Phyllanthus section Choretropsis (Euphorbiaceae) in Brazil

Section Choretropsis genus Phyllanthus (Euphorbiaceae) is formed by plants whose branches are modified into phylloclades. Despite the suitable characteristics of phylloclades within the Phyllanthus genus, the systematics of these species are poorly understood. Morphological data are presented here to allow future revision of this taxonomic group. The section is represented by nine species distributed in South America, eight of them endemic to Brazil. Our taxonomic studies distinguish the species and propose the synonymization of nine varieties of P. klotzschianus and two varieties of P. flagelliformis based on vegetative and reproductive characters. The diversity of branching and the morphology of phylloclades are discussed.  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 150 , 131–164.     

大戟科叶下珠属花粉形态的研究

对叶下珠属Phyllanthus Linn.13种植物花粉进行了光学显微镜和扫描电镜的观察。研究表明,本属花粉虽然在大小、形状和外壁纹饰等方面有不同程度的差异,但差异最大的还是在于萌发孔类型上。可以把本属花粉分为3孔沟,4－5孔沟,多孔沟,散沟（沟短而纺锤形或近圆形）和散孔等类型。这些萌发孔类型虽然与李秉滔在中国植物志第44卷中所划分的组不很吻合,但或多或少有一定的相关性,并且对于属下的分类具有一定的意义。在外壁纹饰方面,均为网状纹饰,可分为具粗网眼,细网眼和中等网眼等几种类型。但纹饰类型在组间或种间的差异不大明显。     

Pollen morphology of Microstachys (Euphorbiaceae) with emphasis on neotropical species

Abstract

In Euphorbiaceae tribe Hippomaneae is the pantropical genus Microstachys, comprised by 24 species, mostly from Brazil. Palynological studies with several representatives of the genus are scarce. The purpose of this study was to describe the pollen morphology of Microstachys and find possible differentiating characteristics between its species. Ten species of the genus were analysed, represented by 21 herbarium collections. Their pollen grains were acetolysed, measured and photographed under light microscopy (LM) and scanning electron microscopy (SEM). The analysed species possess pollen in monads, of small to medium size, isopolars, amb distincticly trilobed, oblate-spheroidal, prolate-spheroidal or subprolate, tricolporate with narrow and very long colpi, almost fused at the poles sometimes forming an invagination with tapering ends and margo of the colporus wide to very wide and psilate; endoapertures lalongate to very lalongate with tapering ends, presence of a narrow costae distinct in the endoaperture; exine microreticulate homobrochate, walls with simple columella; at the mesocolpus the sexine separates from the nexine forming cavea; and sexine always thicker than the nexine. Microstachys is a stenopalynous genus, despite its species varying in pollen size, shape and width of the cavea, margo of the colporus, and length of the endoapertures, demonstrating taxonomic value, and indicating a contribution towards future systematic analyses.     

Pollen morphology of the Swertiinae (Gentianaceae): phylogenetic implications

Recent molecular surveys of the Swertiinae (Gentianaceae–Gentianeae) revealed unexpected phylogenetic relationships, including polyphyly of the genera Gentianella , Jaeschkea , Lomatogonium and Swertia . To find new non-molecular characters supporting the phylogeny, we examined the exine variation of 73 species of all major lineages of subtribe Swertiinae using environmental scanning electron miscroscopy supplementing older, mainly light microscopical, studies. In contrast to previous studies, we were able to pick out taxa from phylogenetic key positions with particular focus on Swertia . Many distantly related taxa such as parts of Frasera , Gentianopsis , Halenia , Gentianella , Megacodon and several lineages of Swertia share a striate–reticulate or reticulate exine pattern. This is interpreted as the plesiomorphic character state of Swertiinae. There is also considerable variation of derived patterns; for example, different types of microechinate or almost smooth pollen was repeatedly observed in distantly related groups. Another extreme was the ring-shaped reticulation found in a North American species of Gentianopsis . Unfortunately, major relationships as revealed by molecular analyses were rarely supported because of the abundance of the plesiomorphic type and homoplasy even on low taxonomic levels; for example, within Lomatogonium . Exine variation was particularly useful in characterizing independent lineages of Swertia . For example, according to pollen characters and in agreement with other data, the Asian Swertia cuneata is a sister group of a strongly diversified African lineage and Swertia yunnanensis , which is rather aberrant in flower morphology, seems close to parts of Lomatogonium .  © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society , 2008, 157 , 323–341.     

Pollen morphology and systematics of palaeotropical Phyllanthus and related genera of subtribe Phyllanthinae (Euphorbiaceae)

The observations of pollen from 27 species of subtribe Phyllanthinae using scanning electron microscopy reveal considerable morphological diversity in palaeotropical Phyllanthus and the related palaeotropical genera Breynia, Glochidion, and Sauropus. The tribe appears to be monophyletic, but the pollen morphology does not support the monophyly of Phyllanthus or Sauropus. Within Phyllanthus, the pollen characters suggest a close relationship between the subgenera Emblica and Phyllanthodendron. They also reveal a surprising morphological congruence between the pollen grains of section Ceramanthus (subgenus Isocladus) and those of subgenus Eriococcus, although it is not clear whether this similarity is homoplastic. The presence of diploporate colpi is a synapomorphy uniting Breynia and Sauropus, but may be homoplastic in Phyllanthus. The affinities suggested by the morphological features of the pollen in the Phyllanthinae are concordant with recent molecular phylogenies. © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 157 , 591–608.     

Phylloclade anatomy in Phyllanthus section Choretropsis (Phyllanthaceae)

The phylloclade anatomy of nine Brazilian species of the genus Phyllanthus section Choretropsis was studied, and its systematic significance was analysed. The arrangement of epidermal cells varied according to the species, with the guard cells of the stomatal apparatus distributed randomly or with the longitudinal axis parallel to the branch. In both branch types, the stomata were distributed on the phylloclade surface, except in P. gladiatus which showed hypostomatic branches. In most of the species, phylloclades developed a vascular system surrounded by clusters of fibres and cortical parenchyma, which could differentiate into palisade‐like tissue and ground tissue like a true leaf blade. The distribution of the vascular system followed the cylindrical or flattened diameter shape of the branch. In subsection Choretropsis, the cylindrical branches of P. choretroides, contrary to other species, developed a second ring formed by clusters of fibres surrounded by large parenchymal cells. In subsection Applanata, the palisade‐like cells of P. flagelliformis phylloclades acquired an isodiametric shape. The highest level of specialization was observed in P. gladiatus, which developed phylloclades with a dorsiventral structure, a main vein, and small sets of lateral vascular bundles like a typical leaf blade. The tissue arrangement of different species showed a gradual pattern, which was correlated with the different branch morphology and plant architecture. Despite the transitional pattern, it was suggested that some of the phylloclades had an independent origin and convergent evolution in the Choretropsis section. The anatomy of different branch types was shown to be useful, either in isolation or in association with other morphological characters, to delimit the species and to further understand their relationships and phylogeny. © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 157 , 91–102.     

Pollen morphology of the genus Cornukaempferia (Zingiberaceae) in Thailand

Cornukaempferia is a recently described genus of Zingiberaceae which only occurs in Thailand as a rare genus with limited geographical distribution. Only three species have been described so far, including a recently described new species, C. larsenii. These three species are morphologically very similar and additional data on other biological aspects are useful for the elucidation of their relationship. Pollen morphology of all three species of genus Cornukaempferia has been studied by light microscopy and scanning electron microscopy. The pollen grains are monad, spherical, inaperturate. The exine sculpture is echinate with psilate between the spines for C. aurantiflora and C. longipetiolata, or echinate with regulate between the spines for C. larsenii. This obser-vation helps support the taxonomic status of C. larsenii.     

The pollen morphology of Phyllanthus (Euphorbiaceae) section Choretropsis

The pollen grain morphology of the eight South American species of the genus Phyllanthus , subgenus Phyllanthus , section Choretropsis , endemic to Brazil, was investigated using light and scanning electron microscopy. The high homogeneity in pollen shape, as well as aperture number and configuration, allowed the definition of a basic pollen type for this section, viz. prolate spheroidal or subprolate, zonocolporate with 4–5 invaginated colpi, sexine reticulate. Two subtypes have been observed, based on pollen grain apertures and exine sculpture. The first subtype is characterized by straight and narrow colpi provided with costae, lalongate endoapertures and a reticulum of rectilineous muri as in P. choretroides and P. spartioides . The second subtype has wider colpi without costae, circular endoapertures and a reticulum with curvilineous muri as in P. flagelliformis , P. goianensis and P. scoparius , or curvilineous-rectilineous muri as in P. angustissimus , P. gladiatus and P. klotzschianus . It was demonstrated that the pollen shape and the number of colpi varied within and between some of the species. The results suggest that pollen grain morphology, in association with plant architecture and the presence of phylloclades, constitute additional evidence to delimit the species of the section Choretropsis and to further understand their evolutionary origin within the genus Phyllanthus.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 144 , 243–250.     

Pollen morphology of Trigonostemon and its relatives (Euphorbiaceae)

The pollen of Trigonostemon and the related genera Dimorphocalyx, Ostodes, Tritaxis and Jatropha (outgroup) has been studied with light microscopy, and scanning and transmission electron microscopy. The two major pollen types within Trigonostemon correlate well with macromorphological characters. Species belonging to the Trigonostemon reidioides type have pollen with ‘croton pattern’ ornamentation, a pistil with deeply divided stigmas (to at least half the length of the stigma arm) and stamens with a protruding appendage on the connective, while species of the Trigonostemon verrucosus type have verrucate (to almost gemmate) pollen, stigmas that are shortly cleft and stamens without an appendage on the connective. Dimorphocalyx, Ostodes, Tritaxis and Jatropha (outgroup) have similar pollen morphology, while Trigonostemon deviates from these genera in the absence of the ‘vertically’ striate ornamentation on the subunits. Therefore, when compared with an existing phylogeny of the Euphorbiaceae, the pollen characters of Trigonostemon appear to be derived. Moreover, because the ‘croton pattern’ ornamentation itself is widely shared by the ‘inaperturate crotonoids’, the loss of that structure in the Trigonostemon verrucosus type pollen is considered a further apomorphy.     

Pollen morphology of the Flueggeinae (Euphorbiaceae,Phyllanthoideae)

A total of 129 species from the subtribe Flueggeinae of the tribe Phyllantheae (Euphorbiaceae, Phyllanthoideae) were investigated using light and scanning electron microscopy, and 10 species using transmission electron microscopy, in order to evaluate the relationships between the eight constituent genera: Breynia, Flueggea, Glochidion, Margaritaria, Phyllanthus, Reverchonia, Richeriella, and Sauropus. Of these genera, Flueggea, Margaritaria and Richeriella share pollen with a prolate spheroidal meridional outline and a 3-colporate aperture system. Pollen of Reverchonia is also 3-colporate, but differs from that of the Flueggea alliance by its clearly prolate shape, tilioid ornamentation and absence of costae endopori. Breynia and Sauropus have 4–12 and 3–16-colporate pollen, respectively, with diploporate colpi. Two pollen types are recognised in Breynia, and four in Sauropus, one of which supports the recognition of Sect. Hemisauropus. Glochidion pollen is 3–6-colporate, and similar to that of Breynia in having reticulate sculpture with Y-shaped sexine structures, but it has monoporate colpi. Of the genus Phyllanthus, only species with pollen with diploporate colpi have been studied. Seven types are described. Diploporate Phyllanthus pollen can be distinguished from that of Breynia and Sauropus by its distinct colpus margins consisting of parallel muri. Colpal irregularities and endoaperture configurations in the subtribe are discussed, and pollen morphological trends are hypothesised. Placed in the successiform aperture series, the Flueggea alliance and Reverchonia form a basal group. Glochidion is considered intermediate, giving rise to the Breynia-Sauropus group. The relationship with Phyllanthus remains unclear.     

Seed and capsule morphology of Iranian perennial species of Euphorbia (Euphorbiaceae) and its phylogenetic application

Of the 480 species of Euphorbia subgenus Esula, c. 290 occur in the Mediterranean and Irano‐Turanian regions. Turkey and Iran are the most species‐rich countries in Asia with 83 and 74 species, respectively. Following our previous paper on annual species of Iranian Euphorbia, we studied the quantitative and qualitative macro‐ and micromorphological traits of seeds and capsules of 47 perennial species, including E . ferdowsiana sp. nov. , E . sulphurea sp. nov. and E. glareosa, as a first report from Iran. A key for all Iranian perennial Euphorbia spp. based on seed and capsule morphology is provided. The phylogenetic relationships of Iranian species based on internal transcribed spacer (ITS) nuclear and ndhF plastid regions are updated and used for the characterization of the synapomorphies of each clade. Capsule shape, seed shape, seed surface and shape of the caruncle have been found to be homoplastic, whereas the presence or absence of granulate elements on seed surfaces represents a phylogenetically important trait for section delimitation. The capsule surface is synapomorphic for several sections, including Helioscopia (tuberculate‐verrucose), Myrsiniteae (vesiculate) and Esula (granulate), and seed shape is synapomorphic for sections Helioscopia (ellipsoidal), Myrsiniteae (ovoid‐quadrangular) and Herpetorrhizae (pseudo‐hexahedral). Reversals have also taken place in some features, including capsule surface (E. mazandaranica, E. altissima) and seed shape (E. densa, E. aleppica). It seems that ecarunculate seeds are plesiomorphic in sections Helioscopia (E. eriophora) and Herpetorrhizae (E. consanguinea and E. turczaninowii). © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 00 , 000–000.     

中国槲寄生属花粉形态的研究

首次系统地报道中国槲寄生属10种植物花粉形态。通过光学显微镜和扫描电镜观察,国产槲寄生属种类可按花粉的外壁纹饰分为两大类群,与形态上划分相一致。类群Ⅰ(8种)具不显著或近光滑的外壁纹饰,这一类型又可分为2个亚型:纹饰在花粉各个部位基本上是一致的和纹饰在赤道和极、沟两侧不一致的;类群Ⅱ(2种)外壁具显著的杆状纹饰。但花粉性状对形态上相近的种类鉴定意义不大。还讨论了槲寄生属花粉的可能的演化趋势。     

Pollen morphology of selected Central European species from subgenera Vignea and Carex (Carex,Cyperaceae) and its relation to taxonomy

This study analysed the taxonomic utility of selected features of pollen grains of 45 Central European species of subgenera Vignea and Carex of the genus Carex. Seven quantitative features were analysed: [length of polar axis (P), length of equatorial axis (E), exine thickness on the pole (Exp), exine thickness on the equator plane (Exe), P/E ratio and the relative thickness of exine (Exp/P and Exe/E ratio)] and outline, shape and exine sculpture. Among the biometrical features, the most important for Carex taxonomy (particularly at the level of subgenus) is the thickness of exine (Exp). Marked differences in the morphological structure of inflorescences at the subgenus level are reflected in the differences of pollen features. Results of our studies clearly show an explicit (and not previously described in the palynological literature) taxonomic division of the genus Carex on the basis of pollen biometrical features into two units corresponding exactly to the subgenera (Vignea and Carex). However, the features of pollen grains analysed did not contribute any essential taxonomic criteria at levels lower than subgenus (e.g. sections). An exception was for C. disticha, where it was observed that the poroid areas occurred in pairs, a new feature for sedges. © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 164 , 422–439.     

Molecular analysis of phylogenetic relationships among Myrmecophytic macaranga species (Euphorbiaceae)

Many species of the paleotropical pioneer tree genus Macaranga Thou. (Euphorbiaceae) live in association with ants. Various types of mutualistic interactions exist, ranging from the attraction of unspecific ant visitors to obligate myrmecophytism. In the latter, nesting space and food bodies are exchanged for protection by highly specific ant partners (mainly species of the myrmicine genus Crematogaster). As a first step toward elucidating the coevolution of ant-plant interactions in the Macaranga-Crematogaster system, we have initiated a molecular investigation of the plant partners' phylogeny. Nuclear ribosomal DNA internal transcribed spacer (ITS) sequences were analyzed for 73 accessions from 47 Macaranga species, representing 17 sections or informally described species groups. Three accessions from the putative sister taxon Mallotus Lour, were included as outgroups. Cladograms of the ITS data revealed Macaranga to be nested within Mallotus. ITS sequences are highly similar within section Pachystemon s.str., suggesting a relatively recent and rapid radiation of obligate myrmecophytes within this section. Forty-three accessions, mainly of ant-inhabited species, were additionally investigated by random amplified polymorphic DNA (RAPD) and microsatellite-primed PCR (MP-PCR) techniques. Phenetic analysis of RAPD and MP-PCR banding profiles generally confirmed the ITS results. Best resolutions for individual clades were obtained when ITS and RAPD/MP-PCR data were combined into a single matrix and analyzed phenetically. The combined analysis suggests multiple (four) rather than a single evolutionary origin of myrmecophytism, at least one reversal from obligate myrmecophytism to nonmyrmecophytism, and one loss of mutualistic specifity.     

Pollen morphology of the tribe Rhinantheae (Orobanchaceae) and its systematic significances

Pollen morphology of 36 species representing 14 genera within the tribe Rhinantheae in the family Orobanchaceae was studied and illustrated with light microscopy (LM) and scanning electron microscopy (SEM). Five major pollen types were recognized on the basis of exine ornamentation. Within these major types, minor types (subtypes) were distinguished based on exine surface pattern, size, shape, amb form, colpi and colpus membrane. These types and subtypes are as follows: type I. retipilate: subtype Ia. regular retipilate: (1) pollen size < 27 μm, (2) pollen size > 27 μm, subtype Ib. irregular retipilate; type II. verrucate: subtype IIa. macro-verrucate, subtype IIb. verrucate, subtype IIc. sparse verrucate; type III. retirugulate; type IV. granulate; type V. micro-reticulate. A key to pollen morphology of genera studied within the Rhinantheae was made based on pollen morphology from our study and earlier work. Combining with other sources of information on the Rhinantheae, the systematic relationships of this tribe are discussed. Rhinantheae pollen displays considerable variation between genera and species, with taxonomically significant characters at genus and species level. Palynological characteristics provide evidence for interpreting the conflicting views concerning the “Pterygiella Complex”. The evolutionary trend in exine sculpture of Rhinantheae could be proposed, namely that retipilate sculpturing which is the most widespread type is more primitive than the other types (such as foveolate, granulate, regulate, reticulate, retirugulate and verrucate). The pollen data in present study and the view of Hong (1986), as well as the molecular data from Bennett and Mathews (2006) indicated that Asia and related regions were likely to the origin centre of the tribe Rhinantheae.     

A Revision of Chinese Phyllanthus (Euphorbiaceae)

The present paper is the outcome of a taxonomic study of Chinese Phyllanthus plants in our flora. Phyllanthus is one of the largest genera in Euphorbiaceae, and is distributed in whole China except for the northwest, extending from 92° to 124° E and 18° to 41°N. It includes 6 subgenera, 7 sections, 33 species and 4 varieties, of which 1 subgenus and 5 species are described as new, and 1 new name and 9 synonyms are proposed.     

Pollen morphology and relationships of the Misodendraceae (Santalales)

The stem–parasitic family Misodendraceae is composed of a single genus, Misoden–drum , of 12 species endemic to the subantarctic Nothofagus forests of Chile and Argentina. Pollen of nine species was examined in the light microscope and scanning and transmission electron microscopes. Pollen is spheroidal (P/E 1 :1), sparsely echi–nate and polyporate. Aperture number is variable within and among species ranging from (3–)4–19 pores scattered randomly over the surface. Ultrastructurally, the pollen wall is composed primarily of endexine with the ektexine represented only by spines and an occasional thin granular layer between these elements. Pollen data indicate ties with the recently resurrected santalalean family Eremolepidaceae including Lepidoceras.     

Pollen morphology and its relation with meiotic irregularities in ten species of Campuloclinium (Eupatorieae,Asteraceae)

Pollen grains of ten species of Campuloclinium (Eupatorieae, Asteraceae) are described and illustrated using light and scanning electron microscopy. The species included in this study are C. burchelli, C. campuloclinioides, C. chlorolepis, C. hirsutum, C. irwinii, C. macrocephalum, C. megacephalum, C. parvulum, C. purpurascens and C. riedelli. Pollen grains of Campuloclinium are typically radially symmetric, echinate, tectate, oblate-spheroidal to prolate-spheroidal (P/E ratio: 0.94–1.10). Pollen is mostly triaperturate, but tetraaperturate or biaperturate pollen also occurs in some species. The pollen diameter was highly variable ranging from 10.54 µm to 36.18 μm. The significance of the variation in type and size of pollen is discussed in relation to diverse meiotic abnormalities and possible reproductive apomictic processes within the polyploid species.     

Pollen morphology in the genus Veronica L. (Plantaginaceae) and its systematic significance

Pollen grains of 30, mainly annual, species from 134 populations of the genus Veronica (Plantaginaceae, formerly Scrophulariaceae) from the Mediterranean Region, have been studied with light and scanning electron microscopy. Three pollen types are defined based on pollen size and exine surface sculpture. In some cases within the study group pollen size can be useful in species determination. Hypothetical relationships of pollen grain size and aperture number with ploidy level, style length and corolla diameter are discussed. The ornamentation of the exine in Veronica, although generally a conservative character within the genus, gives some additional support to the most recent infrageneric classification of Veronica.     

In vitro culture of Phyllanthus caroliniensis (Euphorbiaceae)

An efficient micropropagation protocol was developed for the medicinal plant Phyllanthus caroliniensis (Euphorbiaceae) using nodal segments for axillary shoot proliferation. Maximum multiplication (21–23 shoots per explant) was achieved on MS or AR media supplemented with either 5.0 μM BA, 1.25–5.0 μM kinetin or 2.5–5.0 μM 2iP. Rooting was achieved with 80–100% of the microshoots on MS medium without growth regulators, although 1.25 μM NAA and 1.25–5.0 μM IAA promoted significant increases in the number of roots per explant. Regenerated plants were successfully acclimatized and about 88% of plantlets survived under ex vitro conditions. Flowering was observed on in vitro grown plantlets and after 3–4 weeks of acclimatization. High frequency callus initiation and growth was achieved when nodal segment explants were inoculated in the vertical position on MS medium supplemented with 5.0 μM 2,4-D. Root cultures were successfully established on MS medium containing 1.1 μM NAA. The optimized micropropagation, callus and root culture protocols offer the possibility to use cell/root culture techniques for vegetative propagation and secondary metabolism studies. This revised version was published online in June 2006 with corrections to the Cover Date.