Leaf anatomy of Ficus subsection Urostigma (Moraceae)

Species of Ficus subsection Urostigma show much overlapping variation in vegetative morphology, which often precludes correct identification of the species. The aim of this study was to describe the leaf anatomical characters and their variation and to check their suitability for identification. Included were 41 samples belonging to 25 species of subsection Urostigma, four samples belonging to two species of section Leucogyne and one specimen of Ficus glaberrima subsp. siamensis of subsection Conosycea. Transverse sections of lamina, midrib and petiole and cuticular macerations were used, and the observed anatomical characters are described for each species. On the basis of a limited number of studied samples, leaf anatomy shows little variation within each species and each species has a unique combination of character states, facilitating identification. Ficus arnottiana shows some leaf anatomical characters that are quite different from those of other members of subsection Urostigma, including a multiple epidermis and enlarged lithocysts on both sides of the leaf. Both characters are generally considered as typical for Ficus subsection Conosycea. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 175 , 259–281.     

Leaf epidermal studies in some Nigerian species of Ficus L. (Moraceae)

Forty-two leaf epidermal features in 20 Nigerian species of Ficus, representing three of the four subgenera and four of the seven sections of the genus found in Africa have been examined. Discontinuities in these characters clearly separated the three subgenera studied, namely, Ficus, Sycomorus and Urostigma, and supported the earlier sectional classification of the subgenus Urostigma (i.e. sections Sycidium, Galoglychia and Urostigma). The subsectional boundaries among the members of the section Galoglychia studied, however, had little backing from their leaf epidermal characteristics due to extensive overlap of the features. The results of the present studies support the genetic basis of the qualitative and quantitative variations in the leaf epidermal cells, stomata and trichomes in the Nigerian Ficus and the present and potential applications of these features in diagnostic and taxonomic investigations in the genus are discussed.     

Intraspecific classification of Ficus deltoidea Jack subsp. deltoidea (Moraceae) in Peninsular Malaysia based on morphological and molecular variations

Taxonomic classification of the important medicinal plant, Ficus deltoidea (Moraceae), is challenging because of the variability of its leaves and fig forms that occur within the species. We developed 16 nuclear Simple Sequence Repeat (nSSR) markers, and characterized them using 24 individuals from a natural population. We then studied the intraspecific variation of F. deltoidea subsp. deltoidea in Peninsular Malaysia using morphological and molecular approaches. Based on the morphological variations, we further determined the varieties that occur regionally under the above subspecies based on the leaf characteristics. As for molecular data sets, we used both chloroplast DNA (cpDNA) and nSSR markers to elucidate the phylogenetic relationship among the varieties. The cpDNA dendrogram yielded poorer resolution where most of the clades were forming paraphyletic complex. The cluster analysis based on nSSR is largely congruent with the morphological classification, with F. deltoidea subsp. deltoidea classified into four main varieties, namely var. deltoidea, var. angustifolia, var. kunstleri and var. lutescens. Our study demonstrates the applicability of molecular approach in complementing the conventional taxonomic classification.     

Molecular phylogeny and genetic variation in the genus Lilium native to China based on the internal transcribed spacer sequences of nuclear ribosomal DNA

We present a comprehensive phylogeny derived from nuclear ribosomal DNA (nrDNA) for 214 samples representing 98 species and five varieties, including 44 species and five varieties native to China. Our collection of 25 species and five varieties (44 samples) covering all five sections of the genus (Comber) distributed in China also were included in the internal transcribed spacer (ITS) database. This study incorporates previous research with an emphasis on Chinese species, including the controversial subsection, Sinomartagon 5c Comber. In the phylogenetic tree obtained by maximum parsimony (PAUP) and maximum likelihood (RAxML) analyses, the samples were divided into four major groups. Our results suggest that the subsection (subsect.) 5c Comber should be classified into the true subsect. 5c and the section (sect.) Lophophorum. And the latter was divided into three subsections (subsect. Lophophorum I, subsect. Lophophorum II, and subsect. Lophophorum III). Based on molecular phylogenetic analysis and fluorescence in situ hybridization, we report that L. henryi and L. rosthornii are closely related, and we propose their classification into subsect. Leucolirion 6a. Our results support Comber’s subdivision of sect. Leucolirion, which was primarily based on bulb color. Chinese species were divided into five sections: sect. Martagon, sect. Archelirion, sect. Leucolirion, sect. Sinomartagon, and sect. Lophophorum. These findings contribute to our understanding of the phylogeny, origin, and classification of Lilium.     

Rhododendron kuomeianum (Ericaceae), a new species from northeastern Yunnan (China), based on morphological and genomic data

Rhododendron kuomeianum Y.H. Chang, J. Nielsen & Y.P. Ma, a new species of Rhododendron (Ericaceae) within subsect. Maddenia in sect. Rhododendron from Yiliang County, NE Yunnan, China, is described and illustrated. The new species is similar to R. valentinianum, but it can be easily distinguished by its sparse scales on the abaxial surface of the leaf blade, fewer flowers per inflorescence and white corolla with pale red margins. There are also differences in the widths of calyx lobes, leaf blade shape and indumentum characteristics of the petiole between the new species and Rhododendron linearilobum. We confirmed that R. kuomeianum is a new species closely related to R. valentinianum and R. changii with phylogenomic studies of 10 species within this subsection based on restriction site-associated DNA sequencing (RAD-seq) data. These phylogenomic analyses also clarified additional taxonomic problems in this subsection previously raised by morphological analysis. Our findings make a strong case for using next-generation sequencing to explore phylogenetic relationships and identify new species, especially in plants groups with complicated taxonomic problems.     

Chloroplast phylogenomics and the taxonomy of Saxifraga section Ciliatae (Saxifragaceae)

Comprising ca. 200 species, Saxifraga sect. Ciliatae is the most species-rich section of Saxifraga s.str., whose center of diversity is in the Tibeto-Himalayan region. The infra-sectional classification of sect. Ciliatae is still in debate due to the high level of species richness, as well as remarkable variations of habitat, morphology, physiology and life cycles. Subdivisions of this section proposed in various taxonomic systems have not been adequately tested in previous phylogenetic studies, partly due to low taxonomic sampling density, but also to the use of few DNA markers. In order to achieve a more robust infra-sectional classification of sect. Ciliatae, complete chloroplast genomes of 94 taxa from this section were analyzed, of which 93 were newly sequenced, assembled and annotated. The length of the 94 plastomes of sect. Ciliatae taxa range from 143,479 to 159,938 bp, encoding 75 to 79 unique protein-coding genes (PCGs). Analyses of the 94 plastomes revealed high conservation in structural organization, gene arrangement, and gene content. Gene loss and changes of IR boundaries were detected but in extremely low frequency. The molecular phylogenetic tree from concatenated PCGs and complete chloroplast genome sequences exhibited high resolution and support values and confirms that sect. Ciliatae is monophyletic. Three well-supported clades were revealed within the section that agree relatively well with the subsectional taxonomy of Gornall (1987), but some minor modifications should be made. Firstly, the monotypic subsection Cinerascentes should be abandoned and its constituent species, S. cinerascens, assigned to subsect. Gemmiparae. Secondly, subsections Rosulares and Serpyllifoliae should be merged and become subsect. Rosulares. Section Ciliatae thus comprises: subsect. Hirculoideae Engl. & Irmsch.; subsect. Rosulares Gornall; subsect. Gemmiparae Engl. & Irmsch.; subsect. Flagellares (C. B. Clarke) Engl. & Irmsch. and subsect. Hemisphaericae (Engl. & Irmsch.) Gornall.     

栝楼属基于核糖体ITS序列的系统发育分析

栝楼属（Trichosanthes）是葫芦科(Cucurbitaceae)中一个种类较多、药用价值较大的属。本文基于ITS序列分析了栝楼属16个种的系统发育关系。聚类分析表明：在组的划分上,具有分类争议的截叶组(sect. Truncata)与大苞组(sect. Involucraria)形成一大支,处于一亚分枝位置,不支持独立成组。在属内分类地位上,在核酸水平不支持贵州栝楼(T.guizhouensis)为一独立种,结合形态特征建议并入双边栝楼(T.rosthornii)。大方油栝楼(T.dafangensis)与小苞组(sect. Trichosanthes)成为姊妹群,处于一个向叶苞组(sect. Foliobracteola)过渡位置,结合其形态特征,建议作为小苞组(sect. Trichosanthes)成员。     

Morphological Characters Useful for the Delimitation of Taxa Within Viola Subsect. Viola (Violaceae): A Morphometric Study from the West Carpathians

Forty-nine morphological characters were scored or measured on 44 populations (376 individuals) of Viola subsect. Viola from the West Carpathians and adjacent areas (Slovakia, Czech Republic, Austria and Hungary). The presence of six species, namely V. alba (represented by subsp. alba), V. ambigua, V. collina, V. hirta, V. odorata and V. suavis s.l. was revealed based on pollen fertility, cytological and morphometric analyses. The morphological characters traditionally used to delimit taxa within the subsection and those revealed by our study as most reliable are widely discussed. A key for identifying the taxa and most common hybrids of subsection Viola occurring in the West Carpathians is presented. Chromosome counting and flow cytometry were used to determine the ploidy levels of the populations studied. All individuals of V. alba subsp. alba, V. collina, V. hirta and V. odorata were tetraploid, while those of V. ambigua and V. suavis s.l. were octoploid.     

Primula wawushanica sp. nov. (Primulaceae) from Sichuan,southwestern China

A new species of Primulaceae, Primula wawushanica G. Hao, C. M. Hu & Y. Xu, from Sichuan, China, is described and illustrated. In general morphology it is clearly allied to P. sect. Petiolares subsect. Davidii, characterized by firmly papery or leathery leaves, with adaxially impressed veins which are often prominently raised and alveolate abaxially. However, from other species of this subsection, P. wawushanica is distinguished by its glabrous and sessile leaves, smaller flowers, and by the shorter scape at flowering time.     

Figs (Ficus,Moraceae) in Urban Hong Kong,South China1

Hong Kong (22°N) is on the northern margins of the Asian tropics and has a native fig flora of 24 species. A total of 3.4 km² of the urban area on Hong Kong Island was surveyed for spontaneous and planted fig plants of reproductive size. The 1124 individuals included 14 species in four subgenera: seven native species (F. fistulosa, F. hirta, F. hispida, F. microcarpa, F. pumila, F. subpisocarpa, and F. variegata), four of which were sometimes planted, one probably native (F. virens), one naturalizing species (F. religiosa), and five exotic species that occurred only in cultivation, two of which (F. altissima, F. rumphii) are pollinated and produce viable seeds. The native species in the two most common subgenera form distinct ecological guilds: those in subgenus Sycomorus bear large, many‐seeded, green or yellow figs and are bat‐dispersed pioneers on exposed soil; those in subgenus Urostigma bear small, few‐seeded, dark‐purple figs and are largely bird‐dispersed and epilithic. The density of potentially fruiting fig plants in the study area (2.3/ha) was within the range reported for tropical forests and between them they were visited by the entire urban frugivore fauna. This study shows the importance of the urban fig flora to urban wildlife and also highlights the risk of cultivated Ficus species becoming invasive, despite their obligate species‐specific pollinator mutualisms.     

Fossil records of subsection Pinus (genus Pinus,Pinaceae) from the Cenozoic in Japan

Extant pines of subsection Pinus (section Pinus, genus Pinus, Pinaceae) are predominantly distributed in Eastern Asia. However, the extent of diversification in the section has yet to be fully clarified. We reviewed fossil records of subsection Pinus from Japan and collected permineralized materials, in which anatomical details are preserved for better understanding of the diversification. Our results suggest that this subsection appeared in Japan no earlier than the Middle Eocene, with extant species (i.e., Pinus densiflora and Pinus thunbergii) appearing around the beginning of the Pleistocene. Pinus fujiii (Early Miocene to Early Pleistocene) is inferred to have a close affinity to P. thunbergii based on the medial arrangement of its leaf resin canals. Additionally, P. fujiii has a similar cone morphology to those of extant species living in China, bridging the morphological gap between P. thunbergii and Chinese relatives of P. thunbergii as inferred by molecular phylogenetic analyses. Our results also suggest that taxonomic revisions of Pinus miocenica and Pinus oligolepis are required among the Japanese fossil species reported to date.     

Molecular analyses of <Emphasis Type="Italic">Ficus erecta</Emphasis> and its allies within the subsection <Emphasis Type="Italic">Frutescentiae</Emphasis> (Moraceae)

Ficus (Moraceae) is a keystone group in tropical and subtropical forests with remarkable diversity of species and taxonomical challenges as a consequence of fig–pollinator coevolution. Ficus subsect. Frutescentiae includes about 30 species that are predominantly shrubs or small trees with Terminalia branching. Many of these species are difficult to delimit morphologically, and the group includes a tangle of uncertain taxa and incorrectly applied names. We conducted a phylogenetic analysis with internal and external transcribed spacer data (ITS and ETS) and data from 18 polymorphic microsatellite loci to evaluate the species status of the most perplexing members of this subsection. The results confirm the monophyly of subsect. Frutescentiae, with F. pedunculosa as sister to the rest. The F. erecta complex comprises approximately 17 taxa: F. erecta, F. abelii, F. boninsimae, F. nishimurae, F. iidaiana, F. gasparriniana var. laceratifolia, F. gasparriniana var. viridescens, F. pyriformis, F. stenophylla, F. fusuiensis, F. fengkaiensis, F. sinociliata, F. tannoensis, F. vaccinioides, F. formosana, F. pandurata, and F. periptera. The last five of these were supported as good species, while the others were not well supported by the present evidence. Evidence also supported the status of the non-F. erecta complex species including. F. pedunculosa, F. ischnopoda, F. heteromorpha, and F. variolosa. Ficus filicauda and F. neriifolia are possibly conspecific. The species status of F. potingensis should be restored and it should be treated as a member of section Eriosycea. Identification of the remaining taxa (F. gasparriniana var. esquirolii, F. ruyuanensis, F. daimingshanensis, F. chapaensis, F. changii, F. trivia, and F. tuphapensis) and their relationships to the F. erecta complex were not clarified. As a whole, only ten species in this subsection are confirmed, one is excluded, one is synonymous, and the others are either unresolved or short of samples. There appears to be a consistent genetic background among these unresolved groups, which suggests that repeated hybridization (as a result of pollinator host shifts) has filled up the interspecific gaps during the fig–pollinator coevolution process.     

Seed and gland morphology in Euphorbia (Euphorbiaceae) with focus on their systematic and phylogenetic importance, a case study in Iranian highlands

A comprehensive study based on gland and seed micromorphology in Euphorbia (Euphorbiaceae) for species distributed in Iranian highlands is presented. A total of 86 species were studied. The gland structure was examined by direct field observations. Taxonomically important characters of glands were observed and measured: size, texture, shape, color, and horns. For species out of Iran herbarium materials were studied. Seed characteristics were examined using scanning electron microscopy (SEM) as well as dissecting light microscopy. Significant features are: seed size, seed shape, presence of caruncle, shape of caruncle, and seed surface ornamentation. A phylogenetic study using Maximum Parsimony (MP) and Bayesian Inference (BI) was performed based on sequences of nuclear DNA internal transcribed spacers (ITS) for selected species representing the main clades known in Euphorbia and with special focus on the species distributed in Iranian highlands. ITS sequences for 20 accessions representing 19 species are provided for the first time, and 48 accessions of 47 species were used from GenBank. The topologies of both analyses were congruent. The results indicate: (1) four main clades with high supports in subgen. Esula which are appropriate to be recognized at sectional rank. (2) E. larica is nested within clade A including few members of subgen. Rhizanthium and is closely related to sect. Balsamis, which is suggested here to be transferred from subgen. Esula into subgen. Rhizanthium. (3) E. osyridea of the monotypic subsect. Osyrideae is closely related to E. buhsei and to the members of sect. Esula. Tracing morphological characters on the phylogenetic tree shows that several morphological characters, such as seed ornamentation applied in previous subgeneric classification of the subgen. Esula, are homoplasious, but the gland structure and capsule surface characters are more reliable for classification purposes.     

Biogeography and conservation of the genus Ficus (Moraceae) in Mexico

Aim The main objective of this study is to document the biogeographical patterns, endemism and degree of conservation of the species of Ficus (Moraceae) in Mexico. There are over 750 species of the genus Ficus distributed worldwide, and Mexico practically represents its northernmost limit in the American continent. Detailed studies at regional scales may help to understand the biogeography of large genera such as Ficus. Location Mexico. Methods The biogeographical patterns of Mexican Ficus were obtained from information of fig specimens available in two of the main herbaria of Mexico (2140 vouchers), collecting figs throughout this country, and revising the specialized literature. The presence of each species of Ficus was recorded for every one of Mexico's states and several tropical countries of America. Besides, the Mexican territory was divided into cells of 1° × 1° and the presence or absence of all species of the genus was recorded. Rarity of species was classified based on the width of geographic distribution, habitat specificity and population size. Results A total of 21 species of Ficus occur in Mexico, including six species (28.6%) that are endemic to this country. Five species are included in subgenus Pharmacosycea and 16 species are documented under subgenus Urostigma. Affinities of Ficus flora with other tropical countries in America generally decreased as geographical distances from Mexico increased. Mexican states and cells with highest values of Ficus species richness (both total and endemic species) were located. Ten species, including three endemics, presented a wide distribution. Five species, including two endemics, possess the three attributes of rarity (narrow geographical distribution, high habitat specificity and scarce local populations). Three species of Ficus, including the endemic and very rare Ficuslapathifolia (Liebm.) Miq., are not recorded in any protected area existing in Mexico. Main conclusions Most of the Mexican Ficus show a great morphological variation and occupy different habitats along their geographic distribution. The biogeographical patterns described here establish a fundamental scenario for ongoing studies on Ficus–pollinator interactions. However, many local populations are considered to be at risk, as there have been significant reductions in the number and size of local populations. Further studies are needed to understand the process of colonization, maintenance and persistence of fig–pollinator mutualism in species with different patterns of geographic distribution. Mexican Ficus require special policies for conservation due to their complex degree of rarity, particularly their geographic distribution in different types of vegetation, ranging from dry scrublands to tropical rain forests.     

Geographic and taxonomic distribution of a positive interaction: ant-tended homopterans indirectly benefit figs across southern Africa

Although species pairs and assemblages often occur across geographic regions, ecologists know very little about the outcome of their interactions on such large spatial scales. Here, we assess the geographic distribution and taxonomic diversity of a positive interaction involving ant-tended homopterans and fig trees in the genus Ficus. Previous experimental studies at a few locations in South Africa indicated that Ficus sur indirectly benefited from the presence of a homopteran (Hilda patruelis) because it attracted ants (primarily Pheidole megacephala) that reduced the effects of both pre-dispersal ovule gallers and parasitoids of pollinating wasps. Based on this work, we evaluated three conditions that must be met in order to support the hypothesis that this indirect interaction involves many fig species and occurs throughout much of southern Africa and Madagascar. Data on 429 trees distributed among five countries indicated that 20 of 38 Ficus species, and 46% of all trees sampled, had ants on their figs. Members of the Sycomorus subgenus were significantly more likely to attract ants than those in the Urostigma subgenus, and ant-colonization levels on these species were significantly greater than for Urostigma species. On average, each ant-occupied F.sur tree had 37% of its fig crop colonized by ants, whereas the value was 24% for other Ficus species. H. patruelis was the most common source for attracting ants, although figs were also attacked by a range of other ant-tended homopterans. P. megacephala was significantly more common on figs than other ant species, being present on 58% of sampled trees. Ant densities commonly exceeded 4.5 per fig, which a field experiment indicated was sufficient to provide protection from ovule gallers and parasitoids of pollinators. Forty-nine percent of all colonized F. sur trees sampled had ant densities equal to or greater than 4.5 per fig, whereas this value was 23% for other Ficus species. We conclude that there is considerable evidence to suggest that this indirect interaction occurs across four southern African countries and Madagascar, and involves many Ficus species. Received: 11 December 1997 / Accepted: 6 April 1998     

Histological Comparisons of Parasitism by Schistonchus spp. (Nemata: Aphelenchoididae) in Neotropical Ficus spp.

Syconia (enclosed infructescences) infested with host-specific species of Schistonchus (Aphelenchoididae) were collected from six species of Ficus (Moraceae) native to Florida or Panama. They were sectioned and histologically examined to assess the effects of parasitism. Parasitism by Schistonchus spp. was associated with hypertrophied cells, tissue necrosis, and the presence of an exudate in all species. Occasional hypertrophy of the outer epidermal cells occurred on seed florets, wasp florets, and on the endothecial cells of male florets in F. aurea (subgenus Urostigma) from Florida. Aberrations of the inner mesocarp occurred under the hypertrophied cells on seed florets. In F. laevigata (subgenus Urostigma) from Florida, Schistonchus sp. infested immature male florets and was associated with hypertrophy of endothecial cells, epidermal cells of the anther filaments, and anthers. Schistonchus sp. also caused aberrations of the anther filament, anthers, and pollen. Ficus poponoei (subgenus Urostigma) and F. glabrata (subgenus Pharmacosycea), both from Panama, had hypertrophied outer epidermal cells on seed florets. Ficus poponoei also had Schistonchus sp. within the pedicel of an aborted floret, with hypertrophy of the cortical parenchyma. Ficus trigonata (subgenus Urostigma) from Panama had hypertrophy of the outer epidermis of seed florets. When the outer epidermis on these florets was missing, the inner mesocarp was hypertrophied. Ficus maxima (subgenus Pharmacosycea) from Panama had hypertrophy on the outer epidermis of seed and aborted florets. Schistonchus spp. were not found in wasp larvae or pupae in any of the Ficus spp. examined. Hypertrophy was never observed in the absence of Schistonchus spp.     

The affinities of Chenopodium flabellifolium (Chenopodiaceae): Evidence from seed coat surface and flavonoid chemistry

Evidence from scanning electron microscopy of seed coat surfaces and leaf flavonoid chemistry has provided new insights into the relationships of the problematical Chenopodium flabellifolium from San Martín Island, Baja California. The seed surface of C. flabellifolium is basically smooth and thus is essentially the same as members of the subsection Lejosperma of section Chenopodium. This feature distinguishes the species from the taxon that has commonly been considered its nearest relative, namely C. neomexicanum of subsection Cellulata, which has an alveolate seed surface. Leaf flavonoid chemistry also serves to separate C. flabellifolium and C. neomexicanum. Chenopodium flabellifolium has also been viewed as conspecific with C. inamoenum (= C. hians or C. leptophyllum) of subsection Lejosperma. While the seed surface indicates that C. flabellifolium is best treated as a member of this subsection, other morphological as well as chemical data suggest that its closets affinities within the subsection lie with C. fremontii. The leaves of the two species are essentially of the same shape and collectively differ strikingly from those of C. hians and C. leptophyllum. Leaf flavonoid chemistry indicates that C. flabellifolium is closer to C. fremontii than to other taxa of subsection Lejosperma occurring in the western United States because both contain kaempferol 3-O-glycosides, which have not been detected elsewhere. The two species differ consistently in that the former has the pericarp attached to the seed whereas it is separable from the seed in the latter.     

Naemacyclus culmigenus,a newly reported potential pathogen to Miscanthus sinensis,new to Japan

Since the summer of 2010, a discomycete with erumpent apothecia associated with a leaf blight of Miscanthus leaves, were often collected. The morphological characteristics of the fungus suggested it was a member of the Helotiales rather than the Rhytismatales and this was supported by a phylogenetic analysis. Based on a morphological comparison with the type specimen of Naemacyclus culmigenus, currently known from Poaceae (Andropogon and Panicum), it was identified as N. culmigenus, new to Japan. The molecular phylogenetic analysis showed that the generic delimitation of Naemacyclus and related species requires clarification, as does their higher classification within the Leotiomycetes.