A molecular phylogenetic and fruit evolutionary analysis of the major groups of the paleotropical woody bamboos (Gramineae: Bambusoideae) based on nuclear ITS, GBSSI gene and plastid trnL-F DNA sequences

This study presented the first molecular phylogenetic analysis of the major clades of woody bamboos of the Old World tropics based on nuclear and chloroplast sequences (ITS, GBSSI and trnL-F). Sequence data from 53 species, representing 17 paleotropical woody bamboo genera, were analyzed using the maximum parsimony and Bayesian inference methods. All examined ingroup taxa were clustered into two clades, i.e., the Bambusinae+Dinochloa clade and the Melocanninae clade. The former clade included Bambusa, Bonia, Dendrocalamus, Dendrocalamopsis, Dinochloa, Gigantochloa, Molecalamus, Neomicrocalamus, Neosinocalamus, Oxytenanthera s. str. (sensu stricto), Racemobambos and Thyrsostachys. The Melocanninae clade consisted of Cephalostachyum, Leptocanna (better treated as part of Cephalostachyum), Melocanna, Pseudostachyum and Schizostachyum s. str. The subtribe Racemobambosinae and tribes Dendrocalameae and Oxytenanthereae were not supported and may be better placed in subtribe Bambusinae. The ovary characters seemed to be good criteria to distinguish these two clades. The reconstruction of ancestral fruit characters indicated that the bacoid caryopsis, namely, fleshy or berry-like fruits, was found to be scattered in three lineages of the examined paleotropical woody bamboos. Fruit characters are thus not reliable indicators of phylogeny and bacoid caryopsis likely represents a specialization for particular ecological conditions.     

[竹思]簩竹属及其近缘属叶表皮微形态特征

用扫描电镜和光学显微镜对[竹思]簩竹属（Schizostachyum Nees）7个种,及其近缘的梨竹属（Melocanna Trin．）、泡竹属（Pseudostachyum Munro）、薄竹属（Lepzocanna Chia et H．L．Fung）各1个种以及空竹属（Cephalostachyum Munro）6个种的叶表皮微形态特征进行了观察。结果表明竹叶下表皮乳突的形状和在气孔器上及周围的排列式样在这5个属中各有不同,具有重要的分类学和系统学价值。同时我们的结果不支持广义的笃竹属;而梨竹属、空竹属和泡竹属则被支持独立成属。另外,薄竹（Leptocanna chinensis）和红毛[竹思]簩竹（Schizostachyum sanguineum）的叶表皮微形态特征非常接近空竹属;香糯竹（Cephalostachyum pergracile）和金毛空竹（C．virgatum）的叶表皮微形态特征则十分接近[竹思]簩竹属。     

Micromorphological Study on Leaf Epidermis of Schizostachyum and Its Allies ( Poaceae: Bambusoideae)

Micromorphological characters of leaf epidermis of 16 published species from Schizostachy um Nees and its allies ( including Melocanna Trin. , Pseudostachyum Munro, Leptocanna Chia et H. L. Fung and Cephalostachyum Munro) were examined by both light and scanning electron microscopies. The results indicated that the papillae forms and distributional patterns on the abaxial surface were taxonomically informative at the generic level. The characters of papillae did not support Schiz ostachyum sensu lato; meanwhile, it was supported that Melocanna, Pseudostachyum and Cep halostachyum were better to be treated as separate genera respectively. Furthermore, it seemed to be more appropriate to place Leptocanna chinensis and Schizostachyum sanguineum in Cep halostachyum; and to place Cephalostachyum pergr acile and C1 virgatum in Schizostachy um .     

Revision on Cephalostachyum Munro (Poaceae: Bambusoideae) in China

Species of Cephalostachyum Munro (Poaceae: Bambusoideae) from China are distributed in Yunnan and Tibet, mainly in Yunnan. In this paper, we discussed species of Cephalostachyum and compiled a key to species from China, based on recent studies on micromorphological characters of leaf epidermis and molecular phylogenetics of paleotropical woody bamboos. There is a total of seven species of Cephalostachyum distributed in China, all in Yunnan: Cchinense (Rendle) DZ. Li et HQ. Yang, Cfuchsianum Gamble et Hook. f., Cmannii (Gamble) Stapleton et DZ. Li, Cpallidum Munro, Cpingbianense (Hsueh et YM. Yang ex Yi et al.) DZ. Li et HQ. Yang, Csanguineum (WP. Zhang) DZ. Li et HQ. Yang and Cscandens Bor. Leptocanna Chia et HL. Fung and Cvirulentum YM Yang et F. Du are synonyms of Cephalostachyum Munro and Cfuchsianum Gamble et Hook. f. respectively. On the other hand, Cpergracile Munro and Cvirgatum (Munro) Kurz are morphologically closer to Schizostachyum Nees than to Cephalostachyum, and they should be treated as members of Schizostachyum. This paper is of significance to a worldwide revision of Cephalostachyum.     

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.     

Comparative morphology of the leaf epidermis in Schisandra (Schisandraceae)

The leaf epidermis of 127 samples representing ten species within Schisandra Michaux. and one species of the related genus, Kadsura Kaempf. ex Juss., was investigated using light and scanning electron microscopy. Many characters of the leaf epidermis in Schisandra , such as pattern of epidermal cells, type of stomata, shape of guard cell pairs and cuticular ornamentation, are usually constant within species and thus make good characters for studying the relationship between and within genera. A new character, rim number, of the outer stomatal rim in the genus is introduced. It is shown that double outer stomatal rims occur only in evergreen Schisandra species, whereas a single rim occurs in deciduous species. This character supports the classification of Schisandra into two subgenera based on habit and androecial organs. This classification is also supported by additional morphological and molecular taxonomic characters. Kadsura coccinea (Lem.) Smith A. C. is the most primitive taxon in the related genus Kadsura . The outer stomatal rim of this species also has double rims. Combined with morphological and molecular evidence, this suggests that Schisandra and Kadsura are closely related and may share a recent common ancestor.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 148 , 39–56.     

Taxonomic relationships of the western Asian taxa of Gentiana sect. Pneumonanthe

The taxonomic position of western Asian members of Gentiana sect. Pneumonanthe has long been a matter of conflict. In this paper, the six western Asian species currently recognized as belonging to sect. Pneumonanthe ( G. boissieri , G. calycina , G. freyniana , G. gelida , G. paradoxa , and G. septemfida ) are compared and their relationships established using a morphological phylogenetic analysis. Seed testa and flower and leaf morphological characters were studied and 11 characters were selected for a cladistic analysis. Euro-Siberian and Far Eastern taxa of sect. Pneumonanthe ( G. pneumonanthe and G. scabra ) were used as outgroups. Our results suggest the presence of two morphologically distinct clades within the western Asian gentians: a Septemfida and a Gelida clade. G. calycina and G. freyniana show close affinities to G. boissieri and G. gelida , and are distinct from G. septemfida s.l. Biogeographical aspects of the two groups are discussed.  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 152 , 197–208.     

Cauline anatomy of native woody bamboos in Argentina and neighbouring areas: epidermis

The presence of woody bamboos in Argentine archaeological sites led to attempts to identify them. In general, fragments of culm are recovered with nodes, internodes (or parts of them) and occasionally nodes with buds or their vestiges, but without leaves. Epidermal studies were carried out on developed culms of several bamboo genera growing currently in Argentina and neighbouring areas, with the purpose of detecting characters of taxonomic value that will allow identification at generic or species level. The selected characters did indeed facilitate the determination of the archaeological material. Fifteen species of six native genera from America were studied: Aulonemia Goudot, Colanthelia McClure & E. W. Smith, Chusquea Kunth, Guadua Kunth, Merostachys Spreng. and Rhipidocladum McClure. Epidermal cauline anatomy was described and illustrated using scanning electron and light microscopes. An identification key based on cauline epidermis is presented. © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society , 138 , 45–55.     

Scanning electron microscopy of the leaf epidermis in Arundinarieae (Poaceae: Bambusoideae): evolutionary implications of selected micromorphological features

Arundinarieae (temperate woody bamboos) is a taxonomically challenging group and the taxa in this tribe show a low level of DNA sequence variation, especially for plastid markers. As a result of the scarcity of flowering collections, species identification mainly depends on vegetative features, and leaf epidermal micromorphology has proven to be useful in bamboo taxonomy. In this study, we used scanning electron microscopy to investigate the abaxial leaf epidermal micromorphology of 94 species in 24 genera of Arundinarieae and three species of Bambuseae. Leaf epidermal characteristics differed mainly in the number and distribution pattern of papillae. Seven papilla forms were assigned, which had little taxonomic value at the generic and subtribal levels. However, the papilla patterns combined with other leaf epidermal features were useful in the discrimination of several species. Most papilla types were shared between tribes Arundinarieae and Bambuseae. We assessed the phylogenetic implications of the leaf epidermal micromorphology based on recently published molecular phylogenetic analyses. With one exception, none of the papilla types corresponded to a particular clade in the plastid or nuclear phylogenetic trees. Based on these results, the papilla patterns have limited phylogenetic value. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 176 , 46–65.     

Molecular phylogeny of the arthrostylidioid bamboos (Poaceae: Bambusoideae: Bambuseae: Arthrostylidiinae) and new genus Didymogonyx

We present the first multi-locus chloroplast phylogeny of Arthrostylidiinae, a subtribe of neotropical woody bamboos. The morphological diversity of Arthrostylidiinae makes its taxonomy difficult and prior molecular analyses of bamboos have lacked breadth of sampling within the subtribe, leaving internal relationships uncertain. We sampled 51 taxa, chosen to span the range of taxonomic diversity and morphology, and analyzed a combined chloroplast DNA dataset with six chloroplast regions: ndhF, trnD-trnT, trnC-rpoB, rps16-trnQ, trnT-trnL, and rpl16. A consensus of maximum parsimony and Bayesian inference analyses reveals monophyly of the Arthrostylidiinae and four moderately supported lineages within it. Six previously recognized genera were monophyletic, three polyphyletic, and two monotypic; Rhipidocladum sect. Didymogonyx is here raised to generic status. When mapped onto our topology, many of the morphological characters show homoplasy.     

Styloid crystals in Claoxylon (Euphorbiaceae) and allies (Claoxylinae) with notes on leaf anatomy

Claoxylon and Micrococca are the only Euphorbiaceae genera that have rough dried leaves (fresh ones are smooth) because of protruding styloid (needle-like) crystals more or less perpendicular to the leaf surface, which perforate the epidermis and cuticle. A broad leaf anatomical study of the subtribes Claoxylinae, the monogeneric Lobaniliinae, and Mercurialis of the Mercurialinae (95 of a possible 235 species in all six genera) showed that styloids are present in Claoxylon , Discoclaoxylon , Erythrococca , Lobanilia , and Micrococca , and lacking in Claoxylopsis and Mercurialis . Contrary to Claoxylon , the dried leaves of Discoclaoxylon , Erythrococca , Lobanilia , and Micrococca are not rough, because the styloids do not perforate the epidermis during drying and therefore herbarium leaves generally remain smooth. The presence of styloids supports a clade in a recent molecular phylogenetic study that unites subtribes Claoxylinae and Lobaniliinae as Claoxylinae s.l. Mercurialis (no styloids) is sister to all other taxa (with styloids) in the monophyletic Claoxylinae s.l. The styloids form a synapomorphy for a monophyletic part of the Claoxylinae ( Claoxylon , Discoclaoxylon , Erythrococca , Lobanilia , and Micrococca ). Other leaf anatomical notes are provided, together with an overview of the occurrence of styloids, stomata, and druse crystals for most species in the Claoxylinae. The indumentum ranges from (virtually) absent to a sparse or dense tomentum of straight and thick- or thin-walled unicellular hairs (most common), curly hairs, or two-armed hairs ('Malpighiaceous hairs'). The last two hair types probably form diagnostic characters for species groups within the genus Claoxylon .  © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society , 2008, 156 , 445–457.     

Seed coat morphology of Aizoaceae–Sesuvioideae, Gisekiaceae and Molluginaceae and its systematic significance

Macro- and micromorphological characters of the seeds of 32 species belonging to Aizoaceae (26 species), Gisekiaceae (one species) and Molluginaceae (five species) were analysed for their taxonomic value. Seed morphology is found to be of considerable taxonomic value within the taxa investigated, although additional criteria are usually necessary for distinguishing the species. However, a subgroup of Trianthema , consisting of T. cussackiana , T. megasperma , T. pilosa , T. rhynchocalyptra and T. oxycalyptra var. oxycalyptra (Aizoaceae) shares the synapomorphy of seeds with scattered idioblast cells (papillae). There is little infrageneric variation within Sesuvium and Zaleya in contrast to Trianthema , whose seeds are considerably polymorphic. The fairly different seed structures found in Aizoaceae s.s . and Molluginaceae support their treatment as two distinct families. However, the position of Lineum (Molluginaceae) remains uncertain. Moreover, seed characters support the independence of Gisekia (Gisekiaceae) in a family of its own.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 148 , 189–206.     

Silica bodies and hooked papillae in lemmas of Melica species (Gramineae: Pooideae)

Melica (sensu lato ) is a genus with approximately 80 species distributed in the temperate regions of both hemispheres. The genus has been better studied morphologically than anatomically, but morphological patterns have led to conflicting subdivisions either into sections, or by segregation of the genus Bromelica . The lemma epidermis is examined here under scanning electron microscopy (SEM) for 40 species of Melica to see if it exhibits characters of systematic significance. Silica bodies and hooked papillae are reported in the lemma for the first time in the genus. Patterns of variation are evident within four structures that terminate in points: prickles, hooks, hooked papillae and macro-hairs. Fourteen qualitative characters for lemma epidermis are used in phenetic analyses. The presence or absence of macro-hairs conflicts with some other patterns, but gives some support to the classification of Papp. Characters associated with the split of the clusters were micro-hairs (straight and geniculate) and the distribution and frequency of prickles in both costal and intercostal zones. As a conclusion, Melica is not a coherent taxonomic grouping and a simple segregation of Bromelica from Melica genus is rejected.  © 2003 The Linnean Society of London , Botanical Journal of the Linnean Society , 2003, 141, 447 – 463.     

Comparative floral structure and systematics in Chrysobalanaceae s.l. (Chrysobalanaceae, Dichapetalaceae, Euphroniaceae, Trigoniaceae; Malpighiales)

Chrysobalanaceae s.l. , one of the few suprafamilial subclades of Malpighiales that is supported by molecular phylogenetic analyses, and containing Chrysobalanaceae, Dichapetalaceae, Euphroniaceae, and Trigoniaceae, was comparatively studied with regard to floral structure. The subclade is well supported by floral structure. Potential synapomorphies for Chrysobalanaceae s.l. are the following shared features: floral cup; flowers obliquely monosymmetric; sepals congenitally united at base; sepals of unequal size (outer two shorter); fertile stamens concentrated on the anterior side of the flower and sometimes united into a strap; staminodes absent in the posteriormost antepetalous position; anthers extremely introrse, with thecae almost in one plane; endothecium continuous over the dorsal side of the connective; dorsal anther pit; gynoecium completely syncarpous up to the stigma; carpel flanks slightly bulged out transversely and thus carpels demarcated from each other by a longitudinal furrow; flowers with dense unicellular, non-lignified hairs, especially on the gynoecium; light-coloured, dense indumentum on young shoots and inflorescences. Potential synapomorphies for Chrysobalanaceae + Euphroniaceae include: spur in floral cup; clawed petals; lignified hairs on petals; nectary without lobes or scales and mostly annular. Potential synapomorphies for Dichapetalaceae + Trigoniaceae include: special mucilage cells in sepals in mesophyll (in addition to epidermis); anthers almost basifixed; gynoecium synascidiate up to lower style; nectary with lobes or scales and semi-annular.  © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society , 2008, 157 , 249–309.     

Cytological studies on some representative species of the tribe Orchideae (Orchidaceae) from China

Cytological studies were carried out on 14 taxa belonging to Amitostigma , Chusua , Galearis , Habenaria , Hemipilia , Hemipiliopsis , Herminium , Peristylus and Ponerochis , collected mostly from the south-eastern part of the Hengduan Mountain Region, south-west China. Cytological data on 11 of the taxa are reported for the first time. The interphase nuclei were either of the simple chromocentre type or intermediate between simple and complex chromocentre types. The nuclear morphology of Hemipiliopsis at interphase supports the conclusion that it is related more closely to Chusua and Ponerochis than to Habenaria . At the whole tribe level, however, the results did not indicate a clear correlation between morphological features of the interphase nuclei and phylogeny. The somatic chromosome numbers were 2 n  = 42 in ten species and 2 n  = 32, 38, 40, 64 and 72 in four species. The chromosome counts of 2 n  = 32 and 64 in Habenaria aitchsonii are rare in the genus. It is proposed that the repeated change of chromosome number from x  = 7 to x  = 8 has played an important role in the evolution of the tribe Orchideae. This change has occurred mainly in the European subtribe Orchidinae, but also in the Asian subtribe Habenariinae.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 145 , 231–238.     

A new species of Silene (Caryophyllaceae) from east Anatolia, Turkey

Silene bitlisensis O.Tugay & Ertugrul from the Green Line Afforestation Zone (B9: Bitlis Province) is a new species described and illustrated here. It is closely related to S. caramanica Boiss. & Heldr. var. ilarslanii Aytaç & Dural, from which it mainly differs in its habit, leaf, and floral features.  © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society , 2008, 156 , 463–466.     

Biomechanics and development of rattans: what is special about Plectocomia himalayana Griff. (Calamoideae,Plectocomiinae)?

Mechanical and morphological studies of Plectocomia himalayana (subtribe Plectocomiinae) revealed characteristics that differ strongly from species of subtribe Calaminae ( Calamus and Daemonorops ). In species of Calaminae tested previously, the contribution of the leaf sheath drastically increases stiffness in juvenile axes and towards the apex of older plants. In P. himalayana the relative contribution of the leaf sheath to axis stiffness is less and leaf sheath senescence does not strongly reduce axial stiffness as observed in Calamus and Daemonorops . Natural aerial branching, only described in Korthalsia and Laccosperma among rattans, is common in P. himalayana . Aerial branching and adventitious roots occur frequently along old stems allowing autonomy of stems, following mechanical injury and promoting vegetative propagation. The climbing habit is known to have evolved at least twice within the Calamoideae. The results observed here suggest that climbing habits may differ in detail and that different 'climbing strategies' may have evolved within the subfamily Calamoideae resulting from: (1) variable stem flexibility, (2) the variable mechanical role of the leaf sheath ( Calamus–Daemonorops ) and (3) production of branches and aerial roots conferring a higher degree of architectural plasticity ( Plectocomia ).  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 151 , 83–91.     

Molecular phylogeny of the genus Philodendron (Araceae): delimitation and infrageneric classification

The genus Philodendron (Araceae) is a large neotropical group whose classification remains unclear. Previous classifications are based on morphological characters, mainly from the inflorescence, flower and leaf shape. The classification by Krause, with few modifications, is still the most commonly used system. To examine phylogenetic relationships in the genus, two ribosomal DNA nuclear markers, internal transcribed spacer (ITS) and external transcribed spacer (ETS), and the chloroplast intron rpl 16, were sequenced and analysed for more than 80 species of Philodendron and its close relative Homalomena . According to the resulting phylogeny, the genus Homalomena may be paraphyletic to the genus Philodendron . The inclusion of the American Homalomena species within the genus Philodendron might resolve this taxonomic problem. All three subgenera of Philodendron were revealed as monophyletic. Below the subgeneric level, the groups obtained in our phylogeny globally correspond to sections recognized in previous classifications. Among the morphological characters used by previous taxonomists to build their classifications, and which we optimized onto one of the most parsimonious trees, most characters were found to be homoplasious. However, leaf shape, characteristics of the sterile zone on the spadix and venation patterns are useful for delimiting subgenera and sections within the genus.  © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society , 2008, 156 , 13–27.     

On the confusion surrounding Pareledone charcoti (Joubin, 1905) (Cephalopoda: Octopodidae): endemic radiation in the Southern Ocean

Until recently, all papillated specimens of Pareledone were ascribed to the species Pareledone charcoti (Joubin, 1905), of which P. aurorae (Berry, 1917) was considered a junior synonym. Re-examination of the papillated type material of Pareledone, coupled with extensive fishing over several years off the Antarctic Peninsula, has led to a revision of this position. Seven new species of papillated Pareledone are identified from the Antarctic Peninsula region. They are identified by subtle taxonomic characters, such as the morphology and placement of their papillae, although traditional indices often fail to separate the species. Whilst apparently sympatric, there is some evidence of niche separation of these species with respect to depth. A key is provided for their identification.  © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society , 2005, 143 , 75−108.