosome number (x = 4, 8, 9, 11), variation in chromosome size, and two species have polyploid races. The three species with chromosome numbers based on x = 11 have the smallest chromosomes (1.0–1.5 µm); chromosomes are larger (1.0–3.0 µm) in the other species.P. elongatum has the lowest chromosome number in the genus (2n = 8).P. alchemilloides is exceptional in that it has four cytotypes, 2n = 16, 18, 34 and 36, and the form with 2n = 36 has large chromosomes (2.0–5.0 µm). Evidence from a synthesized hybrid suggests thatP. alchemilloides with 2n = 16 may be of polyploid origin. The three species based on x = 11 appear to be more closely related to species from other sections ofPelargonium that have the same basic chromosome number and small chromosome size, rather than to other species of sect.Eumorpha.