A phylogenetic study of the Plantaginaceae

In a study based on morphological, embryological and chemical data of the Plantaginaceae, within the subclass Sympetalae or Asteridae, the superorder Lamianae is shown to be monophyletic. However, it was not possible to reconstruct the phylogeny within Lamianae or to find a sister-group for the monophyletic Plantaginaceae – Hydrostachyaceae for the latter is rejected. Three or rarely four genera have previously been recognized within Plantaginaceae, but in both cases Plantago appears as polyphyletic, which is considered unacceptable. Six clades are recognized as subgenera within Plantago: subgen. Plantago (c. 131 species), subgen. Coronopus (c. 11 species), subgen. Littorella (three species), subgen. Psyllium (c. 16 species), subgen. Bougueria (one species), and subgen. Albicans (51 species). Within P. subgen. Plantago, the paraphyletic sect. Plantago (c. 42 species) is found in all parts of the world except South America, New Zealand, Australia, and New Guinea, where sect. Oliganthos and sect. Mesembryniae vicariate. It is not possible to infer the phylogeny within sect. Plantago, for example between the endemic species from distant Pacific Islands; their common ancestor might be a species that once had a very wide distribution. Keys to genera, subgenera, sections and series are given. Only one genus, Plantago L., is recognized. The following proposals are made in the Appendix: P. ser. Oliganthos Rahn, ser. nov.; P. ser. Carpophorae (Rahn) Rahn, stat. nov. (= sect. C.); P. ser. Microcalyx (Pilg.) Rahn, stat. nov. (= sect. M.); P. unibrackteata Rahn, nom. nov. (=P. uniflora Hook.f. non L.); P. subgen. Littorella (P. J. Bergius) Rahn, stat. nov. (=Littorella P. J. Bergius); P. araucana Rahn, nom. nov. (=Littorella australis Griseb. non Plantago australis Lam.); P. americana (Fernald) Rahn, comb. nov.(=Littorella a.); P. subgen. Bougueria (Decne.) Rahn, stat. nov. (=Bougueria Decne.); P. nubicola (Decne.) Rahn, comb. nov. (=Bougueria n. Decne.); P. subgen. Albicans Rahn, subgen. nov. Ten lectotypes are selected.     

The Identification of the Seeds and Achenes of the British Plantaginaceae

The morphology of seeds and achenes of the British members ofthe Plantaginaceae has been examined. Keys to the identificationof seeds in the genera Plantago and Littorella are presented.Quantitative data on size and weight, as well as informationabout shape, colour and surface features of the seeds are givenfor: Plantago major L., P. media L., P. lanceolata L., P. martimaL., P. coronopus L., P. indica L., P. sempervirens Crantz, andLittorella uniflora (L.) Aschers. A representative sample ofeach seed is illustrated.     

Evolution of flower shape in <Emphasis Type="Italic">Plantago lanceolata</Emphasis>

Plantago lanceolata produces small actinomorphic (radially symmetric), wind-pollinated flowers that have evolved from a zygomorphic, biotically pollinated ancestral state. To understand the developmental mechanisms that might underlie this change in flower shape, and associated change in pollination syndrome, we analyzed the role of CYC-like genes in P. lanceolata. Related zygomorphic species have two CYC-like genes that are expressed asymmetrically in the dorsal region of young floral meristems and in developing flowers, where they affect the rate of development of dorsal petals and stamens. Plantago has a single CYC-like gene (PlCYC) that is not expressed in early floral meristems and there is no apparent asymmetry in the pattern of PlCYC expression during later flower development. Thus, the evolution of actinomorphy in Plantago correlates with loss of dorsal-specific CYC-like gene function. PlCYC is expressed in the inflorescence stem, in pedicels, and relatively late in stamen development, suggesting a novel role for PlCYC in compacting the inflorescence and retarding stamen elongation in this wind pollinated species.     

中国2种归化植物新记录

报道了发现于辽宁省的2种中国归化植物新记录:白毛马鞭草(Verbena stricta Vent.)和向日葵(Helianthus annuus L.),白毛马鞭草穗状花序粗壮而紧凑,单生枝顶或呈简单的聚伞状或圆锥状排列,且花序轴在果期显著延长,叶卵圆形,无柄,与中国有报道的本属其他物种区别明显。该种原产美国西部至中部地区,近来在辽宁省铁岭市发现有归化。向日葵原产于北美地区,被发现广泛归化于辽宁省大连市,营口市鲅鱼圈也有发现。归化种群形态特征与该地区栽培类型完全不同,其花序极多分枝,头状花序直径较小,瘦果极小,表面纹饰多样性很高。     

Molecular systematics and biogeography of the amphibious genus Littorella (Plantaginaceae)

Littorella (Plantaginaceae) is a disjunct, amphibious genus represented by three closely related species. Littorella uniflora occurs in Europe including Iceland and the Azores, L. americana is found in temperate North America, and L. australis grows in temperate South America. Littorella has been recognized in numerous floristic treatments, but its status as a genus has recently been questioned. Rahn (Botanical Journal of the Linnean Society 120: 145-198, 1996) proposed a new phylogeny for Plantaginaceae based on morphological, embryological, and chemical data in which he reduced Littorella to a subgenus of Plantago. This article compares the phylogeny proposed by Rahn to one based on DNA sequence data from the internal transcribed spacer (ITS) region. In our analysis, Littorella forms a strongly supported monophyletic clade sister to Plantago and its recognition at the generic rank appears warranted. Littorella australis is sister to L. americana, and this clade is sister to the European L. uniflora. This more distant relationship between L. uniflora and L. americana provides support for maintaining both taxa at the specific rank and suggests a European origin for Littorella. Our studies also indicate that the monotypic genus Bougueria is deeply nested within Plantago and that its inclusion within Plantago as proposed by Rahn appears justified.     

NEW WORLD JUGLANDACEAE: A NEW SPECIES OF ALFAROA FROM MEXICO

Alfaroa mexicana is described as new. Certain aspects of the wood anatomy, inflorescence, pollen, staminate flower, and pistillate flower morphology are compared with previous findings on the three other species of Alfaroa and representatives of the closely related genus Engelhardia.     

Genus Colocasiomyia (Drosophilidae: Diptera) in Sabah,Bornean Malaysia: High species diversity and use of host aroid inflorescences

A total of 21 Colocasiomyia species, including 17 undescribed species, are reported from Sabah (Mt. Kinabalu and neighboring areas), Malaysia, based on samples collected from inflorescences of 14 or 15 Araceae species. This number of species is the largest as a local fauna of this genus in the world. The high species diversity is attributed to 12 undescribed species belonging to the Colocasiomyia baechlii species group. A particular breeding habit of Colocasiomyia is sharing of the same inflorescence by a pair of species, with partial niche separation between them: one species uses exclusively the pistillate (lower female‐flower) section of the spadix for oviposition and larval development, whereas the other mostly uses the staminate (upper male‐flower) section. However, the baechlii group species show quite different patterns of host plant use: many (up to eight) species cohabit on the same inflorescence. It is unlikely that they separate their breeding niches micro‐allopatrically within an inflorescence. Instead, species composition and their proportions of individual numbers vary among different localities, seasons and host plants, with partial overlap among them. Such partial separations in local distribution, phenology and host selection would in combination contribute to their coexistence and promote the species diversity of this group.     

The inflorescence structure of Campylotropis (Leguminosae)

Racemose inflorescences were investigated organographically and ontogenetically in three species of Campylotropis. An additional phyllome was found to be borne at the base of the pedicel in C. hirtella and C. giraldii but not in C. polyantha. Organographic observations revealed that the additional phyllome is the prophyll on the reduced lateral branch borne in the axil of the phyllome on the central inflorescence axis, and that the additional phyllome subtends the pedicel terminating in a flower. The flower presents its abaxial side to the additional phyllome in accordance with the organographic observations. Ontogenetic observations, moreover, revealed the presence of a rudimentary apex of the reduced branch in earlier stages of development. On the basis of such evidence, the central inflorescence axis of Campylotropis is interpreted as being composed of reduced lateral branches each of which usually bears only one flower in the additional phyllome axil. The inflorescence structure is kept in spite of the loss of the additional phyllome. In comparison with allied genera, the inflorescence of Campylotropis is regarded as a reduced pseudoraceme. An evolutionary trend in the inflorescence structure of the genus is inferred.     

THE ALLIUM INFLORESCENCE: SOME SPECIES OF THE SECTION MOLIUM

Mann , Louis K. (U. California, Davis.) The Allium inflorescence: some species of the section Molium. Amer. Jour. Bot. 46(10): 730–739. Illus. 1959.—The inflorescence is described for Allium neapolitanum, A. roseum, A. hirsutum, A. subhirsutum, A. zebdanense and A. erdelii, species of Molium, a Mediterranean section of the genus. Inflorescence structure is similar among these 6 species. The single spathe appears to consist of 4 coalesced bracts, each of which bears in its axil a bostryx (helicoid cyme) of 3–7 flowers. Central to these 4 peripheral bostryxes are several smaller ones which differentiate later and decrease in flower number toward the inflorescence center. Bracts and bracteoles (prophylls) are generally absent within the spathe. The inflorescence is not radially symmetrical and the bostryx on the side opposite the uppermost foliage leaf differentiates and develops first. Among the peripheral bostryxes there is a definite sequence of development and certain regularities of coiling (clockwise vs. counterclockwise). Within each bostryx the flowers open in a strict sequence from oldest to youngest; among the bostryxes, the 4 peripheral ones flower first, usually starting with the first bostryx to be differentiated. The central bostryxes flower last.     

POLLEN STUDIES IN THE GENUS CORDIA (BORAGINACEAE)

The pollen of 40 species of Cordia L. was studied by light microscopy and scanning and transmission electron microscopy. Three major types were found, 3-porate with a reticulate sexine, 3-colpate to 3-colporoidate with a striatoreticulate sexine, and 3-colpate to 3-colporate with a spinulose sexine. These pollen types are correlated with significant differences in the inflorescence, flower, and fruit structures found in the sporophyte. The following revisions are suggested and discussed: that Cordia sect. Varronia P. Br. be elevated to generic status; that Cordia sebestena L. and a few close allies be treated as generically distinct; and that Cordia myxa L. be conserved as the lectotype for the genus, thus continuing a wide application of the epithet Cordia.     

A community of submerged aquatic CAM plants in Lake Kalgaard,Denmark

The CO₂ exchange pattern of leaves of dominant Isoetes lacustris L., Littorella uniflora (L.) Aschers. and Lobelia dortmanna L. from oligotrophic, carbon-poor Lake Kalgaard, Denmark was examined by gas-exchange experiments and by following the diurnal acidity rhythm. Both variables suggest Crassulacean Acid Metabolism (CAM) in Littorella and Isoetes. During the dark, both species had a continuous, but declining CO₂ uptake. The maximum uptake in Littorella was 28 μmol CO₂ g^?1 DW h^?1 or about 36% of the light CO₂ assimilation rate. In Isoetes, corresponding figures were 19 μmol CO₂ g^?1 DW h^?1 and 22%.A peak of 49.5 μEq g^?1 FW titratable acidity (Littorella) and 41.9 μEq g^?1 FW (Isoetes) was found at the end of the dark period, reflecting an increase of 11.6 μEq g^?1 FW (9 h)^?1 (Littorella) and 16.3 μEq g^?1 FW (9 h)^?1 (Isoetes). These results agreed with the cumulative CO₂ assimilation during the dark.The assimilation of CO₂ in the dark is important, apparently, for growth of Littorella and Isoetes in Lake Kalgaard. Lobelia, which exhibits little dark CO₂ assimilation, is also the least abundant species, at less than 1% of the total cover.     

Organographic and ontogenetic studies on the inflorescence ofLespedeza cuneata (Dum.-Cours.) G. Don (Leguminosae)

Structure of inflorescence and its variation were organographically and ontogenetically studied inLespedeza cuneata (Dum.-Cours.) G. Don. An axillary inflorescence of the species forms a compound inflorescence which is composed of three or four component inflorescences. Each component inflorescence bears four (rarely six), three, two, or one flowers. Based on the arrangement of inflorescence phyllomes, the component inflorescence with four flowers is interpreted as a pseudoraceme bearing two shortened lateral shoots (partial inflorescences) each of which has two flowers. The component inflorescence with one flower appears to be terminated by the flower and to compose the cyme. Organographic observations revealed that the terminally located flower is not truly terminal, but axillary in origin. Ontogenetic observations showed that the apices of component inflorescence and partial inflorescence exist in early developmental stages in spite of variation in the form of component inflorescence. The terminally located flower in the cyme-like inflorescence was thus demonstrated to be laterally borne on the partial inflorescence axis. The component inflorescence composing the cyme-like one inL. cuneata is a reduced form in the number of partial inflorescences and of flowers from the pseudoraceme. The cyme-like inflorescence inL. cuneata resembles the inflorescence ofKummerowia.     

Fruit production and floral traits: correlated evolution in <Emphasis Type="Italic">Govenia</Emphasis> (Orchidaceae)

The most common estimate of reproductive success in orchids is usually fruit set. Factors such as resource limitation and certain floral traits may influence reproductive success in animal-pollinated plants. Correlated evolution of reproductive success vs. seven floral traits (inflorescence length, flower number, flower distribution along inflorescence, dorsal sepal length, lateral sepal length, flower color, and column position) was studied in eight species of Govenia. Taxa represented three lineages in the genus. Independent contrasts were calculated on a phylogeny inferred from chloroplast (trnL-F IGS) DNA sequences, and a correlation test and multiple regression were then performed. Two data sets were evaluated, one including all eight species and another excluding G. utriculata, which is autogamous. The historical analyses showed that there is a correlation between reproductive success and dorsal sepal length, column position, and flower number, these correlations suggest that changes in these floral traits usually accompany evolutionary shifts in reproductive success. Multiple regression tests suggest that changes in reproductive success can be explained by shifts in flower number, inflorescence length, column position and by dorsal sepal length. When phylogeny is taken into account, our analyses showed that evolutionary shifts in these floral traits were correlated with changes in reproductive success. Evolutionary correlation between reproductive success and floral traits might be explained by the natural selection of certain floral phenotypes by pollinators.     

Molecular phylogeny and evolution of inflorescence types in Eperua

Premise

The Amazonian hyperdominant genus Eperua (Fabaceae) currently holds 20 described species and has two strongly different inflorescence and flower types, with corresponding different pollination syndrome. The evolution of these vastly different inflorescence types within this genus was unknown and the main topic in this study.

Methods

We constructed a molecular phylogeny, based on the full nuclear ribosomal DNA and partial plastome, using Bayesian inference and maximum likelihood methods, to test whether the genus is monophyletic, whether all species are monophyletic and if the shift from bat to bee pollination (or vice versa) occurred once in this genus.

Results

All but two species are well supported by the nuclear ribosomal phylogeny. The plastome phylogeny, however, shows a strong geographic signal suggesting strong local hybridization or chloroplast capture, rendering chloroplast barcodes meaningless in this genus.

Conclusions

With our data, we cannot fully resolve the backbone of the tree to clarify sister genera relationships and confirm monophyly of the genus Eperua. Within the genus, the shift from bat to bee and bee to bat pollination has occurred several times but, with the bee to bat not always leading to a pendant inflorescence.

     

Pollination mutualism between a new species of the genus Colocasiomyia de Meijere (Diptera: Drosophilidae) and Steudnera colocasiifolia (Araceae) in Yunnan, China

A new species of the genus Colocasiomyia de Meijere (Diptera: Drosophilidae) was discovered from inflorescences of Steudnera colocasiifolia K. Koch (Araceae) in Yunnan, China. The new species is described as Colocasiomyia steudnerae Takenaka and Toda, sp. nov., and we investigated the reproductive ecology of both the fly and the plant species. This fly species reproduces in the inflorescences/infructescences of the plant, and depends almost throughout its entire life cycle on the host plant. The fly species is the most abundant flower visitor for S. colocasiifolia and behaves intimately with the flowering events, suggesting that it is the unique and most efficient pollinator for the host plant. Bagging (insect‐exclusion) treatment of inflorescences resulted in no fruits. These findings strongly suggest that intimate pollination mutualism has evolved between the fly and the host plant, as are known in other Colocasiomyia flies and Araceae plants. One notable feature of this system is that the new species almost monopolizes the host‐plant inflorescence as a visitor, without any cohabiting Colocasiomyia species. In comparison to other cases where two Colocasiomyia species share the same inflorescence and infructescence of Araceae host plants for reproduction by separating their breeding niches microallopatrically between the staminate (upper male‐flower) and the pistillate (lower female‐flower) regions on the spadix, C. steudnerae exhibits a mixture of stamenicolous and pistillicolous breeding habits.     

A revision of Gibsoniothamnus L. O. Williams (Schlegeliaceae)

Gibsoniothamnus L. O. Williams is a genus of 11 species native to Mexico, Central America, and northern Colombia. The species are shrubs or small trees with opposite, entire leaves, tubular corollas, four stamens, 2-locular ovary, and fleshy, indehiscent fruits. Calyces are unlobed, or they may have lateral, triangular or winglike lobes. Most species are epiphytes and appear to be bird pollinated and bird dispersed.     

ISOZYME DIVERGENCE BETWEEN EASTERN ASIAN,NORTH AMERICAN,AND TURKISH SPECIES OF LIQUIDAMBAR (HAMAMELIDACEAE)

The deciduous woody genus Liquidambar has four morphologically similar species in eastern and western Asia, eastern North America, and Central America. Liquidambar styraciflua is found in the eastern United States and Central America, L. orientalis is native only to southwest Turkey, and L. formosana and L. acalycina occur in eastern Asia. This genus is one of many that contributes to the floristic similarities observed between these different regions. Allelic variation was scored at 22 isozyme loci from 41 populations. The level of genetic divergence between species on different continents is high. Nei's genetic identity was 0.431 between L. formosana and L. styraciflua, 0.485 between L. acalycina and L. styraciflua, 0.512 between L. orientalis and L. styraciflua, 0.256 between L. formosana and L. orientalis, and 0.305 between L. acalycina and L. orientalis. Estimates of time of divergence from the isozyme data suggest that the current species diverged before or during the Miocene. The pattern of relationships portrayed by the isozyme data suggest a longer period of separation between the eastern and western Asian forms of this genus. In addition, the eastern North American and Turkish species appear to be the most closely related intercontinental pair of species providing evidence for a North Atlantic land bridge as late as the Miocene. It would appear, therefore, that the North American populations were in contact with the Asian populations over the North Pacific and North Atlantic possibly as late as the Miocene, but that the separation between the two Asian populations occurred much earlier. The time of divergence as measured from the isozyme data correlates with an independent assessment of the origin of these disjuncts as determined from the fossil record.     

Seven new species of Miliusa (Annonaceae) from Thailand

Seven new species of the genus Miliusa are described from Thailand (M. fragrans, M. hirsuta, M. intermedia, M. nakhonsiana, M. sessilis, M. thailandica, and M. umpangensis). A key to the 19 species of Miliusa in Thailand is provided. In addition, the complete taxonomic nomenclature of all known species of Miliusa in Thailand is given, with several new proposed synonyms. The new as well as the known species of Miliusa in Thailand are classified into four morphological groups on the basis of a combination of flower and/or inflorescence position and inner petal morphology proposed earlier.     

TRIGONOBALANUS (FAGACEAE): TAXONOMIC STATUS AND PHYLOGENETIC RELATIONSHIPS

The fagaceous genus Trigonobalanus as recently treated includes 3 species, two in Malaysia and Southeast Asia and a single species in Colombia, South America. Character analysis suggests that the genus as currently circumscribed is paraphyletic, without synapomorphies to unite the three species. Each of the three species is a morphologically distinct relict of a group that probably was ancestral to the modern genera Quercus and Fagus. Each of the three species also has at least one autapomorphy which is unique within Fagaceae. Analysis of cupule morphology in Fagaceae provides an interpretation of evolution in cupules which differs substantially from Forman's interpretation. We interpret trigonobalanoid cupules as indicative of an ancestral type of inflorescence within Fagaceae. This inflorescence type is a dichasial structure in which the outermost axes are cupular valves, but the degree of branching and subsequent number of fruits are variable. Following this model, a strict relationship exists between valve number and fruit number as seen in cupules of Trigonobalanus (valves = fruits + 1). Fossil evidence is consistent with our interpretation of the phylogenetic position of the trigonobalanoids. We propose to segregate the three species of Trigonobalanus as three monotypic genera; two of these require names which we provide here: Formanodendron and Colombobalanus.