A molecular phylogeny of celtidaceae and ulmaceae (Urticales) based onrbcL Nucleotide sequences

On the basis of 1,290 bp sequences of the chloroplast generbcL, a molecular phylogeny of seven of nine genera of the Celtidaceae and four of six genera of the Ulmaceae was produced. These data were analyzed together with some other urticalean genera using three methods (i.e., maximum parsimony, maximum likelihood, and neighbor joining methods). Maximum likelihood topology among 18 trees obtained indicated that the Urticales are monophyletic with its common clade splitting basally into two: one leading to a line comprisingAmpelocera (traditionally placed in Celtidaceae) and Ulmaceae, and the other leading to a line comprising the remaining genera of Celtidaceae, Moraceae, and other Urticales. Ulmaceae, to whichAmpelocera is a sister group, are monophyletic, as supported by many lines of morphological evidence. In contrast to Ulmaceae, the monophyly of Celtidaceae (excludingAmpelocera) was not supported, and resolution of relationships of Celtidaceae with other Urticales, as well as of those within the family, is left for future study.     

Seed coat morphology and evolution in Celtidaceae and Ulmaceae (Urticales)

The seed coat surface morphology of Celtidaceae and Ulmaceae (Urticales) indicates a significant evolutionary diversity.Celtis, Chaetachme andPteroceltis (Celtidaceae) have a unique sculpturing with many crateriform holes; such holes occasionally sparsely occur in seeds ofAphananthe, Gironniera (Celtidaceae) andPlanera (Ulmaceae), but not in those of the nine remaining genera of the two families. The perforated seed coat further occurs in at least some genera of all other urticalean families. A pattern of its occurrence in families and genera suggest that the perforation represents a common archaic feature of all Urticales, rather than a feature derived many times independently within the order. The seed coat of Celtidaceae and Ulmaceae seems to have lately lost the holes probably by a neotenic evolution: one or more times within Celtidaceae, and one time in an ancestral line leading to all Ulmaceae. The derived reticulate seed coat surface sculpturing, which is shared byGironniera (Celtidaceae) and some Ulmaceae, is probably the result of parallel evolution. On the basis of evidence from seed coat morphology and other sources, close relationships ofLozanella, Parasponia andTrema within Celtidaceae, as well as variously distinct positions ofAmpelocera, Aphananthe andGironniera, are also discussed.     

The fatty acid compositions of seed oils and their significance in the taxonomy of the family Ulmaceae

22 kinds of seed oils were extracted from 8 genera of the family Ulmaceae in China The seed oils were examined for their characteristics and fatty acid compositions by gas liquid chromatography. The fatty acid compositions of these oils were found to fall into two classes. Some genera (such as Ulmus, Zelkova) contain mainly lower saturated acids, in which the chief acid is capric acid 10:0, while the genera (such as Celtis, Pteroceltis, Aphananthe, Trema, Gironniera) contain mainly unsaturated acids, in which the chief acid is linoleic acid 18:2. Hemiptelea davidii (Hance) Planch contain however either certain amount of short-chain saturated acids or higher unsaturated acids, it appears a intermediate genus between the two classes. According to the component acids we support that the Ulmaceae be split into two subfamilies. The genera arrangement based on the component acids corresponds basically with the view based on mophological characters and flavonoids found in leaves of Ulmaceae, but there are some discrepancies in certain genera, for example, the Aphananthe should beplaced in Celtoid instead of Ulmoid by the present study.     

Gynoecial vascular anatomy and its systematic implications in Celtidaceae and Ulmaceae (Urticales)

Vasculature in the bicarpellate, pseudomonomerous gynoecium with two distinct styles is examined and compared in all of 15 genera of Celtidaceae and Ulmaceae (Urticales). Gynoecial vasculature is diversified in the families but consistet in a genus or a group of genera. Our observations corroborate the earlier suggestion that Ulmaceae (six genera) basically have three-bundled styles, while Celtidaceae (nine genera) always have one-bundled styles. Comparisons with other Urticales and with Eucommiaceae as an outgroup of Urticales indicate that Celtidaceae are more closely related to Moraceae in sharing one-bundled style (a synapomorphy), rather than to Ulmaceae. This supports a separation of Celtidaceae as a distinct family from Ulmaceaesensu lato. Based on the degree of fusion of major vascular bundles running in the gynoecium, we further distinguish three types of gynoecial vasculature in Ulmaceae and, likewise, three types in Celtidaceae, and discuss evolutionary trends of gynoecial vasculature as well as some generic relationships within the families.     

Karyomorphology and relationships of Celtidaceae and Ulmaceae (Urticales)

Based on karyomorphological features, six (examined in this study) of nine genera of Celtidaceae are divided into three groups: 1)Celtis, Parasponia, Pteroceltis andTrema; 2)Aphananthe; 3)Gironniera, and six genera of Ulmaceae into two: 1)Holoptelea andPhyllostylon; 2)Hemiptelea, Planera, Ulmus andZelkova. The first four genera share the simple chromocenter type at the resting stage andx-10, with all chromosomes with submedian or median centromeres (frequency of chromosomes with subterminal or terminal centromeres 0%, although uncertain inTrema).Aphananthe hasx=13, but resembles the above four genera in other features.Gironniera is distinct from all other Celtidaceae in having the diffuse-complex chromocenter type andx=14, features which occur in Ulmaceae. InGironniera the frequency of chromosomes with subterminal or terminal centromeres is 43%, a proportion similar to those inHoloptelea (36%) andPhyllostylon (58%) of Ulmaceae. All six genera of Ulmaceae havex=14, yetHemiptelea, Planera, Ulmus andZelkova are distinct fromHoloptelea andPhyllostylon (with the simple chromocenter type) in having the diffuse-complex chromocenter type and in predominantly possessing chromosomes with subterminal or terminal centromeres (93%). Evidence from karyomorphology, as well as from other sources, suggests 1) thatAphananthe (x=13) is most distantly related to all genera withx=10 within Celtidaceae, 2) thatGironniera may have a key role for understanding evolutionary relationships between Celtidaceae and Ulmaceae, and 3) thatHoloptelea andPhyllostylon represent derivatives of a line that diverged early from a common ancestor or all Ulmaceae. On the basis of comparisons with other Urticales and the putative outgroups of the order, it is also suggested that the chromosome morphology of Ulmaceae represents the more derived state in Urticales.     

Phylogenetic analysis ofUlmaceae

A phylogenetic analysis of theUlmaceae, Cannabaceae, Barbeyaceae, andBroussonetia of theMoraceae produced nine equally parsimonious trees with 127 steps. TheUlmoideae (Ulmaceae, sensuGrudzinskaya) are a monophyletic group and distinct from theCeltidoideae. The genusAmpelocera occupies an isolated taxonomic position among the celtidoids. The similarity ofAmpelocera to the fossil celtidoid flowerEoceltis of North America suggests thatAmpelocera posesses an archaic suite of characters, and occupies a primitive position among the celtidoids, theCannabaceae and theMoraceae. The relationships among the other celtidoid taxa,Cannabaceae, andBroussonetia are problematic. TheCannabaceae andBroussonetia of theMoraceae are nested within the celtidoids suggesting that this is a paraphyletic group. The close, but unresolved, relationship of the celtidoids to theMoraceae andCannabaceae observed in this analysis, and the appearance of the celtidoids in the fossil record prior to the ulmoids suggests that the evolutionary relationship of the ulmoids and celtidoids may be more distant than current taxonomic treatments reflect.     

Urticalean rosids: circumscription, rosid ancestry, and phylogenetics based on rbcL, trnL-F, and ndhF sequences

To address the composition of the urticalean rosids, the relationships of the component families (maximally Cannabaceae, Cecropiaceae, Celtidaceae, Moraceae, Ulmaceae, and Urticaceae) and analyze evolution of morphological characters, we analyzed sequence variation for a large sampling of these families and various rosid outgroups using rbcL, trnL-F, and ndhF plastid regions. Urticalean rosids are derived out of a lineage including Barbeyaceae, Dirachmaceae, Elaeagnaceae, and Rhamnaceae, with Rosaceae less closely related; thus, they are imbedded within Rosales. Ulmaceae are the sister to all remaining families. Cannabaceae are derived out of a subclade of Celtidaceae; this expanded family should be called Cannabaceae. Cecropiaceae are derived within Urticaceae and are polyphyletic with Poikilospermum derived elsewhere within Urticaceae; this expanded family should be called Urticaceae. Monophyletic Moraceae are sister to this expanded Urticaceae. Support for these relationships comes from a number of morphological characters (floral sexuality, presence or absence of hypanthium, stamen type and dehiscence, pollen pore number, ovule position, and embryo alignment) and chromosome numbers. Most fruit types, in terms of ecological dispersal, are derived independently multiple times and are strongly correlated with habitat.     

A GENETIC SYNDROME AFFECTING LEAF DEVELOPMENT IN PISUM

A recessive foliage mutant of Pisum, designated ‘sinuate leaf’ (sil), was found to have two distinct forms of expression, depending on the background genotype. In an af/af background—wherein leaflets are converted to tendrils—sil/sil plants had adventitious tendrils arising from clefts in the distal portion of the stipule. These adventitious tendrils were morphologically modified, just as were the true tendrils on the same plant, by different allelic combinations at the tl locus. In the standard Af background, sil/sil plants had neither incised stipules nor adventitious tendrils, although they did have undulated and somewhat distorted leaflets and stipules. Because mutant expression was variable in an Af background, classification of segregating populations was uncertain. This uncertainty was removed by taking advantange of pleiotropic effects exerted by sil in the presence of one of the wax mutants, wlo, wb, or wsp. Homozygous wlo plants ordinarily have uniformly waxy stipule surfaces, but when the plants were also homozygous for sil the stipule tips were waxless. Conversely, wb/wb and wsp/wsp plants ordinarily have uniformly waxless stipules, but when wb/wb or wsp/wsp plants were also homozygous recessive for sil the stipule tips were waxy. However, sil had no observable effects of any kind on the stipule tips of plants with stipules reduced in size by the action of st/st. By their individual and combined effects, the foliage mutants used in this study revealed developmental relationships among leaf parts not otherwise evident in non-mutant plants.     

Combined LM and SEM study of the middle Miocene (Sarmatian) palynoflora from the Lavanttal Basin,Austria: part IV. Magnoliophyta 2 – Fagales to Rosales

An ongoing investigation of the middle Miocene (Sarmatian) palynoflora from the Lavanttal Basin continues to show that it contains an extremely rich assemblage of angiosperm taxa. The Fagales to Rosales pollen record documented here contains 34 different taxa belonging to the Betulaceae (Alnus, Betula, Carpinus, Corylus, Ostrya), Fagaceae (Castanea, Fagus, Quercus Groups Cerris, Ilex, Cyclobalanopsis, Quercus/Lobatae), Juglandaceae (Engelhardioideae, Carya, Juglans, Pterocarya), Myricaceae (Morrella vel Myrica), Cannabaceae (Celtis), Elaeagnaceae (Elaeagnus), Rhamnaceae, Rosaceae (Prunus) and Ulmaceae (Cedrelospermum, Ulmus, Zelkova). Two of the pollen types represent extinct genera, Trigonobalanopsis and Cedrelospermum, and are also reported for the first time from the Lavanttal Basin along with pollen of Rhamnaceae and Prunus. The different types of Quercus pollen are now affiliated with Groups Cerris, Cyclobalanopsis, Ilex and Quercus/Lobatae based on sculpturing elements observed using scanning electron microscopy (SEM). Köppen signatures of potential modern analogues of the fossil Fagales and Rosales suggest a subtropical (Cfa, Cwa) climate at lower elevation and subsequent subtropical to temperate climate with altitudinal succession (Cfa → Cfb/Dfa→ Dfb; Cwa → Cwb → Dwb) in the Lavanttal area during accumulation of the palynoflora. Most of the fossil taxa have potential modern analogues that can be grouped as nemoral and/or merido-nemoral vegetation elements, and the diversity of Fagales indicates a varying landscape with a high variety of niches.     

Trichomes and stomata,and their taxonomic significance in theUrticales

Structure and ontogeny of stomata and trichomes have been studied in 23 species and 3 varieties of theUrticales. Stomata are anomocytic, more rarely paracytic; anisocytic and sometimes helicocytic and transitorial types are found inUrticaceae andDorstenia, rarely inArtocarpus. The ontogeny of anomocytic and actinocytic stomata is perigenous, of paracytic either mesogenous or perigenous, of anisocytic either mesogenous or mesoperigenous, and of helicocytic and transitional types mesogenous. Among trichomes eglandular unicellular (wide spread), bicellular or uniseriate filiform (Cannabis); glandular capitate with uni- or bicellular (Moraceae, Ulmaceae, Cannabaceae), uniseriate filiform (Ulmaceae) or multiseriate stalk (Cannabis); sunken glands (Artocarpus); uniseriate glandular with uniseriate stalk (Celtis), and stinging emergences (Urticaceae) have been observed. It is concluded that theUrticales represent a natural order with four families:Ulmaceae, Moraceae, Urticaceae andCannabaceae which are distinct but interrelated with each other.     

Immature Stages of Tomapoderus(T.) ruficollis Fabricius (Coleoptera: Attelabidae) from Korea

Eggs, larvae and pupae of Tomapoderus (T.) ruficollis Fabricius are described and illustrated. The species is found on host plants, Zelkova serrata Makino and Ulmus davidiana var. japonica Nakai. and is a well‐known forest pest. Taxonomic notes and cradle structure of this species are provided.     

榆属（榆科）一新种——绵竹榆

绵竹榆的花秋季开放,翅果柱头面被毛,其两侧的翅较果核为窄,果核位于翅果上端接近缺口处,与榔榆(Ulmus parvifolia Jacq.)相似,但树皮深灰色,不规则鳞块状浅裂,叶片先端渐尖,花被片裂至基部,宿存,边缘上部生纤毛,翅果狭椭圆形,中部最宽,向两端渐变窄,果梗与花被等长,长约2 mm,果序梗长约1 mm,而明显不同。     

Leaf-cutting ants and avoided plants: Defences against Atta texana attack

Summary Leaf-cutting ants (Formicidae; Attini) characteristically never attack some common plant species in their habitats. These plants may be defended against the ants in several ways. In Texas, mature leaves of Sapindus saponaria (Sapindaceae) and Celtis reticulata (Ulmaceae) are unpalatable to Atta texana Buckley foragers, while mature leaves of Berberis trifoliata (Berberidaceae) are palatable to the ants, but are too tough to cut. Young Celtis leaves and and young Berberis leaves are palatable and can be cut by the ants, however. These young leaves may escape attack by remaining palatable a brief amount of time (new Celtis leaves), or by occurring patchily in space and time (new Berberis leaves).     

江苏省地带性植被的基本特点与分布规律

江苏省地跨南暖温带、北亚热带和中亚热带三个生物气候带,典型地带性植被类型依次为落叶阔叶林、落叶常绿阔叶混交林和常绿阔叶林。前两个类型也分别见于北亚热带和中亚热带地区。三个地带性植被类型的主要类型及其分布与组成特点,分别列表概述。落叶常绿阔叶混交林是落叶林与常绿阔叶林之间的过渡类型,在北亚热带地区,林内落叶树的种类与数量往往超过常绿树,落叶层片占优势地位。江苏境内的常绿阔叶林,因地处中亚热带北缘,具有一定的过渡性特征,林内落叶阔叶树的种类往往超过常绿阔叶树,但后者的多度与盖度均占显著优势地位,常绿层片为主,所以外貌为常绿阔叶林。 江苏没有高山,东西间狭窄,故植被的垂直地带性和经度地带性分布规律均未显示。但南北间宽广,跨纬度4?以上,从北向南气候梯度变化明显,所以植被分布的纬度地带性规律明显。     

Leaf Development, Metamorphic Heteroblasty and Heterophylly in Berberis s. l. (Berberidaceae)

Shoot development of temperate and tropical members of Berberis s. l. was examined in order to assess: (1) the homology of the spines along the long shoots and the foliage leaves that form on the short shoots; (2) the occurrence of heterophylly and/or heteroblasty in the genus; and (3) the structural correspondence between cataphylls, spines, and foliage leaves. The 1-5-armed spines have been interpreted as modified compound leaves lacking stipules, as a modified lamina (central spine) with stipules (lateral spines), or less often, as transformed branches, or as epidermal outgrowths. On the other hand, the foliage leaves of the short shoots have been interpreted as leaflets of palmately compound leaves. Our results indicate that there are three distinct leaf types per node: (1) Leaves modified in spines spirally arranged in long shoots; (2) foliage, expanded leaves densely arranged in short shoots; and (3) cataphylls protecting axillary buds. The spines are leaf homologs with a clear distinction between the leaf base with stipules, and a laminar portion modified into the 1-5-armed spine; the lateral spines are not stipules as they arise from the marginal meristem of the laminar portion, and not from the leaf base. The foliage leaves also have stipules flanking the leaf base. Both spiny leaves and foliage leaves develop an articulation between the base and the laminar portion. Cataphylls of the short shoots of Berberis s. str. and those of the reproductive short shoots of Mahonia correspond to the entire leaf base, but those of the renewal (vegetative) shoots of Mahonia are spiny and have an odd vestigial pinnately compound lamina. Heterochrony due to ontogenetic truncation caused by the formation of the terminal inflorescence at the apex of the short shoots could be responsible for the lack of petiole/lamina differentiation in the foliage leaves. The spiny long-shoot/foliose short-shoot system of branching in Berberis s. str. appears to be genetically and phylogenetically fixed and not environment-dependent. This represents a clear example of metamorphic heteroblasty sensu Zotz et al. (Botanical Review 77:109–151, 2011) with further occurrence of heterophylly along the short shoots.     

A new flavonoid from the bark of Ulmus pumila L

A new flavonoid (6), together with eight (1–5 and 7–9) known flavonoids, were isolated from the n-butyl alcohol soluble portion of the EtOH extract of Ulmus pumila L. The chemical structures of the compounds were determined by using spectroscopic methods and further supported by comparison with previously literature values. Among them, flavonoids 4, 6 and 9 were isolated for the first time from the family Ulmaceae. Furthermore, the chemotaxonomic significance of the isolates was also discussed.     

Distribution profile of stomatal conductance and its interrelations to transpiration rate and water dynamics in young maize laminas under sulfate deprivation

Seven-day-old maize (Zea mays) plants were grown hydroponically for 10 days in S-deprived nutrient solution. The distribution profiles according to the position on the stem of the S-deprived laminas’ stomatal conductance, transpiration rate, photosynthetic rate, dry mass, water content, and specific surface area were monitored relative to control among others. Photochemical efficiency of photosystem II remained unaffected by the deprivation, as well as the specific surface area of all but the embryonic laminas after d2. In S-deficient plants, the embryonic (L0) and the uppermost lamina or the one below it presented mostly significant changes. The response ratios (Rr) of the L0 stomatal conductance oscillated; the oscillation started with an increase at d2. The corresponding Rr values of L0 transpiration and photosynthetic rates started oscillating at d4 in the same fashion. At d8, an increasing gradient appeared in water-content Rr values from L1 to the uppermost lamina. At d10, all but the embryonic laminas presented significantly reduced Rr values in water content. Changes in dry mass and surface area of laminas were synchronized. In control, the transpiration rate expressed per DM unit remained constant during the examined period, while under the deprivation it followed a power function of surface area.     

ANGIOSPERM PALEOBIOCHEMISTRY OF THE SUCCOR CREEK FLORA (MIOCENE) OREGON,USA

The organic chemical profiles of fossil Acer and Quercus leaf tissues are presented and correlated with those of previously described fossil Celtis, Ulmus and Zelkova and interpreted in conjunction with referable extant genera. Intrageneric comparisons among fossil and extant taxa indicate that relatively minor phytochemical differences exist suggesting that little flavonoid and steroid evolution since post-Miocene times has occurred. Biosystematic relationships between living North American and Asian genera indicate that in some cases (Quercus, Zelkova) a greater affinity exists between living Asiatic species and elements of the Succor Creek Flora. The chemical data are proposed as an independent parameter in assessing angiosperm biogeography and proposed migration patterns of the Fagaceae and Ulmaceae. The high chemical fidelity seen between some living and fossil genera preserved in ash-fall deposits is ascribed to the reaction of membrane bound lipids with various organic acids and to subsequent rapid dehydration.     

Ulmus davidiana var. japonica Nakai Upregulates Eosinophils and Suppresses Th1 and Th17 Cells in the Small Intestine

The bark of Ulmus davidiana var. japonica Nakai (Ulmaceae) has been used in traditional Korean medicine for chronic inflammation in the gastrointestinal tract. Here we investigated the frequency and cytokine profile of the major immune cells in the small intestinal lamina propria (SI LP), spleen, and mesenteric lymph nodes (MLNs) of mice treated orally with Ulmus davidiana var. japonica Nakai bark water extract (UDE) to address the immunomodulatory role of this herb in intestinal homeostasis. B6 mice were given 5g/kg UDE once daily for 14 days. They were then sacrificed, and cells were isolated from the spleen, MLNs, and SI LP. The proportion of B versus T lymphocytes, CD4⁺ versus CD8⁺ T lymphocytes, Th1 and Th17 cells, and Foxp3⁺ regulatory T cells in the spleen, MLNs, and SI LP were analyzed. The frequency of antigen-presenting cells (APCs), including dendritic cells, macrophages, and eosinophils in the SI LP and the expression of costimulatory molecules on APCs were also evaluated. The numbers and frequencies of Th1 and Th17 cells in the SI LP were significantly reduced in the UDE-treated mice compared with PBS controls. In addition, the proportion of IL-4-producing eosinophils in the SI LP was significantly elevated in the UDE-treated mice compared with controls. Taken together, these data indicate that UDE up-regulates the number and frequency of SI LP eosinophils, which can down-regulate the Th1 and Th17 responses via IL-4 secretion and contribute to intestinal homeostasis.     

TheUlmaceae, one family or two? Evidence from chloroplast DNA restriction site mapping

The Ulmaceae is usually split into two subgroups, referred to as either tribes or more commonly subfamilies (Ulmoideae andCeltidoideae). The two groups are separated, with some exceptions, on the basis of leaf venation, fruit type, seed morphology, wood anatomy, palynology, chemistry, and chromosome number. Propositions to separate the two groups as distinct families have never gained general acceptance. Recent morphological and anatomical data have suggested, however, that not only is family status warranted but thatCeltidaceae are more closely related toMoraceae and otherUrticales than toUlmaceae. In order to test these alternative sets of relationships, restriction site mapping of the entire cpDNA was done with nine rare cutting enzymes using 11 genera ofUlmaceae s. l., three other families of theUrticales, and an outgroup family from theHamamelidae. Cladistic analysis of the data indicates thatUlmaceae s. l. is not monophyletic and that distinct families (Ulmaceae andCeltidaceae) are warranted; thatUlmaceae is the sister group toCeltidaceae plus all other families in the order; and thatCannabaceae might be nested withinCeltidaceae. Familial placements of various problematic genera (e.g.Ampelocera, Aphananthe) are resolved and character evolution of key morphological, anatomical, chemical, and chromosomal features are discussed.