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1.  Floral development of Hydrocera and Impatiens reveals evolutionary trends in the most early diverged lineages of the Balsaminaceae 
Annals of Botany  2012;109(7):1285-1296.
Background and Aims
Balsaminaceae consist of two genera, the monospecific Hydrocera and its species-rich sister Impatiens. Although both genera are seemingly rather similar in overall appearance, they differ in ecology, distribution range, habitat preference and morphology. Because morphological support for the current molecular phylogenetic hypothesis of Impatiens is low, a developmental study is necessary in order to obtain better insights into the evolutionary history of the family. Therefore, the floral development of H. triflora and I. omeiana was investigated, representing the most early-diverged lineage of Impatiens, and the observations were compared with the literature.
Flowers at all developmental stages were examined using scanning electron microscopy and light microscopy.
Key results
In Hydrocera, two whorls of five free perianth primordia develop into a less zygomorphic perianth compared with its sister genus. The androecial cap originates from five individual stamen primordia. Post-genital fusion of the upper parts of the filaments result in a filament ring below the anthers. The anthers fuse forming connivent anther-like units. The gynoecium of Hydrocera is pentamerous; it is largely synascidiate in early development. Only then is a symplicate zone formed resulting in style and stigmas. In I. omeiana, the perianth is formed as in Hydrocera. Five individual stamen primordia develop into five stamens, of which the upper part of the filaments converge with each other. The gynoecium of I. omeiana is tetramerous; it appears annular in early development.
Comparison of the present results with developmental data from the literature confirms the perianth morphocline hypothesis in which a congenital fusion of the parts of the perianth results in a shift from pentasepalous to trisepalous flowers. In addition, the development of the androecial cap and the gynoecium follows several distinct ontogenetic sequences within the family.
PMCID: PMC3359920  PMID: 22499855
Balsaminaceae; androecium; floral development; gynoecium; Hydrocera triflora; Impatiens omeiana
2.  The multiple fuzzy origins of woodiness within Balsaminaceae using an integrated approach. Where do we draw the line? 
Annals of Botany  2011;109(4):783-799.
Background and Aims
The family Balsaminaceae is essentially herbaceous, except for some woodier species that can be described as ‘woody’ herbs or small shrubs. The family is nested within the so-called balsaminoid clade of Ericales, including the exclusively woody families Tetrameristaceae and Marcgraviaceae, which is sister to the remaining families of the predominantly woody order. A molecular phylogeny of Balsaminaceae is compared with wood anatomical observations to find out whether the woodier species are derived from herbaceous taxa (i.e. secondary woodiness), or whether woodiness in the family represents the ancestral state for the order (i.e. primary woodiness).
Wood anatomical observations of 68 Impatiens species and Hydrocera triflora, of which 47 are included in a multigene phylogeny, are carried out using light and scanning electron microscopy and compared with the molecular phylogenetic insights.
Key Results
There is much continuous variation in wood development between the Impatiens species studied, making the distinction between herbaceousness and woodiness difficult. However, the most woody species, unambiguously considered as truly woody shrubs, all display paedomorphic wood features pointing to secondary woodiness. This hypothesis is further supported by the molecular phylogeny, demonstrating that these most woody species are derived from herbaceous (or less woody) species in at least five independent clades. Wood formation in H. triflora is mostly confined to the ribs of the stems and shows paedomorphic wood features as well, suggesting that the common ancestor of Balsaminaceae was probably herbaceous.
The terms ‘herbaceousness’ and ‘woodiness’ are notoriously difficult to use in Balsaminaceae. However, anatomical observations and molecular sequence data show that the woodier species are derived from less woody or clearly herbaceous species, demonstrating that secondary woodiness has evolved in parallel.
PMCID: PMC3286280  PMID: 22190560
Balsaminaceae; herbaceousness; Hydrocera; Impatiens; insular woodiness; light microscopy; primary woodiness; secondary woodiness; wood anatomy
3.  Symbiotic ß-Proteobacteria beyond Legumes: Burkholderia in Rubiaceae 
PLoS ONE  2013;8(1):e55260.
Symbiotic ß-proteobacteria not only occur in root nodules of legumes but are also found in leaves of certain Rubiaceae. The discovery of bacteria in plants formerly not implicated in endosymbiosis suggests a wider occurrence of plant-microbe interactions. Several ß-proteobacteria of the genus Burkholderia are detected in close association with tropical plants. This interaction has occurred three times independently, which suggest a recent and open plant-bacteria association. The presence or absence of Burkholderia endophytes is consistent on genus level and therefore implies a predictive value for the discovery of bacteria. Only a single Burkholderia species is found in association with a given plant species. However, the endophyte species are promiscuous and can be found in association with several plant species. Most of the endophytes are part of the plant-associated beneficial and environmental group, but others are closely related to B. glathei. This soil bacteria, together with related nodulating and non-nodulating endophytes, is therefore transferred to a newly defined and larger PBE group within the genus Burkholderia.
PMCID: PMC3555867  PMID: 23372845
4.  The need to re-investigate the nature of homoplastic characters: an ontogenetic case study of the ‘bracteoles’ in Atripliceae (Chenopodiaceae) 
Annals of Botany  2011;108(5):847-865.
Background and Aims
Within Chenopodioideae, Atripliceae have been distinguished by two bracteoles enveloping the female flowers/fruits, whereas in other tribes flowers are described as ebracteolate with persistent perianth. Molecular phylogenetic hypotheses suggest ‘bracteoles’ to be homoplastic. The origin of the bracteoles was explained by successive inflorescence reductions. Flower reduction was used to explain sex determination. Therefore, floral ontogeny was studied to evaluate the nature of the bracteoles and sex determination in Atripliceae.
Inflorescences of species of Atriplex, Chenopodium, Dysphania and Spinacia oleracea were investigated using light microscopy and scanning electron microscopy.
Key Results
The main axis of the inflorescence is indeterminate with elementary dichasia as lateral units. Flowers develop centripetally, with first the formation of a perianth primordium either from a ring primordium or from five individual tepal primordia fusing post-genitally. Subsequently, five stamen primordia originate, followed by the formation of an annular ovary primordium surrounding a central single ovule. Flowers are either initially hermaphroditic remaining bisexual and/or becoming functionally unisexual at later stages, or initially unisexual. In the studied species of Atriplex, female flowers are strictly female, except in A. hortensis. In Spinacia, female and male flowers are unisexual at all developmental stages. Female flowers of Atriplex and Spinacia are protected by two accrescent fused tepal lobes, whereas the other perianth members are absent.
In Atriplex and Spinacia modified structures around female flowers are not bracteoles, but two opposite accrescent tepal lobes, parts of a perianth persistent on the fruit. Flowers can achieve sexuality through many different combinations; they are initially hermaphroditic, subsequently developing into bisexual or functionally unisexual flowers, with the exception of Spinacia and strictly female flowers in Atriplex, which are unisexual from the earliest developmental stages. There may be a relationship between the formation of an annular perianth primordium and flexibility in floral sex determination.
PMCID: PMC3177680  PMID: 21852278
Atriplex; Atripliceae; bract/bracteole; Chenopodiaceae; Chenopodioideae; Chenopodium; Dysphania; floral ontogeny; floral sex determination; perianth modification; SEM/LM; Spinacia
5.  Endosymbiont Transmission Mode in Bacterial Leaf Nodulation as Revealed by a Population Genetic Study of Psychotria leptophylla 
Leaf-nodulated plants are colonized by vertically inherited bacterial endosymbionts, which maintain symbioses throughout host generations. The permanent character of the interaction implies phylogenetic congruence between the host and the endosymbiont. However, the present population genetic study of Psychotria leptophylla provides evidence for a mixed symbiont transmission involving both vertical inheritance and horizontal transfers from the environment.
PMCID: PMC3255619  PMID: 22038605
6.  Bacterial Leaf Symbiosis in Angiosperms: Host Specificity without Co-Speciation 
PLoS ONE  2011;6(9):e24430.
Bacterial leaf symbiosis is a unique and intimate interaction between bacteria and flowering plants, in which endosymbionts are organized in specialized leaf structures. Previously, bacterial leaf symbiosis has been described as a cyclic and obligate interaction in which the endosymbionts are vertically transmitted between plant generations and lack autonomous growth. Theoretically this allows for co-speciation between leaf nodulated plants and their endosymbionts. We sequenced the nodulated Burkholderia endosymbionts of 54 plant species from known leaf nodulated angiosperm genera, i.e. Ardisia, Pavetta, Psychotria and Sericanthe. Phylogenetic reconstruction of bacterial leaf symbionts and closely related free-living bacteria indicates the occurrence of multiple horizontal transfers of bacteria from the environment to leaf nodulated plant species. This rejects the hypothesis of a long co-speciation process between the bacterial endosymbionts and their host plants. Our results indicate a recent evolutionary process towards a stable and host specific interaction confirming the proposed maternal transmission mode of the endosymbionts through the seeds. Divergence estimates provide evidence for a relatively recent origin of bacterial leaf symbiosis, dating back to the Miocene (5–23 Mya). This geological epoch was characterized by cool and arid conditions, which may have triggered the origin of bacterial leaf symbiosis.
PMCID: PMC3168474  PMID: 21915326
7.  Correction: Endophytic Bacteria in Toxic South African Plants: Identification, Phylogeny and Possible Involvement in Gousiekte 
PLoS ONE  2011;6(6):10.1371/annotation/e7fa3cdf-c7bc-439f-83a2-36da63ef6c08.
PMCID: PMC3124557
8.  Endophytic Bacteria in Toxic South African Plants: Identification, Phylogeny and Possible Involvement in Gousiekte 
PLoS ONE  2011;6(4):e19265.
South African plant species of the genera Fadogia, Pavetta and Vangueria (all belonging to Rubiaceae) are known to cause gousiekte (literally ‘quick disease’), a fatal cardiotoxicosis of ruminants characterised by acute heart failure four to eight weeks after ingestion. Noteworthy is that all these plants harbour endophytes in their leaves: nodulating bacteria in specialized nodules in Pavetta and non-nodulating bacteria in the intercellular spaces between mesophyll cells in Fadogia and Vangueria.
Principal Findings
Isolation and analyses of these endophytes reveal the presence of Burkholderia bacteria in all the plant species implicated in gousiekte. Although the nodulating and non-nodulating bacteria belong to the same genus, they are phylogenetically not closely related and even fall in different bacterial clades. Pavetta harborii and Pavetta schumanniana have their own specific endophyte – Candidatus Burkholderia harborii and Candidatus Burkholderia schumanniana – while the non-nodulating bacteria found in the other gousiekte-inducing plants show high similarity to Burkholderia caledonica. In this group, the bacteria are host specific at population level. Investigation of gousiekte-inducing plants from other African countries resulted in the discovery of the same endophytes. Several other plants of the genera Afrocanthium, Canthium, Keetia, Psydrax, Pygmaeothamnus and Pyrostria were tested and were found to lack bacterial endophytes.
The discovery and identification of Burkholderia bacteria in gousiekte-inducing plants open new perspectives and opportunities for research not only into the cause of this economically important disease, but also into the evolution and functional significance of bacterial endosymbiosis in Rubiaceae. Other South African Rubiaceae that grow in the same area as the gousiekte-inducing plants were found to lack bacterial endophytes which suggests a link between bacteria and gousiekte. The same bacteria are consistently found in gousiekte-inducing plants from different regions indicating that these plants will also be toxic to ruminants in other African countries.
PMCID: PMC3082559  PMID: 21541284
9.  Spikelet structure and development in Cyperoideae (Cyperaceae): a monopodial general model based on ontogenetic evidence 
Annals of Botany  2010;105(4):555-571.
Background and Aims
In Cyperoideae, one of the two subfamilies in Cyperaceae, unresolved homology questions about spikelets remained. This was particularly the case in taxa with distichously organized spikelets and in Cariceae, a tribe with complex compound inflorescences comprising male (co)florescences and deciduous female single-flowered lateral spikelets. Using ontogenetic techniques, a wide range of taxa were investigated, including some controversial ones, in order to find morphological arguments to understand the nature of the spikelet in Cyperoideae. This paper presents a review of both new ontogenetic data and current knowledge, discussing a cyperoid, general, monopodial spikelet model.
Scanning electron microscopy and light microscopy were used to examine spikelets of 106 species from 33 cyperoid genera.
Ontogenetic data presented allow a consistent cyperoid spikelet model to be defined. Scanning and light microscopic images in controversial taxa such as Schoenus nigricans, Cariceae and Cypereae are interpreted accordingly.
Spikelets in all species studied consist of an indeterminate rachilla, and one to many spirally to distichously arranged glumes, each subtending a flower or empty. Lateral spikelets are subtended by a bract and have a spikelet prophyll. In distichously organized spikelets, combined concaulescence of the flowers and epicaulescence (a newly defined metatopic displacement) of the glumes has caused interpretational controversy in the past. In Cariceae, the male (co)florescences are terminal spikelets. Female single-flowered spikelets are positioned proximally on the rachis. To explain both this and the secondary spikelets in some Cypereae, the existence of an ontogenetic switch determining the development of a primordium into flower, or lateral axis is postulated.
PMCID: PMC2850794  PMID: 20197291
10.  Rate accelerations in nuclear 18S rDNA of mycoheterotrophic and parasitic angiosperms 
Journal of Plant Research  2010;124(5):561-576.
Rate variation in genes from all three genomes has been observed frequently in plant lineages with a parasitic and mycoheterotrophic mode of life. While the loss of photosynthetic ability leads to a relaxation of evolutionary constraints in genes involved in the photosynthetic apparatus, it remains to be determined how prevalent increased substitution rates are in nuclear DNA of non-photosynthetic angiosperms. In this study we infer rates of molecular evolution of 18S rDNA of all parasitic and mycoheterotorphic plant families (except Lauraceae and Polygalaceae) using relative rate tests. In several holoparasitic and mycoheterotrophic plant lineages extremely high substitution rates are observed compared to other photosynthetic angiosperms. The position and frequency of these substitutions have been identified to understand the mutation dynamics of 18S rRNA in achlorophyllous plants. Despite the presence of significantly elevated substitution rates, very few mutations occur in major functional and structural regions of the small ribosomal molecule, providing evidence that the efficiency of the translational apparatus in non-photosynthetic plants has not been affected.
PMCID: PMC3159761  PMID: 21188459
18S rDNA; Mycoheterotrophy; Parasitism; Substitution rates; Relative rate test
11.  Evolution of fruit and seed characters in the Diervilla and Lonicera clades (Caprifoliaceae, Dipsacales) 
Annals of Botany  2009;104(2):253-276.
Background and Aims
The Diervilla and Lonicera clades are members of the family Caprifoliaceae (Dipsacales sensu Donoghue et al., 2001, Harvard Papers in Botany 6: 459–479). So far, the intergeneric relationships of the Lonicera clade and the systematic position of Heptacodium remain equivocal. By studying fruit and seed morphology and anatomy, an attempt is made to clarify these issues. In addition, this study deals with the evolution of fruit and seed characters of the Diervilla and Lonicera clades with reference to allied taxa.
Light and scanning electron microscopy were used for the morphological and anatomical investigations. Phylogenetic analyses were carried out by applying the parsimony and Bayesian inference optimality criteria. Character evolution was studied by means of parsimony optimization and stochastic character mapping.
Key Results
Diervilla and Weigela (Diervilla clade) are characterized by several unique traits in Dipsacales, including capsules with numerous seeds, seed coats without sclerified outer tangential exotestal cell walls, and dehiscent fruits. Seeds with completely sclerified exotestal cells and fleshy fruits characterize the Lonicera clade. Leycesteria and Lonicera have berries, ovaries without sterile carpels and several seeds per locule, whereas Symphoricarpos and Triosteum have drupes, ovaries with one or two sterile carpels and a single seed per locule. Heptacodium shares several characteristics with members of the Linnina clade, e.g. achenes, single-seeded fruits and a compressed, parenchymatous seed coat.
The results confirm the monophyly of the Diervilla and Lonicera clades and allow us to hypothesize a close relationship between Leycesteria and Lonicera and between Symphoricarpos and Triosteum. Fruit and seed morphology and anatomy point to a sister relationship of Heptacodium with the Linnina clade, rather than with the Lonicera clade.
PMCID: PMC2710890  PMID: 19502353
Diervilla; Weigela; Symphoricarpos; Lonicera; Triosteum; Leycesteria; Heptacodium; Caprifoliaceae; Dipsacales; fruit; seed; evolution
12.  Woodiness within the Spermacoceae–Knoxieae alliance (Rubiaceae): retention of the basal woody condition in Rubiaceae or recent innovation? 
Annals of Botany  2009;103(7):1049-1064.
Background and Aims
The tribe Spermacoceae is essentially a herbaceous Rubiaceae lineage, except for some species that can be described as ‘woody’ herbs, small shrubs to treelets, or lianas. Its sister tribe Knoxieae contains a large number of herbaceous taxa, but the number of woody taxa is higher compared to Spermacoceae. The occurrence of herbaceous and woody species within the same group raises the question whether the woody taxa are derived from herbaceous taxa (i.e. secondary woodiness), or whether woodiness represents the ancestral state (i.e. primary woodiness). Microscopic observations of wood anatomy are combined with an independent molecular phylogeny to answer this question.
Observations of wood anatomy of 21 woody Spermacoceae and eight woody Knoxieae species, most of them included in a multi-gene molecular phylogeny, are carried out using light microscopy.
Key Results
Observations of wood anatomy in Spermacoceae support the molecular hypothesis that all the woody species examined are secondary derived. Well-known wood anatomical characters that demonstrate this shift from the herbaceous to the woody habit are the typically flat or decreasing length vs. age curves for vessel elements, the abundance of square and upright ray cells, or even the (near-) absence of rays. These so-called paedomorphic wood features are also present in the Knoxieae genera Otiophora, Otomeria, Pentas, Pentanisia and Phyllopentas. However, the wood structure of the other Knoxieae genera observed (Carphalea, Dirichletia and Triainolepis) is typical of primarily woody taxa.
In Spermacoceae, secondary woodiness has evolved numerous times in strikingly different habitats. In Knoxieae, there is a general trend from primary woodiness towards herbaceousness and back to (secondary) woodiness.
PMCID: PMC2707911  PMID: 19279041
Knoxieae; LM; primary woodiness; Rubiaceae; Rubioideae; secondary woodiness; Spermacoceae; wood anatomy
13.  Floral and Inflorescence Morphology and Ontogeny in Beta vulgaris, with Special Emphasis on the Ovary Position 
Annals of Botany  2008;102(4):643-651.
Background and Aims
In spite of recent phylogenetic analyses for the Chenopodiaceae–Amaranthaceae complex, some morphological characters are not unambiguously interpreted, which raises homology questions. Therefore, ontogenetic investigations, emphasizing on ‘bracteoles’ in Atripliceae and flowers in Chenopodioideae, were conducted. This first paper presents original ontogenetic observations in Beta vulgaris, which was chosen as a reference species for further comparative investigation because of its unclarified phylogenetic position and its flowers with a (semi-)inferior ovary, whereas all other Chenopodiaceae–Amaranthaceae have hypogynous flowers.
Inflorescences and flowers were examined using scanning electron microscopy and light microscopy.
Key Results
Floral development starts from an inflorescence unit primordium subtended by a lateral bract. This primordium develops into a determinate axis on which two opposite lateral flowers originate, each subtended by a bracteole. On a flower primordium, first five tepal primordia appear, followed by five opposite stamen primordia. Simultaneously, a convex floral apex appears, which differentiates into an annular ovary primordium with three stigma primordia, surrounding a central, single ovule. A floral tube, which raises the outer floral whorls, envelops the ovary, resulting in a semi-inferior ovary at mature stage. Similarly, a stamen tube is formed, raising the insertion points of the stamens, and forming a staminal ring, which does not contain stomata. During floral development, the calyces of the terminal flower and of one of the lateral flowers often fuse, forming a compound fruit structure.
In Beta vulgaris, the inflorescence is compound, consisting of an indeterminate main axis with many elementary dichasia as inflorescence units, of which the terminal flower and one lateral flower fuse at a later stage. Floral parts develop starting from the outer whorl towards the gynoecium. Because of the formation of an epigynous hypanthium, the ovary becomes semi-inferior in the course of floral development.
PMCID: PMC2701786  PMID: 18694878
Beta vulgaris; Chenopodiaceae; floral ontogeny; gynoecial development; epigynous hypanthium; semi-inferior ovary; inflorescence ontogeny; LM; SEM
14.  Diversification of myco-heterotrophic angiosperms: Evidence from Burmanniaceae 
Myco-heterotrophy evolved independently several times during angiosperm evolution. Although many species of myco-heterotrophic plants are highly endemic and long-distance dispersal seems unlikely, some genera are widely dispersed and have pantropical distributions, often with large disjunctions. Traditionally this has been interpreted as evidence for an old age of these taxa. However, due to their scarcity and highly reduced plastid genomes our understanding about the evolutionary histories of the angiosperm myco-heterotrophic groups is poor.
We provide a hypothesis for the diversification of the myco-heterotrophic family Burmanniaceae. Phylogenetic inference, combined with biogeographical analyses, molecular divergence time estimates, and diversification analyses suggest that Burmanniaceae originated in West Gondwana and started to diversify during the Late Cretaceous. Diversification and migration of the species-rich pantropical genera Burmannia and Gymnosiphon display congruent patterns. Diversification began during the Eocene, when global temperatures peaked and tropical forests occurred at low latitudes. Simultaneous migration from the New to the Old World in Burmannia and Gymnosiphon occurred via boreotropical migration routes. Subsequent Oligocene cooling and breakup of boreotropical flora ended New-Old World migration and caused a gradual decrease in diversification rate in Burmanniaceae.
Our results indicate that extant diversity and pantropical distribution of myco-heterotrophic Burmanniaceae is the result of diversification and boreotropical migration during the Eocene when tropical rain forest expanded dramatically.
PMCID: PMC2492876  PMID: 18573195
15.  Elaborate Petals in Australian Spermacoce (Rubiaceae) Species: Morphology, Ontogeny and Function 
Annals of Botany  2006;98(6):1167-1178.
• Background and Aims Australian Spermacoce species display various types of elaborate petals. Their precise morphology, ontogenetic origin, and function are hitherto unknown. The aim of the present paper is to unravel the development and nature of the diverse types of elaborate petals in Spermacoce through a floral ontogenetic study.
• Methods The floral ontogeny of six species characterized by different types of corolla appendages was studied by scanning electron microscopy and light microscopy. In order to elucidate the possible functions of the elaborate petals, field observations were conducted as well.
• Key Results Scanning electronmicrographs show that full-grown petals of Spermacoce lignosa, S. phaeosperma and S. redacta bear appendages on their ventral side. Despite their different appearance at anthesis, the appendages develop very similarly in all three species. They are initiated at the same developmental stage and are first visible as two arcs of primordia converging from the upper margins of the petal towards its midvein and downwards. In S. brevidens, S. caudata and S. erectiloba, the full-grown petals have two long, concave protuberances, which develop from the tissue at both sides of the petal's mid-vein. In these three species, initiation of appendages on the ventral side of the petals is also observed, but they are hardly visible on the mature petals. The two types of elaborate petals tightly enclose the anthers, both in bud and during most of the flowering period.
• Conclusions Among Australian Spermacoce species, two types of elaborate petals can be distinguished. The former hypothesis that the two types of elaborate petals are essentially homologous is here rejected. Field investigations point out that the elaborate petals might play a role in the pollination biology of the species.
PMCID: PMC3292270  PMID: 17028295
Australia; corolla appendages; elaborate petals; floral ontogeny; pollination; Rubiaceae; Spermacoce brevidens; S. caudata; S. erectiloba; S. lignosa; S. phaeosperma; S. redacta
16.  The Micromorphology of Pit Membranes in Tracheary Elements of Ericales: New Records of Tori or Pseudo-tori? 
Annals of Botany  2006;98(5):943-951.
• Background and Aims Intervascular pit membranes were examined within Ericales to determine the distribution and structure of torus-like thickenings.
• Methods Forty-nine species representing 12 families of the order Ericales were investigated using light, scanning and transmission electron microscopy. They were compared with four species of Oleaceae to determine the true nature of the thickenings.
• Key Results Pit membranes with torus-like thickenings were observed in seven species of Ericaceae and were found to be amorphous, plasmodesmata-associated structures with an irregular distribution. These pseudo-tori show major differences compared with true tori with respect to their distribution and ultrastructure. Genuine tori, which are strongly correlated with round pit apertures in narrow tracheary elements, were found in two species of Osmanthus (Oleaceae).
• Conclusions The pseudo-tori found in some Ericaceae are considered to be similar to pit membrane thickenings previously recorded in Rosaceae. While true tori appear to be functionally significant in terms of efficiency and safety of water transport, the possible function of pseudo-tori could be associated with the role of plasmodesmata during differentiation of tracheids, fibre-tracheids or narrow vessels.
PMCID: PMC3292242  PMID: 16935867
Ericaceae; Ericales; Oleaceae; pit membrane; plasmodesmata; pseudo-torus; torus; tracheary elements
17.  Palynological Variation in Balsaminoid Ericales. I. Marcgraviaceae 
Annals of Botany  2005;96(6):1047-1060.
• Background and Aims Marcgraviaceae are a rather small family of seven genera and approx. 130 neotropical species. This study aims to present a detailed palynological survey of the family in order to comment on the intrafamily relationships and possible correlations with pollinators.
• Methods In total, 119 specimens representing 67 species and all genera are observed using light microscopy and scanning electron microscopy. Furthermore, eight species from five genera are studied with transmission electron microscopy.
• Key Results Our results show that pollen grains of Marcgraviaceae are small (20–35 µm), have three equatorial apertures, granules on the colpus membrane, oblate spheroidal to prolate spheroidal shapes, mainly psilate to perforate ornamentations, and lalongate colpus-shaped thinnings at the inner layer of the exine, and show the presence of orbicules. Based on our fragmentary knowledge of the pollination biology of the family, there are no clear correlations between pollinators and pollen features.
• Conclusions The genus Marcgravia has a high percentage of reticulate sexine patterns and a relatively thin nexine. Sarcopera can be defined by the presence of an oblate spheroidal to even suboblate shape, while Ruyschia and Souroubea typically show prolate spheroidal to subprolate pollen grains. The presence of a thick foot layer in the pollen wall is characteristic of the genera Norantea, Sarcopera and Schwartzia. Pollen features that are taxonomically useful within the family are the shape, sexine sculpturing, and ultrastructure of the pollen wall.
PMCID: PMC4247093  PMID: 16157628
Balsaminoids; Ericales; Marcgraviaceae; neotropics; orbicules; palynology; SEM; TEM
18.  Palynological Variation in Balsaminoid Ericales. II. Balsaminaceae, Tetrameristaceae, Pellicieraceae and General Conclusions 
Annals of Botany  2005;96(6):1061-1073.
• Background and Aims The objective of this study is to examine the palynological diversity of Balsaminaceae (two genera/±1000 species), Tetrameristaceae (two genera/two species) and Pellicieraceae (one genus/one species). The diversity found will be used to infer the systematic value of pollen features within the balsaminoid clade.
• Methods Pollen morphology and ultrastructure of 29 species, representing all families of the balsaminoid clade except Marcgraviaceae, are investigated by means of light microscopy, scanning electron microscopy and transmission electron microscopy.
• Key Results Balsaminaceae pollen is small to medium sized with three to four apertures, which can be either colpate or porate, and a sexine sculpturing varying from coarsely reticulate to almost microreticulate. Tetrameristaceae pollen is small sized, 3-colporate, with a heterobrochate reticulate sculpturing and granules present in the lumina. Pellicieraceae pollen is large sized, 3-colporate with long ectocolpi and a perforate sexine sculpturing with large verrucae. Furthermore, Pelliciera is characterized by the occurrence of aggregated orbicules, while orbicules are completely absent in both Balsaminaceae and Tetrameristaceae. Balsaminaceae pollen differs from the other balsaminoid families due to the occurrence of colpate or porate grains with an oblate to peroblate shape, a very thin foot layer and a lamellated endexine.
• Conclusions From a pollen morphological point of view, Balsaminaceae are completely different from the other balsaminoid families. Therefore, no pollen morphological synapomorphies could be defined for the balsaminoid clade. However, various pollen features were observed that could indicate a possible relationship between Tetrameristaceae, Pellicieraceae and Marcgraviaceae. Despite the palynological similarities in the latter three families, it remains unclear to what extent they are related to each other.
PMCID: PMC4247094  PMID: 16157626
Balsaminaceae; Ericales; orbicules; Pellicieraceae; pollen; SEM; TEM; Tetrameristaceae
19.  Morphological and Ultrastructural Diversity of Orbicules in Gentianaceae 
Annals of Botany  2003;92(5):657-672.
Minute granules of sporopollenin, called orbicules, can be observed on the innermost tangential and/or radial walls of secretory tapetum cells. Orbicules were investigated in 53 species of 34 Gentianaceae genera using light microscopy, scanning electron microscopy and transmission electron microscopy. This selection covered all different tribes and subtribes recognized in Gentianaceae (87 genera, ±1650 species). Orbicules were found in 38 species (23 genera) distributed among the six tribes recognized in Gentianaceae. The orbicule typology is based on those described previously in Rubiaceae. Of the six orbicule types described previously, Type II orbicules are lacking. Type III orbicules are most common (17 species). Hockinia Gardner is the only representative with Type I orbicules. The number of representatives with orbicules belonging to the other orbicule types are equally distributed among the species studied: seven species possess Type IV orbicules, six species Type V and six species Type VI. The systematic usefulness of this typology is discussed in comparison with the latest systematic insights within the family, and palynological trends in Gentianaceae. Orbicule data have proven to be useful for evaluating tribal delimitation within Rubiaceae and Loganiaceae s.l.; however, they seem not to be useful for tribal delimitation in Gentianaceae. In the tribes Potalieae and Gentianeae orbicule data may be useful at subtribal level.
PMCID: PMC4244851  PMID: 14500324
Orbicules; Ubisch bodies; pollen; Gentianaceae; SEM; TEM; morphology; ultrastructure; typology; tapetum; systematics
20.  Comparative Wood Anatomy of Epacrids (Styphelioideae, Ericaceae s.l.) 
Annals of Botany  2003;91(7):835-856.
The wood anatomy of 16 of the 37 genera within the epacrids (Styphelioideae, Ericaceae s.l.) is investigated by light and scanning electron microscopy. Several features in the secondary xylem occur consistently at the tribal level: arrangement of vessel‐ray pits, distribution of axial parenchyma, ray width, and the presence and location of crystals. The primitive nature of Prionoteae and Archerieae is supported by the presence of scalariform perforation plates with many bars and scalariform to opposite vessel pitting. The wood structure of Oligarrheneae is similar to that of Styphelieae, but the very narrow vessel elements, exclusively uniseriate rays and the lack of prismatic crystals in Oligarrheneae distinguish these two tribes. The secondary xylem of Monotoca tamariscina indicates that it does not fit in Styphelieae; a position within Oligarrheneae is possible. Like most Cosmelieae, all Richeeae are characterized by exclusively scalariform perforation plates with many bars, a very high vessel density and paratracheal parenchyma, although they clearly differ in ray width (exclusively uniseriate rays in Cosmelieae vs. uniseriate and wide multiseriate rays in Richeeae). Several wood anatomical features confirm the inclusion of epacrids in Ericaceae s.l. Furthermore, there are significant ecological implications. The small vessel diameter and high vessel frequency in many epacrids are indicative of a high conductive safety to avoid embolism caused by freeze–thaw cycles, while the replacement of scalariform by simple vessel perforation plates and an increase in vessel diameter would suggest an increased conductive efficiency, which is especially found in mesic temperate or tropical Styphelieae.
PMCID: PMC4242392  PMID: 12770843
Ecological wood anatomy; epacrids; Ericaceae s.l.; Styphelioideae; systematic wood anatomy
21.  A Comparative Study of Metal Levels in Leaves of Some Al‐accumulating Rubiaceae 
Annals of Botany  2003;91(6):657-663.
Concentrations of Al, Si, Fe, Mn, Cu and Ca were analysed in leaves of ten Rubiaceae species, most of which are Al accumulators, and these were compared with concentrations in one species of Melastomataceae. Quantitative data confirmed the distribution of Al accumulation as previously determined by semi‐quantitative tests, and suggest that there is an apparent congruency between the shoot Al concentration and the number of accumulators within a certain genus or tribe. Al accumulators within the Rubiaceae are most characteristic of the Rubioideae subfamily, although a second origin is likely in at least a few members of the tribes Vanguerieae and Alberteae. While the leaf Si concentration in Melastomata malabathricum L. (Melastomataceae) was negligible, all Rubiaceae studied showed relatively high Si levels (mostly >3000 mg kg–1). It is hypothesized that an Al–Si complex is formed in the shoot tissues of Al‐accumulating Rubiaceae and that this may contribute to Al detoxification. However, the Si : Al mole ratio tended to differ widely among species. There was no significant correlation between Al and the other metals analysed. A remarkably high Mn concentration was found in Coptosapelta olaciformis Elm.
PMCID: PMC4242354  PMID: 12714364
Accumulation; aluminium; Melastomataceae; metals; phylogeny; Rubiaceae; silicon
22.  Morphological and Ultrastructural Diversity of Orbicules in Relation to Evolutionary Tendencies in Apocynaceae s.l. 
Annals of Botany  2002;90(5):647-662.
Minute granules of sporopollenin, called orbicules, can be observed on the innermost tangential and/or radial walls of secretory tapetum cells. Orbicules were investigated in 62 species (50 genera) of Apocynaceae s.l. using light microscopy, scanning electron microscopy and transmission electron microscopy. Orbicules were found in 43 species (34 genera) distributed amongst the subfamilies Rauvolfioideae, Apocynoideae, Periplocoideae, and in the genus Riocreuxia (Asclepiadoideae). Absence of orbicules is apparent in Secamonoideae and Asclepiadoideae (except Riocreuxia). The orbicule types described are based on observed morphological and ultrastructural variation. Of the six orbicule types previously described, Type I and Type II orbicules are lacking. In the majority of species, Type III orbicules were recorded in addition to Types IV, V and VI. In this study we suggest that embedded Type VI orbicules are more derived. A correlation between orbicule typology and evolutionary tendencies in Apocynaceae s.l. palynology was found. A trend was observed from the presence of Type III orbicules in the majority of species belonging to the basal group of genera characterized by colporate to porate single pollen grains, or 3–6‐porate tetrads, towards the more derived embedded Type VI orbicules in the more advanced Periplocoideae genera with multiporate tetrads or pollinia. Orbicule data have proven not to be useful for evaluating tribal delimitation within the Apocynaceae s.l. contrary to the Rubiaceae and Loganiaceae s.l.
PMCID: PMC4240458  PMID: 12466106
Orbicules; Ubisch bodies; Apocynaceae s.l.; SEM; TEM; morphology; ultrastructure; typology; tapetum; systematics
23.  Aluminium Accumulation in Leaves of 127 Species in Melastomataceae, with Comments on the Order Myrtales 
Annals of Botany  2002;90(1):53-64.
The distribution and systematic significance of aluminium accumulation is surveyed based on semi‐quantitative tests of 166 species, representing all tribes and subfamilies of the Melastomataceae as well as a few members of related families within the Myrtales. The character is strongly present in nearly all members of the Memecylaceae and in most primitive taxa of the Melastomataceae, while non‐accumulating taxa are widespread in the more derived tribes of the Melastomataceae. The variable distribution of Al accumulation in advanced clades of the latter family is probably associated with the tendency to herbaceousness, although it is unclear whether the more herbaceous representatives have developed more specialized Al‐response mechanisms that may exclude high Al levels from the shoot. It is hypothesized that Al accumulation is symplesiomorphic for Melastomataceae and Memecylaceae, and that the feature characterizes the most primitive families in the Myrtales. Indeed, Al accumulation is also characteristic of Crypteroniaceae, Rhynchocalycaceae and Vochysiaceae. Crypteroniaceae and Rhynchocalycaceae probably take a basal position in a sister clade of the Memecylaceae and Melastomataceae, while Al accumulation suggests a basal position for Vochysiaceae in the Myrtaceae clade.
PMCID: PMC4233848  PMID: 12125773
Aluminium accumulation; systematics; phylogenetic relationships; Crypteroniaceae; Melastomataceae; Memecylaceae; Myrtales; Rhynchocalycaceae; Vochysiaceae

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