When looks can kill: the evolution of sexually dimorphic floral display and the extinction of dioecious plants

Dioecious plants (with separate male and female individuals) more often have drab, inconspicuous flowers than related bisexual plants. Models indicate, however, that similar conditions favour the evolution of showy floral displays in dioecious and bisexual plants. One difference, however, is that dioecious plants may evolve floral displays that are sexually dimorphic. We show that males are more likely to evolve showy flowers than females in animal-pollinated plants, especially when pollinators are abundant. We demonstrate that this dimorphism places showy dioecious plants at a much higher risk of extinction during years of low pollinator abundance because pollinators may fail to visit female flowers. The higher extinction risk of showy dioecious plants provides an explanation for the fact that dioecious plants that do persist tend to have inconspicuous flowers and are more often wind pollinated. It may also help explain why dioecious plants are less species-rich than related bisexual plants.     

Foliar architecture of vanilloid orchids: insights into the evolution of reticulate leaf venation in monocotyledons

Leaves of representative taxa within the vanilloid clade of Orchidaceae were cleared and their venation patterns studied. Within subtribe Pogoniinae, Isotria and Pogonia exhibit a prominent reticulate venation pattern, although only Pogonia ophioglossoides is characterized by free vein endings. Within subtribe Vanillinae, all species of Epistephium , as well as the New Caledonian endemics, Eriaxis rigida and Clematepistephium smilacifolium , have reticulate-veined leaves that are characterized by numerous free vein endings. Leaves of most species of Vanilla exhibit a parallel-veined pattern stereotypical of monocots, although branched secondaries with free endings were observed in V. africana. Most members of subtribe Galeolinae are 'saprophytes' with reduced bract-like leaves. Vascular bundles enter these leaf-homologues in a parallel manner, but quickly ram+ throughout the tissue in an irregular manner. Leaf venation is used to hypothesize patterns of generic relationships within the vanilloid clade. Molecular evidence for phylogenetic relationships among angiosperms indicates that reticulate leaf venation has arisen independently in several unrelated monocot families, including the vanilloid Orchidaceae, perhaps by a similar evolutionary scenario.     

NONFRUITING HERMAPHRODITIC FLOWERS OF CALOCHORTUS LEICHTLINII (LILIACEAE): POTENTIAL REPRODUCTIVE FUNCTIONS

The breeding system, flowering display, and fruiting pattern of Calochortus leichtlinii Hook. (Liliaceae) was studied in the eastern Sierra Nevada. This species is capable of both autogamy and insect-mediated outcrossing. Either of these two means of pollination alone could affect full seed set; the populations studied were not pollination limited in 1983. Most individuals produced two flowers but set only one fruit. Flowers are not selectively matured on the basis of the quantity or purported quality of the pollen they receive; the first flower to open was matured in all experiments. Nonfruiting flowers have at least three potentially important reproductive functions: disseminating pollen, allowing plants to produce more seeds in response to an abundance of soil moisture, and setting seeds if the first flower fails to develop fruit.     

Micropropagation of Sego Lily

The Sego lily (Calochortus nuttallii T. & G.) produces showy flowers and is a wild species indigenous to the western United States. A 3- to 5-fold increase in shoots per month was achieved when basal sections of bulbs were cultured on Schenk and Hildebrandt medium containing 88 mM sucrose and 8.9 μM 6-benzyladenine (BA) and subcultured every 28 d on the same medium but containing 88 mM sucrose, 2.2 μM BA, and 1.0 μM indole-3-butyric acid. Roots formed from 94% of shoots cultured at 13 °C on medium containing 88 mM sucrose and 2.7 μM α-naphthalene acetic acid. Bulbs formed at the base of shoots cultured on medium containing 263 mM sucrose. This revised version was published online in June 2006 with corrections to the Cover Date.     

Plant distribution and dispersal mechanisms at the Hassayampa River Preserve, Arizona, USA

This study focuses on the relationships between plant dispersal syndromes and plant distributions at the community scale. Species composition and cover are reported from 29 10 × 20-m vegetation plots along five topographic cross-sections in the riparian zone of the Hassayampa River Preserve, Arizona. We find that spatial patterns of dispersal guilds vary within the flood plain of this semiarid region river. Our main results are: (1) wind-dispersed species are fairly evenly distributed at all elevations and distances from the river, whereas cover of animal-dispersed species increases with elevation above, and at greater distances from, the river; (2) wind-dispersed species are proportionally more abundant in the pioneer Populus–Salix community, whereas plants in the late-seral Prosopis community are predominantly animal-dispersed; (3) most of the species classified as obligate-wetland and facultative-wetland are wind-dispersed, whereas facultative-upland and obligate-upland species are mostly animal-dispersed; and (4) there are significantly fewer wind-dispersed species in areas of high total vegetation cover. These results may reflect successional patterns resulting from periodic flooding. Low areas close to the river flood more frequently and with greater intensity than areas farther from the river. Many pioneer species that establish in disturbed areas are wind-dispersed. Over successional time, pioneer species cede to more drought tolerant species that are predominantly animal-dispersed.     

百合目叶绿体rbcL基因核苷酸替代速率检测

分析了百合目主要类群叶绿体中编码核酮糖1,5二磷酸羧化氧化酶大亚基rbcL基因的42条序列,使用RRTree相对速率检测方法,详细研究rbcL基因在百合目7科间同义替代速率和非同义替代速率的变化.相对速率检测显示:百合目内秋水仙科(Colchicaceae)的同义替代速率和非同义替代速率均最快,金梅草科(Campynemat-aceae)同义替代速率最慢,百合科(Liliaceae)的非同义替代速率最慢,但在百合目各科间,无论同义替代速率还是非同义替代速率差异均不显著.     

单子叶植物高级分类阶元系统演化: matK、rbcL和18S rDNA序列的证据

基于两个叶绿体基因（matK和rbcL）和一个核糖体基因（18S rDNA）的序列分析,对代表了基部被子植物和单子叶植物主要谱系分支的86科126属151种被子植物（单子叶植物58科86属101种）进行了系统演化关系分析。研究结果表明由胡椒目Piperales、樟目Laurales、木兰目Magnoliales和林仙目Canellales构成的真木兰类复合群是单子叶植物的姐妹群。单子叶植物的单系性在3个序列联合分析中得到98％的强烈自展支持。联合分析鉴定出9个单子叶植物主要谱系（广义泽泻目Alismatales、薯蓣目Dioscorcales、露兜树目Pandanales、天门冬目Asparagalcs、百合目Liliales、棕榈目Arecales、禾本目Poales、姜目Zingiberales、鸭跖草目Commelinales）和6个其他被子植物主要谱系（睡莲目Nymphaeales、真双子叶植物、木兰目、樟目、胡椒目、林仙目）。在单子叶植物内,菖蒲目Acorales（菖蒲属Acorus）是单子叶植物最早分化的一个谱系,广义泽泻目（包括天南星科Araceae和岩菖蒲科Toficldiaccae）紧随其后分化出来,二者依次和其余单子叶植物类群构成姐妹群关系。无叶莲科Petrosaviaceac紧随广义的泽泻目之后分化出来,无叶莲科和剩余的单子叶植物类群形成姐妹群关系,并得到了较高的支持率。继无叶莲科之后分化的类群形成两个大的分支：一支是由露兜树目和薯蓣目构成,二者形成姐妹群关系：另一支是由天门冬目、百合目和鸭跖草类复合群组成,三者之间的关系在单个序列分析和联合分析中不稳定,需要进一步扩大取样范围来确定。在鸭跖草类复合群分支内,鸭跖草目和姜目的姐妹群关系在3个序列联合分析和2个序列联合分析的严格一致树中均得到强烈的自展支持,获得的支持率均是100％。但是,对于棕榈目和禾本目在鸭跖草类中的系统位置以及它们和鸭跖草目-姜目之间的关系,有待进一步解决。值得注意的是,无叶莲科与其他单子叶植物类群（除菖蒲目和泽泻目外）的系统关系在本文中获得较高的自展支持率,薯蓣目和天门冬目的单系性在序列联合分析中都得到了较好的自展支持,而这些在以往的研究中通常支持率较低。鉴于菖蒲科和无叶莲科独特的系统演化位置,本文支持将其分别独立成菖蒲目和无叶莲目Petrosavialcs的分类学界定。     

动物对松属植物种子的传播作用研究进展

松属植物约110种,根据种子传播方式可分为风传播松和动物传播松。风传播松占绝大多数,种子多具有适应风力的翅。动物传播松大约23种,都具有大、可食用、无翅或短翅的种子,无法借助风力传播。动物传播松的分布生境多为贫瘠的山地,而且多位于高海拔地区。目前已知9种松树的动物传播种类,其余14种可推测为动物传播。动物传播者包括鸦科鸟类和啮齿类动物,动物将获得的种子分散贮藏,未被重取的种子可能萌发,完成传播。动物传播是定向传播,微生境多适合种子萌发。啮齿类的传播距离可达数10 m,而鸦科鸟类的传播距离可达数公里。动物传播的松树会出现树丛和多树干现象,一般由同一贮点内贮藏的多粒种子萌发造成的。动物贮藏的种子大部分被重取,称传播后取食。一些具有大种子的风传播松在种子落地后,啮齿类和鸟类会再次埋藏而形成二次传播,可看做是一个单独的传播类型,即风-动物传播松。动物传播者与依赖传播松树之间可看作是互利共生关系。     

Liana diversity and reproductive attributes in two tropical forests in Mexico

We describe liana diversity and variation in morphology of flowers and diaspores in two tropical forests in Mexico: a seasonally deciduous forest at Chamela, Jalisco and a lowland rain forest at Chajul, Chiapas. Flowers were classified as inconspicuous (1 cm in length and white or pale green flowers) or conspicuous (> 1 cm and brightly colored flowers). Dispersal syndrome was classified as anemochory, barochory, and zoochory. We recorded a higher number of lianas species in Chajul (128 spp.) than in Chamela (71 spp.). In both sites, families with the highest number of species were Bignoniaceae, Leguminosae, Sapindaceae, and Malphigiaceae, and almost half of the liana species had inconspicuous flowers. Most of the species had an abiotic syndrome of dispersal with 40 (56.3%) and 71 (55.5%) wind-dispersed species in Chamela and Chajul, respectively. Zoochory was significantly associated with inconspicuous flowers and anemochory with conspicuous flowers. Our results suggest that (1) flower morphology and dispersal type are not related with the amount of rainfall and (2) lianas are more prone to be wind-dispersed.     

Effects of distinct types of disturbance on seed rain in the Atlantic forest of NE Brazil

In disturbed sites, some groups of seeds might be excluded from the seed rain due to their dispersal modes or seed size, and some groups might be successful as a result of disturbance effects. In the present study, we examined the seed rain in natural treefall gaps and in an area of regenerating forest following an accidental burning, which occurred 4 years before this study. Both of these disturbed areas were compared with nearby forest understorey. The number of seeds, number of species, and proportion of wind-dispersed seeds were compared between these disturbed and undisturbed areas. The treefall gaps have received lower numbers of seeds and species than the nearby understorey, but the number of wind-dispersed seeds did not differ between these areas. The lowest seed number observed in treefall gaps can be attributed to a lower number of animal-dispersed seeds, suggesting that animals may be avoiding treefall gap areas. A higher number of seeds and a lower number of species were observed in the burned area when compared to the adjacent understorey. The high number of small-sized seeds and of wind-dispersed seeds in the burned area was almost surely a consequence of the local production of the pioneer plants established after the burning. In this study, substantial differences were observed in the characteristics of the seed rain at disturbed sites, when compared with undisturbed understorey. However, these two distinct types of disturbance showed quite differing patterns, as treefall gaps received lower number of seeds while the burned area received a higher number of seeds, with a greater proportion of wind-dispersed seeds. The exception was for species richness, which was quite low at both these disturbed sites.     

Leaf flavonoid patterns in the genus Tricyrtis (Tricyrtidaceae sensu stricto, Liliaceae sensu lato)

In a leaf flavonoid aglycone survey of 19 Tricyrtis species, the flavone lulcolin was detected in 16% of the taxa and the flavonols myricetin, qucrcetin, kaempfcrol and isorhamnetin in 32%, 63% and 68% and 58% of the sample, respectively. Kaempfcrol occurred more frequently than qucrcetin in sections Hirtae and Tricyrtis. Section Brachycyrtis was characterized by the presence of lulcolin, while flavoncs were absent from sections Flavae, Hirtae and Tricyrtis. Myricetin was found only in T J'ormosana and T. hirta (section Hirtae) and in T affirm and T. macropoda (section Tricyrtis). This is the first report of myricetin in Tricyrtis and in the Liliales. Also the co-occurrence of flavones and flavonols within the same plant, as in 7. ishiiana var. surugensis and T macrantha , is a rare event in this order. The flavonoid data are congruent with data from morphology, anatomy and molecular analysis in indicating the separation of section Brachycyrtis from the other three sections and in supporting a close relationship between sections Hirtae and Tricyrtis.     

The radiation of Calochortus: generalist flowers moving through a mosaic of potential pollinators

Few studies have surveyed the insects visiting flowers at many sites and for many related species. Calochortus , a genus of about 60 species with its center of diversity in California, has flowers with a wide range of appearances. Insects visiting Calochortus flowers in 25 species were censused at 40 sites. Visitors at coflowering plants were also censused at each site. Calochortus flowers attracted insects in several orders, especially beetles that feed at glands on the petals and bees that often collect pollen. Species in Calochortus section Calochortus had a somewhat narrower spectrum of visitors than species in section Mariposa . The diversity of visitor species at Mariposa flowers was often greater than the diversity at coflowering species. On the whole, Calochortus flowers seem to be generalists in terms of their pollinators. Differences in visitor assemblage were greater between Calochortus populations at different sites than between visitors to Calochortus versus coflowering species at a site. Nevertheless, there were always significant differences in the proportions of various insects at Calochortus versus coflowering plants. In addition, proportions of visitors also differed between species of Calochortus at the same site. Thus, Calochortus flowers have diverged in the visitors they attract, and evidently they have done so without permanently specializing (except to a limited degree at the sectional level). What we know of the radiation of Calochortus lilies is consistent with an interpretation of adaptive wandering through a spatiotemporal mosaic of pollinator communities.     

旅游开发对景观边缘植物溢出效应的影响

人类活动导致的景观改变能形成大量的生境缀块和边缘结构,对植物扩散产生重要影响。以植物个体运动生态学——种子扩散模式作为分类依据,对兴凯湖自然保护区游憩带与非游憩带、交通廊道与非交通廊道中动物扩散物种、风扩散物种、无助力扩散物种(包括重力扩散、弹射扩散等短距离扩散的物种)以及全部物种的边缘溢出效应分别进行对比分析。结果显示,游憩带动物扩散物种和全部物种的边缘溢出效应均明显弱于非游憩带;而风扩散物种在游憩带和非游憩带均有大量溢出;无助力扩散物种在这两个实验区的溢出效应均不明显。交通廊道中动物扩散物种的溢出效应显著弱于非交通廊道,但风扩散物种却明显强于非交通廊道;无助力扩散物种在两种廊道类型中同样只有少量溢出,且距离相对较短;总体溢出效应并未发现显著差异。这些结果表明,旅游开发对植物运动生态学产生了影响,最终导致了溢出效应的改变。     

A Phylogenetic Analysis of the Plastid matK Gene with Emphasis on Melanthiaceae sensu lato

Abstract: The presented mat K tree primarily agrees well with the previously presented rbc L tree and combined rbc L + atp B + 18SrDNA tree. According to the mat K tree, the monocotyledons are monophyletic with 100 % bootstrap support. Acorus diverges first from all other monocotyledons (90 % bootstrap support) in which two major clades are recognized: one (89 %) consisting of Alismatanae and Tofieldia (Nartheciaceae), and the other (< 50 %) comprising Lilianae, Commelinanae and Nartheciaceae other than Tofieldia. Within the latter major clade, Petrosavia and Japonolirion (Nartheciaceae) (82 %) diverge first from the remaining taxa (< 50 %) in which two clades are formed: one (81 %) consisting of Pandanales, Dioscoreales and Nartheciaceae-Narthecioideae, and the other (< 50 %) comprising Liliales, Asparagales and Commelinanae. In the former clade, Dioscoreales and Narthecioideae are grouped together (88 %). In the latter clade, Asparagales and Commelinanae are grouped together (< 50 %). Differences between the mat K and rbc L tree topologies appear in the positions of Tricyrtis (Calochortaceae) and Dracaenaceae. Differences between the mat K and combined rbc L + atp B + 18SrDNA tree topologies exist in the positions of the Petrosavia-Japonolirion pair (Nartheciaceae) and Pandanales. The stop codon position of the mat K gene appears to be highly variable among the monocotyledons, especially in the Liliales.     

Anatomy and morphology of Behnia (Behniaceae) and its relationships within Lilianae: Asparagales

The vegetative and floral anatomy, morphology, embryology and seedlings of Behnia reticulata (Thunb.) Didr. are described, and the relationships of the recently described endemic southern African monocot family Behniaceae Reveal are discussed. Recent molecular studies place the family with the Agavaceae, Hostaceae, Anthericaceae, Herreriaceae and Ane-marrhenaceae. Behnia shows some morphological similarities with this clade, such as the woody shrub habit, a secondary thickening meristem in the rhizomes, small flowers and baccate fruits; but differs in several features, such as reticulate leaf venation, cryptodioecious flowers and seeds lacking phytomelan. Embryologically, the Behniaceae agree with the other members of their clade, although there are no clear synapomorphies by which the group may be defined from other Asparagalean clades.     

Comparative evolution of flower and fruit morphology

Angiosperm diversification has resulted in a vast array of plant morphologies. Only recently has it been appreciated that diversification might have proceeded quite differently for the two key diagnostic structures of this clade, flowers and fruits. These structures are hypothesized to have experienced different selective pressures via their interactions with animals in dispersal mutualisms, resulting in a greater amount of morphological diversification in animal-pollinated flowers than in animal-dispersed fruits. I tested this idea using size and colour traits for the flowers and fruits of 472 species occurring in three floras (St John, Hawaii and the Great Plains). Phylogenetically controlled analyses of nearest-neighbour distances in multidimensional trait space matched the predicted pattern: in each of the three floras, flowers were more divergent from one another than were fruits. In addition, the spacing of species clusters differed for flowers versus fruits in the flora of St John, with clusters in flower space more divergent than those in fruit space. The results are consistent with the idea that a major driver of angiosperm diversification has been stronger selection for divergent floral morphology than for divergent fruit morphology, although genetic, physiological and ecological constraints may also play a role.     

Early seed fall and seedling emergence: precursors to tropical restoration

We explore processes of seed immigration and seedling recruitment before an experimental rainforest restoration matures enough to affect either. Twenty-four 30 × 30-m plots were fenced in 12 ha of pasture in 2006. Seeds were collected in ninety-six 1-m⁻² seed traps; recruits were censused in ~12,000 m² in which establishment was allowed. We tested effects of distance from forest, living trees, and stumps of trees cut during site preparation on seed rain in 2007 and effects of these and soil depth on recruits through June 2008. Seed fall and recruitment were not correlated with distance to forest 90–400 m away, nor to living shade trees outside the 160 × 485-m experimental grid. Recruitment differed for animal- and wind-dispersed species in a topographically complex landscape. Recruitment of wind-dispersed species was random with respect to soil depth or distance to recent stumps. Recruitment of animal-dispersed species was multimodal; partial correlations with number of stumps within 30 m of plots were significant with soil depth held constant (P < 0.025), as were correlations of recruitment with soil depth with number of stumps held constant (P < 0.01). Animal-dispersed recruits were often not conspecifics of adults that had been cut, indicating a legacy of attraction by fruiting trees of animals bearing seeds from distant sources. Ecological implications are that recruitment in pastures released from grazing reflects a mix of widely scattered wind-dispersed pioneers and, where animal-dispersed trees exist, multi-modal and decidedly non-random recruitment of pioneer and later successional animal-dispersed trees from seed banks.     

An extinct genus of Salicaceae based on twigs with attached flowers, fruits, and foliage from the Eocene Green River Formation of Utah and Colorado, USA

A newly recovered twig with attached leaves and flowers from the Eocene Green River Formation of Utah provides the basis for recognizing a new, extinct genus of Salicaceae sensu lato (s.l.). Pseudosalix handleyi gen. et sp. nov. has alternate lanceolate leaves with pinnate, semicraspedodromous venation and a serrate margin with glandular teeth. The inflorescence is terminal on the twig and is unisexual, composed of flowers organized in a paniculoid cyme, with lateral paraclades of pedicellate flowers. The attached pistillate flowers have four prominent sepals that are valvate in bud, spreading but basally fused at anthesis; the single pistil of each flower is ovoid with three or four longitudinal sutures, indicating development to a capsular fruit. Three or four recurved styles radiate from the apex of the pistil, each with a distal globose stigma. The infructescence, verified by attachment to twigs with the same kind of leaves, bore capsular fruits of three and four valves. Associated but unattached, staminate flowers also have four well-developed, basally connate sepals. They are pedicellate and bear several stamens, each with a short filament and globose anther. The available morphological characters place the fossil species within the Salicaceae s.l. as an immediate sister to the clade containing Populus and Salix. Although the likely outgroup genera (including Itoa, Poliothyrsis, Carrierea, and Idesia) to tribe Saliceae all occur in Asia today and not North America, the occurrence of both Pseudosalix and Populus in the Eocene of Utah raises the possibility of a North American origin for the Saliceae.     

Complete cpDNA genome sequence of Smilax china and phylogenetic placement of Liliales--influences of gene partitions and taxon sampling

The complete nucleotide sequence of the chloroplast genome (cpDNA) of Smilax china L. (Smilacaceae) is reported. It is the first complete cp genome sequence in Liliales. Genomic analyses were conducted to examine the rate and pattern of cpDNA genome evolution in Smilax relative to other major lineages of monocots. The cpDNA genomic sequences were combined with those available for Lilium to evaluate the phylogenetic position of Liliales and to investigate the influence of taxon sampling, gene sampling, gene function, natural selection, and substitution rate on phylogenetic inference in monocots. Phylogenetic analyses using sequence data of gene groups partitioned according to gene function, selection force, and total substitution rate demonstrated evident impacts of these factors on phylogenetic inference of monocots and the placement of Liliales, suggesting potential evolutionary convergence or adaptation of some cpDNA genes in monocots. Our study also demonstrated that reduced taxon sampling reduced the bootstrap support for the placement of Liliales in the cpDNA phylogenomic analysis. Analyses of sequences of 77 protein genes with some missing data and sequences of 81 genes (all protein genes plus the rRNA genes) support a sister relationship of Liliales to the commelinids-Asparagales clade, consistent with the APG III system. Analyses of 63 cpDNA protein genes for 32 taxa with few missing data, however, support a sister relationship of Liliales (represented by Smilax and Lilium) to Dioscoreales-Pandanales. Topology tests indicated that these two alignments do not significantly differ given any of these three cpDNA genomic sequence data sets. Furthermore, we found no saturation effect of the data, suggesting that the cpDNA genomic sequence data used in the study are appropriate for monocot phylogenetic study and long-branch attraction is unlikely to be the cause to explain the result of two well-supported, conflict placements of Liliales. Further analyses using sufficient nuclear data remain necessary to evaluate these two phylogenetic hypotheses regarding the position of Liliales and to address the causes of signal conflict among genes and partitions.