Dormancy and dispersal in dimorphic achenes of tansy ragwort,Senecio jacobaea L. (Compositae)

Summary Marginal and central florets of the capitula of tansy ragwort Senecio jacobaea yield different kinds of fruit. The central (disk) achenes are lighter (x±SE=199±5g), more numerous (x±SE=58±0.6 achenes per head), and are equipped with a pappus aiding wind transport and rows of trichomes aiding animal transport. The marginal (ray) achenes are heavier (x±SE=286±7g), less numerous (virtually invariant at 13 achenes per head), and lack dispersal structures. Whereas disk achenes are relased shortly after they mature, ray achenes are retained by the parent for a period of months following maturity.Germination at constant temperature (20°C) and with alternating light (12 h light: 12 h dark) demonstrated that disk and ray achenes exhibit different germination syndromes. Germination percentage increases linearly with achene fresh weight in both types; for a given weight, disk achenes have a higher germination percentage than ray achenes. Germination time decreases with increasing achene weight in disk achenes, but increases with achene weight in ray achenes.The germination percentages and germination times for disk and ray achenes diverge progressively with increasing achene weight. The divergence in behavior is a result of diverging patterns of dry matter allocation in the two achene types. Increase in the size of disk achenes favors the embryo fraction, thereby speeding germination while reducing protection. Increase in the size of the ray achenes favors the pericarp fraction, thereby increasing protection while delaying germination.Reduced germination percentage and germination speed of the ray achenes were shown by experimental manipulation to be caused by physical inhibition by their thicker pericarps.Dimorphism in ragwort likely speards germination out in space and time, thereby increasing the number of safe sites an individual parent can exploit in disseminating offspring. The syndrome in other heteromorphic composites resembles that of ragwort, generally combining reduced dispersal-delayed germination in the outer achenes and distance dispersal-quick germination in the central achenes. The outer achenes are generally less numerous and larger. Dispersal traits (large numbers, early release and light wieght) are the direct opposite of dormancy traits (small numbers, delayed release and heavier weight). Thus conflicts between the properties determining dormancy and dispersal appear to require separate dormancy and dispersal phenotypes.     

Comparative capitular morphology and anatomy of Coreopsis L. and bidens L. (compositae), including a review of generic boundaries

The capitular and floral morphology and anatomy ofBidens L. andCoreopsis L. were studied. All the North American species ofCoreopsis were studied. Selected species ofBidens from North and South America andCoreopsis from South America were included. The results were compared with previous observations on African species ofBidens (incl.Coreopsis). Emphasis was given to character states of the ray florets, paleae, stylearm apices, outer phyllaries, achenes, and pollen grains. Some of the character states are unique features ofCoreopsis, e.g., globular and elongately conical receptacles, deltoid outer phyllaries, truncate and indistinctly 3–5-dentate, 3–4-lobed ray florets, narrowly spathulate paleae, subulate paleae with linear-filiform upper half, hairy and apically 3-cleft paleae, truncate, convex or shallowly conical stylearm apices with the sweeping hairs limited to the area above the stigmatic surfaces and the orbicular to circular achenes. The cylindric setaceous pappus bristles so commonly encountered inBidens are unknown inCoreopsis. The pappus bristles inCoreopsis are paleaceous but similar, though thicker ones are also found in African species ofBidens (incl.Coreopsis) with winged achenes. Twin-celled hairs (setulae) with differing degrees of wall thickness are found on the achenes ofCoreopsis sect.Pseudoagarista (Mexico and South America),Coreopsis sect.Pugiopappus (California), AfricanBidens with winged achenes (e.g.,B. prestinaria, B. macroptera) and some North AmericanBidens (e.g.,B. aristosa). Similar sclerotic parenchyma make up the achenial wings of species in both genera. These may be interpreted as homologous structures, indicating the underlying similarity of these taxa and their derivation from a common ancestral stock.     

Variability of the inflorescence ofMicroseris laciniata (Compositae: Cichorieae)

Quantitative characters of the flowering head of a garden population ofMicroseris laciniata were scored during the second, third, and fourth season of growth. Number of achenes per head, number of phyllaries per head and the average number of pappus parts per achene in single heads show significant plant to plant variation. Achenes per head and pappus parts per achene were scored in identical plants in two subsequent seasons. The number of pappus parts per achene varies freely between five and ten. This contrasts with annual species ofMicroseris in which either five or ten pappus parts are found, depending on the species. In spite of a clear plant-specific average of pappus parts, both high and low pappus part determination can be demonstrated in all specimens. The number of pappus parts depends on the position of an achene on the receptacle, marginal achenes usually having fewer pappus parts than central ones. This gradient is not closely correlated with the position of an achene on the genetic spiral.     

Lipochaeta and Melanthera (Asteraceae: Heliantheae subtribe Ecliptinae): establishing their natural limits and a synopsis

We restrict the genusLipochaeta to the allopolyploid species of the typical section.Lipochaeta s.str. is interpreted to be the result of an intergeneric hybridization betweenMelanthera and a presently unknown taxon, perhaps of the genusWedelia. Lipochaeta is characterized, in addition to its allopolyploidy (n=26), by having both flavonols and flavones, disk corollas with 4 lobes, achenes tuberculate at maturity, the disk achenes flattened to slightly biconvex, and ray achenes obcompressed.Lipochaeta sect.Aphanopappus andWollastonia are here reduced to synonymy underMelanthera. We transfer 14 HawaiianLipochaeta and one New Caledonian species as well as the AsianWedelia prostrata toMelanthera. These transfers, along with the species in Africa and North America, bring the number of species in the genus to 35.Melanthera is delimited by an abruptly narrowed to truncate and flattened top of the achene, (0−)1–15(−20) often unequal, ciliate or barbellate, caducous pappus bristles immediately surrounding the corolla, involucral bracts and receptacular paleae with many veins forming longitudinal striations, and n=15. The florets are 5-merous, the corollas are yellow or white, and rays are absent (in white-flowered species) or present and neutral or fertile. In dealing with species formerly placed inLipochaeta, the GalapagosL. laricifolia is here transferred from the illegitimate generic nameMacraea toTrigonopterum and the BrazilianL. goyazensis is transferred toAngelphytum. We maintain the earlier reduction ofEchinocephalum underMelanthera and reduce all three taxa originally described in it to one,M. latifolia.     

DIFFERENTIAL GERMINATION OF RAY AND DISC ACHENES IN HEMIZONIA INCRESCENS (ASTERACEAE)

The differential germination responses of ray and disc achenes of Hemizonia increscens (Asteraceae) were compared in field and laboratory investigations in order to gain insight into the ecological and evolutionary significance of heterocarpy. In the field, 200 ray and 200 disc achenes were placed in native, sterilized soil in a series of cleared, randomized, replicated plots. In a nearby plot a similar number of achenes were placed in plastic petri dishes in which high moisture conditions were maintained. Disc achene germination occurred under relatively minimal moisture conditions (<1 cm rainfall for 19 days) and relatively mild temperature regimes (21–7 C). Disc achene germination began three days after planting in the field plot and four days after they were put into the field petri dishes. In contrast, the onset of ray achene germination occurred 21 days after planting in the field plot and 19 days after planting in the field petri dishes. Averages of 2.05 and 2.71 disc achenes/day germinated in the field plot and field petri dishes, respectively. These contrasted with averages of 0.57 and 0.50 ray achenes/day germinated in the field plot and in the field petri dishes, respectively. A total of 67.5% and 69% disc achenes germinated in the field plot and the field petri dishes, but only 18% and 16.5% ray achenes germinated in the field plot and field petri dishes, respectively. Three separate treatments, using 100 ray and 100 disc achenes in each, were performed in laboratory growth chambers: 1) nicking the fruit coat, 2) excising the embryo, and 3) leaving the fruit coat intact. Onset of germination for all disc achene treatments occurred after three days. No significant differences were found among the three disc achene treatments in timing, rate, or germination percentage. All three disc treatments in the laboratory closely paralleled those for disc achenes in the field plots in time and germination percentage, but rates of germination were not as high. Germination of the nicked and excised ray achenes treatments began after four days, while germination of the untreated ray achenes began after 27 days. Untreated ray achenes in the laboratory paralleled the ray achenes in both field experiments in rate and germination percentage, but were delayed in time of germination. The nicked and excised ray achene treatments, however, were similar to the disc achene treatments in time of germination, and were not significantly different from disc achenes in rate of germination. These data suggest 1) that ray and disc achenes are markedly different in germination under identical conditions in field and laboratory experiments, and 2) this difference in germination response may be due to the thicker pericarp of the ray achenes.     

POPULATION DYNAMIC CONSEQUENCES AND EVOLUTION OF SEED TRAITS OF HETEROSPERMA PINNATUM (ASTERACEAE)

We summarize research on variation in achene morphology of Heterosperma pinnatum. Each flowering head has a broad range of achene morphologies. There is no between-year seed bank. There are no ecologically relevant differences in achene size, seedling size, or seedling growth rates of the different morphs. Achenes located centrally in the heads lose dormancy earlier during the period between autumn achene production and the onset (May-July) of the rainy season. This results in relatively more early emergence of central achenes. Awned central achenes also have greater adhesive dispersibility. Emergence, survival, and fecundity were measured for achenes sown into natural populations in central Mexico. When harsh conditions result in few seeds surviving to reproduce, the advantage tends to go to the more “conservative” peripheral achenes. Greenhouse experiments show a strong genetic component to the determination of achene proportions within and among populations and a variety of genetic correlations. Populations with a high proportion of awned achenes tend to be found in vegetation types where they are likely to be ephemeral. A high proportion of central achenes tend to be found in sites with little pre-rainy season precipitation. These patterns are interpreted in terms of selection for dispersal and the within-year timing of germination.     

Significance of achene characteristics and within-achene resource allocation in the germination strategy of tetraploid Aster pilosus var. pilosus (Asteraceae)

The interrelationships among achene weight, allocation to embryo and pericarp, and germination time were determined for 500 stratified achenes of tetraploid Aster pilosus Willd. var. pilosus. Only 52.6% of the achenes germinated. Germinated achenes were significantly heavier than ungerminated achenes. Germination time was independent of achene weight and embryo weight, but varied inversely with pericarp weight. Variable achene weight is evolutionarily advantageous. Heavy achenes are at an advantage in that their proportionately larger embryos and thinner pericarps facilitate germination, promoting competitive establishment of seedlings. Lighter achenes are also at an advantage through increased dispersibility, and their relatively thick pericarp provides a persistent seed bank. Evolutionary pressures presumably maintain the variability in achene weight of var. pilosus. These results are discussed in the context of the early midsuccessional ecology of var. pilosus.     

Aster chusanensis (Asteraceae), a new species from Korea

Asterchusanensis is described and illustrated, and compared with its most closely related species,A. pseudoglehni andA. spathulifolius. This new species differs markedly from the latter two by its unequal inner and outer phyllaries, pubescent stem, leaves, and corolla tube, and ray florets partly in two series.     

Morphological variability of Tragopogon pterodes Pančić ex Petrović achenes: taxonomic evaluation of heterocarpy

This study presents the carpological heterogeneity of Tragopogon pterodes, an endemic plant species from Balkan Peninsula. The goal of this paper was to determine the variability of morphological characters of achenes and assess the degree of their morphological differentiation relative to the position they occupy in the receptacle. Analysis included four different populations of T. pterodes and examination of 18 (8 qualitative and 10 quantitative) characters of achenes in total. Data analysis employed basic statistic, univariate (ANOVA, Tukey HSD) and multivariate (CDA, CA) analyses to check the level of heterocarpy in this species. The results indicate possible morphological differentiation of three groups of achenes (outer, median and inner achenes), depending of their position on the receptacle. The members of all three types were described, illustrated and compared. The variability of investigated carpological characters in this species is providing important information prior to their further taxonomic treatment.     

一年生短命植物疏齿千里光果实异形性的生态学意义

疏齿千里光(Senecio subdentatus)是分布在新疆北部古尔班通古特沙漠中的一种具异形瘦果的菊科一年生短命植物。将野外观测与室内实验相结合, 对该物种异形瘦果的形态、扩散和萌发特性, 以及异形瘦果产量与植株大小的关系进行了研究, 并对其生态学意义进行了探讨。结果表明: 疏齿千里光果序中的外围果和中央果均为柱形, 但前者为淡黄色, 后者为褐色, 且二者在大小、冠毛长度及果皮微形态等方面均存在明显差异。两种瘦果均以单个果实为扩散单元, 且在静止空气中的降落速度和在1 m·s^-1与2 m·s^-1风速下的扩散距离无显著差异, 说明虽然果实大小和冠毛长度对瘦果扩散具有不同的影响, 但对其整体扩散能力无明显影响。在各温变周期(5/2、15/2、20/10、25/15和30/15 ℃)处理中, 淡黄色外围果的萌发率均高于褐色中央果的, 且不同温度间两种瘦果的萌发率均存在显著差异, 但光照条件对其无显著影响。果序中的中央瘦果数明显多于外围果的, 且植株中外围果所占比例与植株大小间呈显著负相关关系, 而中央果所占比例与植株大小间呈显著正相关关系。这些特点说明, 该物种的小植株倾向于产生较多较易萌发的外围果, 大植株倾向于产生较多不易萌发的中央果。在古尔班通古特沙漠不可预测的极端环境中, 疏齿千里光可通过异形瘦果间的萌发差异及调节其不同大小植株中异形瘦果的比例, 来减少同胞后代之间的竞争, 增加其对不同微环境条件的生态适应性。     

RESURRECTION OF THE GENUS ENDOLEPIS AND CLARIFICATION OF ATRIPLEX PHYLLOSTEGIA (CHENOPODIACEAE)

The genus Endolepis was first described by Torrey in 1860 with E. suckleyi (E. dioica [Nutt.] Standley) as the only species, noting that it differed from species of Atriplex by the presence of perianth parts in the female flowers. Later, Standley described two additional species: E. covillei and E. monilifera. H. M. Hall and Clements merged Endolepis with Atriplex because they thought that the presence of perianth parts in female flowers was variable in Atriplex phyllostegia. It is now clear that this observation was erroneous; A. phyllostegia never has perianth parts in its female flowers. The plants that they examined that had female flowers with perianth parts were specimens of Endolepis covillei; only those with female flowers devoid of perianth parts were specimens of Atriplex phyllostegia. These two taxa differ by several other major attributes including differences in leaf shape, leaf anatomy, fruiting-bract size and shape, fruiting-bract appendages, and flowering habit, and therefore justify taxonomic separation. Also, because the presence of perianth parts in bracteolate female flowers is a rare attribute in the tribe Atripliceae, consistently absent in Atriplex, but always present in Endolepis, the retention of the genus Endolepis, separate from Atriplex, is deemed warranted. We propose that the genus Endolepis comprise two species, E. dioica and E. covillei. The species named E. monilifera by Standley is based on a specimen of Atriplex serenana Nels.     

Systematic overview of Krigia (Asteraceae-Lactuceae)

Cytological and morphological variation among Krigia species is examined. Krigia exhibits a broad range of chromosome numbers including n = 4, 5, 6, 9, 10, 15, and 30. Section Krigia is characterized by reflexed phyllaries and a base chromosome number of x = 5, while section Cymbia is characterized by erect phyllaries and chromosome numbers of n = 4, 6, and 9. The micromorphological characteristics of achenes, pappus, styles, corolla, pollen, stomata, and trichomes are documented using scanning electron microscopy. Among these, the pappus shows the greatest diversity and three major types are identified: 1) a pappus of many bristles and scales, as in K. dandelion, K. montana, and K. biflora; 2) a pappus of five bristles and five scales, as in K. virginica and K. occidentalis; and 3) an absent or highly reduced pappus, as in the K. cespitosa complex and K. wrightii. Thirty-five cytological and morphological characters are subjected to phylogenetic analyses. The two sections, Krigia and Cymbia, form monophyletic lineages. Within section Krigia, the annual species, K. virginica, forms an independent clade, while the perennial species, K. dandelion, K. biflora, and K. montana, form a monophyletic clade. Krigia montana and K. biflora are identified as sister species and a hybrid between these has been identified. The hybrid is more similar morphologically to K. montana than K. biflora. Within section Cymbia, phylogenetic relationships among K. wrightii, K. occidentalis, and K. cespitosa are uncertain. Nine taxa of Krigia are herein recognized: K. dandelion, K. biflora, K. biflora var. viridis (comb. nov.), K. montana, K. virginica, K. wrightii (comb. nov.), K. occidentalis, K. cespitosa, and K. cespitosa f. gracilis (comb. nov.). Phylogenetic relationships among 12 taxa of Krigia species are compared using various combinations of morphology, chloroplast DNA, and nuclear ribosomal DNA data. Tree topologies from different combinations of data are largely congruent. The most resolved phylogenetic tree is obtained using the combined data from morphology, chloroplast DNA, and nuclear ribosomal DNA.     

Bidirectional Secretions from Glandular Trichomes of Pyrethrum Enable Immunization of Seedlings

Glandular trichomes are currently known only to store mono- and sesquiterpene compounds in the subcuticular cavity just above the apical cells of trichomes or emit them into the headspace. We demonstrate that basipetal secretions can also occur, by addressing the organization of the biosynthesis and storage of pyrethrins in pyrethrum (Tanacetum cinerariifolium) flowers. Pyrethrum produces a diverse array of pyrethrins and sesquiterpene lactones for plant defense. The highest concentrations accumulate in the flower achenes, which are densely covered by glandular trichomes. The trichomes of mature achenes contain sesquiterpene lactones and other secondary metabolites, but no pyrethrins. However, during achene maturation, the key pyrethrin biosynthetic pathway enzyme chrysanthemyl diphosphate synthase is expressed only in glandular trichomes. We show evidence that chrysanthemic acid is translocated from trichomes to pericarp, where it is esterified into pyrethrins that accumulate in intercellular spaces. During seed maturation, pyrethrins are then absorbed by the embryo, and during seed germination, the embryo-stored pyrethrins are recruited by seedling tissues, which, for lack of trichomes, cannot produce pyrethrins themselves. The findings demonstrate that plant glandular trichomes can selectively secrete in a basipetal direction monoterpenoids, which can reach distant tissues, participate in chemical conversions, and immunize seedlings against insects and fungi.     

Germination,reproduction and interference in the amphicarpic annual Emex spinosa (L.) Campd

Summary A comparison was made of the dimorphic subterranean and aerial achenes of the annual Emex spinosa (L.) Campd. Subterranean achenes were less dormant, had a higher percentage of viability, germinated at a faster rate, were less temperature-dependent in germination and produced larger seedlings than aerial achenes. Subterranean achenes germinated, equally well in light or dark but aerial achenes had a strong light requirement. There was a linear relationship between numbers of weight of aerial achenes and plant size in the field. More resources were allocated to reproduction in plants restricted in size because of environmental conditions. Subterranean achene production was less affected than aerial by density, nitrogen levels, varying field conditions and interference between plants grown, from subterranean and aerial achenes. In mixtures of such plants, those from subterranean achenes had larger leaf area and stems and more aerial achenes. These differences were not apparent in monocultures by the time of harvest. Seed dimorphism is more likely in short-lived fugitive species and the different roles associated with dimorphism (persistence in situ versus dispersal in E. spinosa) are seen as an ecological advantage in such species.     

The Relationship Between Achene Weight,Embryo Weight and Germination inTaraxacumApomicts

Apomicts provide the opportunity to investigate the variation in achene size and performance that is due to factors other than genetic differences between achenes. Achene characteristics of a single capitulum from each of 15 biotypes of apomicticTaraxacumwere investigated. Within a capitulum, achene fresh weight ranged from 4.45-fold to 13.74-fold. Overall, 59.8% of achenes germinated. The probability of any one achene germinating was influenced both by its source and by its weight and ranged from 0.049 to 0.902 between capitula and from 0.028 to 0.873 between weight classes. An investigation of the relationship between achene parts in three biotypes identified a linear trend for embryos to acquire a greater proportion of total achene weight as achene weight increases. Although the heaviest achenes showed the best germination, it is predicted that, for a given investment of maternal resources, seedling number would be maximized by producing achenes of a single intermediate weight.     

Inbreeding effects on fitness traits in the heterocarpic herb Leontodon autumnalis L. (Asteraceae)

Heterocarpic plants are characterized by the production of distinct types of fruits that usually differ in their ecological behavior. In the Asteraceae, differences are mainly found between peripheral non-dispersal and central dispersal achenes (single-seeded fruits). Inbreeding depression is considered as an evolutionary force as it may reduce several fitness traits, and in the case of heterocarpic plants, it could influence fitness traits (e.g., seed set, germination rate, growth rate) of each fruit morph, which may have important ecological and evolutionary consequences. In particular, differential effects on fitness traits and dispersal of selfed and outcrossed progeny can strongly determine the viability of extant populations and the potential to colonize new habitats. We conducted a hand-pollination experiment in greenhouse conditions to test whether inbreeding affects the fitness of achene morphs in the heterocarpic herb Leontodon autumnalis (Asteraceae). Results show that achene morphs significantly differ in their ecological behavior, peripheral achenes germinating more and faster than central achenes. The significant interaction between pollination treatment and achene morph for germination probability might indicate a link between dormancy and mating system in L. autumnalis: germination was higher for outcrossed achenes in central achenes whereas the opposite pattern was exhibited by peripheral achenes. Selfing dramatically reduced seed set, probably as a consequence of strong self-incompatibility mechanisms rather than inbreeding effects. Inbreeding depression significantly affected late life-cycle traits, such as growth rate and biomass at flowering. Overall, results suggest that inbreeding depression seems to be an important selective force maintaining outcrossing in L. autumnalis.     

Additions To The Flora Of Shetland

ABSTRACT

Background: Pallenis spinosa (Asteraceae) produces both winged and wingless achenes. Both achene morphs are non-dormant and show a similar embryo size, rendering dispersal ability as their only apparent functional difference.

Aims: We studied morph-specific release and spatial dispersal patterns to ascertain whether the common view of seed dimorphism as a mixed strategy, that is functionally fully differentiated morphs, is appropriate for this system.

Methods: For three years, at the onset of achene release, in early autumn, we placed achene traps at different distances from source plants, censusing achene arrival at 3–4 day intervals. We constructed morph-specific dispersal kernels and related release intensity to prevailing meteorological conditions in census intervals. Selected kernel models were used to describe dispersal effects of observed changes in the proportion of winged achenes (p_w) in successive released fractions.

Results: Achene release extended up to early-mid winter, peaking in rainy, windy intervals. Throughout the season, p_w decreased progressively. Unexpectedly, the wingless morph produced the longest dispersal tails and it only showed ability for fat-tailed dispersal. Consequently, maximum dispersal distances steadily increased throughout the season.

Conclusions: Achene dimorphism in P. spinosa appears to allow a within-season continuous reshaping of the seed-dispersal kernel instead of representing a mixed strategy.     

Morphometric Analysis of Sunflower (<Emphasis Type="Italic">Helianthus annuus</Emphasis> L.) Achenes from Mexico and Eastern North America<Superscript>1</Superscript>

Morphometric Analysis of Sunflower ( Helianthus annuus L.) Achenes from Mexico and Eastern North America. Sunflower (Helianthus annuus L.) has played a major role in the evolution of agricultural systems in the Americas. The discovery of ancient domesticated remains from archaeological deposits in pre-Columbian Mexico offers new dimensions to widely accepted viewpoints on the domestication pattern of H. annuus. Although American sunflower populations north of Mexico have been examined extensively, Mexican indigenous domesticated landraces have not been studied in any detail. In this study, we morphologically assessed wild and domesticated sunflower achenes from Mexico and compared them to similar datasets from eastern North America. Additionally, we evaluated the utility of four computer-assisted shape measurements in discriminating between wild and domesticated sunflower achenes (fruits) and compared variation in achene size among modern wild and cultivated populations from both Mexico and the U.S. We found that, of the shape parameters tested, none were informative in distinguishing wild achenes from domesticated varieties. Subsequent size analysis, using conventional parameters of length, width, and thickness, showed that modern wild populations from Mexico had smaller achenes compared to modern populations from eastern North America. Domesticated achenes unearthed from Mexican archaeological sites, however, were significantly larger than the early domesticated specimens recovered from eastern North America. Our methodological results indicate that variation in archaeological sunflower achenes is better described by conventional size parameters rather than computerized shape analysis.     

Eleocharis khandwaensis sp. nov. (Cyperaceae) from the Madhya Pradesh,India

A new species, Eleocharis khandwaensis Mujaffar, Chandore & S. R. Yadav is described from the Bamangaon village of Khandwa district in Madhya Pradesh state, India. This species belongs to Eleocharis subgen. Limnochloa (P. Beauv. ex Lestib.) Torr. and it is similar to E. lankana T. Koyama but differs in having stolons, weak bristles spinules of perianth, achenes with annulate apex with a short neck, truncate achene base, dark brown to black coloured achenes and nut surface epidermal cells that are isodiametrically hexagonal to pentagonal. Eleocharis khandwaensis grows in ponds along road sides in the Bamangaon and Tirandaz villages.     

长柄马兰（菊科）的形态补充描述及细胞学研究

结合模式产地采集的长柄马兰（Aster longipetiolatus C.C.Chang）标本以及查阅相关文献资料,发现已有的形态描述不充分且缺乏细胞学方面的报道。本研究对长柄马兰的形态特征进行补充描述和核型分析,为探讨其分类地位提供资料。结果包括：①发现并补充长柄马兰新的形态特征：花序托凸起、总苞片3~4层、舌状花具有退化雄蕊、舌片具柔毛、瘦果5肋。②细胞学结果表明,长柄马兰的染色体数目为2n=18,核型公式为2n= 18=16m+2sm,核型属于1A。