ECOTYPIC DIFFERENTIATION WITHIN FOUR NORTH AMERICAN PRAIRIE GRASSES. II. BEHAVIORAL VARIATION WITHIN TRANSPLANTED COMMUNITY FRACTIONS

Ecotypic differentiation in transplanted clones of Andropogon scoparius Michx., A. gerardi Vitman, Panicum virgatum L. and Sorghastrum nutans (L.) Nash showed north-south correlations. In Austin, Texas, during 1959–1962, southern populations had earliest spring activity, latest flowering and latest dormancy. Northern community fractions from Massachusetts to North Dakota showed early flowering and a short span separating initial anthesis among the species-populations. Community fractions from Virginia to Nebraska had initial anthesis over a longer period than in more northern or more southern samples. Community fractions from South Carolina to eastern Texas had late flowering over a short period. Western community fractions, latitudinally and altitudinally diverse, were behaviorally uniform. The length of the growing period and its recurrent selection have sorted out ecological variants in harmony with the habitat gradients. Whether they are of the predominant physiognomic types, as in the true prairie region, or understory plants in the pine forests, the 4 grass taxa are in the climax matrix.     

Clonal identification by microsatellite loci in sporadic flowering of a dwarf bamboo species, <Emphasis Type="Italic">Sasa cernua</Emphasis>

Dwarf bamboo species are monocarpic. They flower simultaneously and die after several decades. The type of flowering in the genus Sasa ranges from sporadic to gregarious. In order to determine whether or not the sporadic flowering of dwarf bamboo is fixed genetically, we investigated the distribution of clones using eight microsatellite (SSR) loci in a sporadic flowering patch of Sasa cernua Makino, a major dwarf bamboo species found in central Hokkaido. In May 2006, flowering occurred on 60.5% of living culms in a 1600 m² patch. We established a 50 × 10 m study plot in this patch and noted 1267 clumps consisting of 2529 living culms. We investigated all 1267 clumps and identified six multilocus genotypes as clones using five variable SSR loci. All flowering clumps belonged to the same clone. On the other hand, non-flowering vegetative clumps were also discovered to be of the same clone. These data suggest that all flowering culms originated from a single clone of a sporadic flowering patch of S. cernua. Clonal analysis for investigation of sporadic flowering of S. cernua revealed that only a portion of a clone flowers and dies instead of the whole clone.     

A CHEMOTAXONOMIC STUDY OF LEMNACEAE

One hundred eighty-six clones of Lemnaceae, representing world-wide collections of 22 taxa, were grown in axenic cultures reproduced by clonal subcultures. Following morphological examinations under a range of culture conditions, a “key” clone was selected to represent each taxon. These key clones were completely representative of the qualitative flavonoid chemistry of the taxa as determined from a number of clones. With the exceptions of Spirodela polyrhiza and S. biperforala which produce identical flavonoids, the flavonoid associations of each species of Lemnaceae were unique in the family. Through paper chromatographic comparisons of purified compounds, plus visible and ultraviolet absorption spectroscopy of isolated individual flavonoids, a total of 47 different compounds was described. These substances included: glycosides of anthocyanidins (cyanidin and petunidin); glycosides of flavonols (kaempferol and quercetin); glycosides of flavones (apigenin and luteolin); and several glycoflavones (C-glycosyl-flavones). When grown under equivalent controlled conditions of culture, infraspecific variation in the qualitative production of these 47 flavonoids was detected in only one flavone glycoside of the species Lemna perpusilla. The reliability of the flavonoid patterns under various conditions of culture was investigated. Under 62 different regimes S. oligorhiza and S. polyrhiza showed only minor variations in their flavonoid glycosides. Studies of other taxa supported this generalization. Under controlled conditions, morphological intergradation obscured identification of many collections. Each could be conclusively identified by its flavonoid chemistry.     

NATURE OF THE PLANT COMMUNITY. VI. TEXAS GRASSLAND COMMUNITIES UNDER TRANSPLANTED CONDITIONS

Mc Millan , Calvin . (U. Texas, Austin.) Nature of the plant community. VI. Texas grassland communities under transplanted conditions. Amer. Jour. Bot. 48(9): 778–785. Illus. 1961.—Clones of 7 grass taxa, Bouteloua gracilis (H. B. K.) Lag., B. eriopoda (Torr.) Torr., B. curtipendula (Michx.) Torr., Panicum virgatum L., the Andropogon scoparius Michx. complex, the Andropogon gerardi Vitman-hallii Hack. complex, and Sorghastrum nutans (L.) Nash, were transplanted from throughout their distribution in Texas and studied in an experimental garden at Austin. Restricted to western Texas and Panhandle areas, Bouteloua gracilis and B. eriopoda contained similar early-flowering clones throughout their distribution. Less restricted to western sites, B. curtipendula contained later-flowering types from eastern and central areas. In the 4 remaining, widespread taxa, early-flowering potential characterized clones from western sites. These 4 widespread taxa contain the latest-flowering clones from the coast of southern Texas. Clones of Stipa leucotricha Trin. and Rupr. from a broad area in Texas lacked a flowering gradient. Grassland communities of western Texas and the Panhandle, attuned to short growing seasons and low rainfall, were composed of opportunists, the Bouteloua species, and early-flowering variants within the widespread species. Communities of central Texas in habitats of highly unpredictable moisture pattern and a relatively long growing season contained later-flowering variants. Coastal communities attuned to a long growing season contained the latest-flowering variants.     

BIOCHEMICAL HETEROPHYLLY AND FLAVONOID EVOLUTION IN NORTH AMERICAN POTAMOGETON (POTAMOGETONACEAE)

Morphologically heterophyllous species of Potamogeton also commonly display biochemical heterophylly with respect to flavonoid compounds. Generally, floating leaves contain an assortment of flavonoids, whereas submersed leaves often exhibit reduced flavonoid profiles. In strictly submersed (homophyllous) species, two patterns occur. Linear-leaved species have few flavonoids and their biochemical profiles resemble those of submersed leaves of heterophyllous species. Broad-leaved homophyllous species possess flavonoid profiles more similar to those of the floating leaves of heterophyllous species. Numerical analysis of these chemical data is consistent with phylogenetic relationships within the genus derived independently on the basis of morphological and chromosomal data. Glycoflavones, which are probably maintained in floating leaves because of their UV filtering ability, exhibit the most pronounced biochemical heterophylly in Potamogeton. The lack of glycoflavones in submersed leaves of heterophyllous species and in linear-leaved homophyllous species is attributable to the ability of naturally colored water to significantly absorb harmful UV radiation. These observations provide strong support for earlier hypotheses suggesting the importance of flavonoid evolution in the conquest of exposed terrestrial habitats by plants.     

Experimental studies of adaptation in Clarkia xantiana. III. Phenotypic selection across a subspecies border

Sister taxa with distinct phenotypes often occupy contrasting environments in parapatric ranges, yet we generally do not know whether trait divergence reflects spatially varying selection. We conducted a reciprocal transplant experiment to test whether selection favors “native phenotypes” in two subspecies of Clarkia xantiana (Onagraceae), an annual plant in California. For four quantitative traits that differ between subspecies, we estimated phenotypic selection in subspecies’ exclusive ranges and their contact zone in two consecutive years. We predicted that in the arid, pollinator‐scarce eastern region, selection favors phenotypes of the native subspecies parviflora: small leaves, slow leaf growth, early flowering, and diminutive flowers. In the wetter, pollinator‐rich, western range of subspecies xantiana, we expected selection for opposite phenotypes. We investigated pollinator contributions to selection by comparing naturally pollinated and pollen‐supplemented individuals. For reproductive traits and for subspecies xantiana, selection generally matched expectations. The contact zone sometimes showed distinctive selection, and in ssp. parviflora selection sometimes favored nonnative phenotypes. Pollinators influenced selection on flowering time but not on flower size. Little temporal variation in selection occurred, possibly because of plastic trait responses across years. Though there were exceptions and some causes of selection remain obscure, phenotypic differentiation between subspecies appears to reflect spatially variable selection.     

VARIATION IN THE DEGREE OF CLEISTOGAMY WITHIN AND AMONG SPECIES OF THE GRASS DANTHONIA

North American species of the grass Danthonia bear distinct chasmogamous and cleistogamous flowers. Four taxa occurring in North Carolina (D. compressa, D. epilis, D. sericea, and D. spicata) were examined for variation in the degree of cleistogamy. The flowering culms bear a single terminal panicle consisting of 4 to 12 spikelets of chasmogamous flowers. In each leaf axil, at the nodes of the flowering culm, is a single spikelet of cleistogamous flowers completely surrounded by the leaf sheath. The percentage of cleistogamous flowers produced on any culm depends on the number of chasmogamous and cleistogamous spikelets and the number of flowers in each type of spikelet. All four characters vary among the taxa examined. Danthonia compressa produces, on the average, 50% cleistogamous flowers; D. spicata, 25% cleistogamous flowers; D. sericea and D. epilis, 5% cleistogamous flowers. The degree of cleistogamy in D. spicata is associated with certain habitat features. Populations from mountain sites, disturbed sites, and non-woodland sites produced higher percentages of cleistogamous flowers than did populations from piedmont, undisturbed, and woodland sites, respectively. Grazing may favor increased cleistogamy because cleistogamous flowers are produced lower on the plant. In North Carolina, the most frequently grazed Danthonia taxa also produce the highest percentage of cleistogamous flowers.     

TILLERING AND MORTALITY OF THE SALT MARSH CORDGRASS,SPARTINA ALTERNIFLORA

Recruitment and mortality of Spartina alterniflora tillers and culms were estimated from measurements of live standing crops at Wallops Island, Virginia. Stands of tall and medium form grass produced from 452 to 1,227 tillers m⁻² yr⁻¹. Those tillered in spring lengthened into culms their first year, while those tillered in late summer and fall may have overwintered and lengthened into culms their second year. Biennial growth was most prevalent in tall form Spartina. Fifty-six to sixty-one percent of all tillers died before lengthening into culms. Of the remainder, 9 to 56% died as culms before the time of flowering. Sixty-three percent of the surviving culms of tall form Spartina flowered, compared to only 14 to 23% in medium form. Mortality of tillers and culms removed 16 to 35% of the annual production of Spartina biomass from live standing crops before the time of flowering.     

Carex yandangshanica sp. nov. (Cyperaceae; C. sect. Rhomboidales) from Zhejiang,China

Carex yandangshanica C. Z. Zheng & X. F. Jin, a new species of Carex sect. Rhomboidales Kükenth. from Zhejiang, China, is described and illustrated. The new species is similar to C. thibetica Franch. and C. brevicuspis C. B. Clarke, but differs from the former by having trabeculate leaves, pistillate spikes with 6–10 staminate flowers or without staminate flower at the apex, and achenes at apex contracted into a slightly curved beak, and from the latter by having lateral culms, trabeculate leaves, and elliptic, rubiginous and awned pistillate scales. SEM microphotographs of perigynia and achenes are provided for the new species and related taxa.     

SURVIVAL PATTERNS IN FOUR PRAIRIE GRASSES TRANSPLANTED TO CENTRAL TEXAS

Transplanted clones of four widespread prairie grasses, Andropogon scoparius, A. gerardii, Panicum virgatum, and Sorghastrum nutans, that had survived in cultivation 1958-1962 in central Texas were studied without cultivation 1963-67 to determine survival patterns. In all four species, clones from northern and eastern sites in the United States were eliminated. Survival of A. scoparius was restricted to plants originating in central and southern Texas and in northern Mexico. Surviving clones of A. gerardii, P. virgatum, and S. nutans were chiefly of Texas origin but included other clones mostly from the south central United States. Population samples of the four species from a central Texas grassland community showed greatest survival in a multi-ramet comparison of clones originating from North Dakota to Mexico City and in a multi-clone comparison from six sites in Texas and one in New Mexico. While the superior adaptation to the local habitat by the local populations might have been expected, this study documented the survival potential of organisms in the local ecosystem.     

Potential host colonization by insect herbivores in a warmer climate: a transplant experiment

We conducted a transplant experiment to investigate the potential colonization of a plant species by insect herbivores under a warmer climate. Acacia falcata seeds collected from four latitudes, encompassing the current coastal range of the species (1150 km), were grown in the same soil type and climatic conditions in a glasshouse. Plants were then transplanted to two sites, 280 km north of A. falcata's current coastal range; the transplant sites were 1.2 and 5.5°C warmer than the northernmost and southernmost boundaries of the species' current range, respectively. We compared the structure and composition of the herbivorous Hemiptera and Coleoptera communities on the transplants (i) to that of A. falcata within its current distribution, (ii) to a closely related Acacia species (Acacia leptostachya) that naturally occurred at the transplant sites, and (iii) among the A. falcata transplants originating from seeds collected at different latitudes. Herbivory on A. falcata was also compared between the transplants and the current distribution, and among transplant originating from different latitudes. Thirty species of externally feeding herbivorous Coleoptera and Hemiptera were collected from the transplanted A. falcata over a period of 12 months following transplantation. Guild structure of this herbivore community (based on the proportion of species within each of seven groups based on taxonomy and feeding style) did not significantly differ between the transplants and that found on A. falcata within its natural range, but did differ between the transplants and A. leptostachya. Rates of herbivory did not significantly differ between the transplants and plants at sites within the natural range. There were no significant differences in herbivore species richness or overall rates of herbivory on the transplants originating from different latitudes. In conclusion, host plant identity was apparently more important than climate in influencing the structure of the colonizing herbivore community. If this result holds for other plant–herbivore systems, we might expect that under a warmer climate, broad patterns in insect community structure and rates of herbivory may remain similar to that at present, even though species composition may change substantially.     

Flowering regimes of terrestrial orchids: unpredictability or regularity?

Abstract. Empirical data on many species of terrestrial orchids suggest that their flowering pattern over the years is extremely irregular and unpredictable. A long search for the reason has hitherto proved inconclusive. Irregular flowering was attributed to costs associated with sexual reproduction, to herbivory, or to the chaotic behaviour of the system represented by difference equations describing growth of the vegetative and reproductive organs. Data on the seasonal growth of leaves and inflorescence of Dactylorhiza majalis are used here to test alternative explanations of the irregular flowering patterns of orchids. These patterns are found to be extremely rare in our data set. Neither costs of reproduction nor grazing seem to explain the rare events of a transition from flowering one year to sterility or absence the next year. These transitions are almost exclusively characteristic of one of four experimental sites, the only unmown site, where the mean leaf area and incidence of flowering in the whole population is also in decline. It is therefore hypothesized that irregular flowering regimes may be characteristic of sites with temporarily or steadily declining populations and that they are usually not present in prosperous ones - at least for D. majalis.     

ECOTYPIC DIFFERENTIATION WITHIN FOUR NORTH AMERICAN PRAIRIE GRASSES. I. MORPHOLOGICAL VARIATION WITHIN TRANSPLANTED COMMUNITY FRACTIONS

Ecological variation within transplanted community fractions, containing Andropogon scoparius Michx., A. gerardi Vitman, Panicum virgatum L. and Sorghastrum nutans (L.) Nash, included broad behavioral and morphological differentiation. In Austin, Texas, during 1958–1962, southern community fractions that contained early spring activity, late flowering and late fall dormancy included taller plants than early-flowering, northern community fractions. The tallest populations in all 4 species originated in Texas. In P. virgatum, size of leaves and panicles showed north-south trends similar to culm height values. Leaf pubescence and leaf glaucousness in all 4 species showed geographic patterning that was not correlated with north-south maturity gradients. Greatest pubescence was on clones in community fractions from pine woods in the Southeast. Glaucousness was concentrated in community fractions from central Texas northward to Nebraska.     

Flavonoid chemistry ofPolygonum sect.Tovara (Polygonaceae): A systematic survey

Polygonum sect.Tovara includes three controversial species;P. virginianum, P. filiforme, andP. neofiliforme. The flavonoid chemistry of these was examined to provide additional information on their delimitation and levels of differentiation. Eight flavonoid compounds were isolated and identified, all of which were 3-O-glycosides of the flavonols kaempferol, quercetin, and myricetin, and their acylated derivatives. Although they exhibit relatively simple flavonoid profiles, the three taxa are readily distinguished by their flavonoid constituents. In addition, they show fundamental differences in flavonol types and glycosylation patterns. These results, in conjunction with evidence from the morphology, strongly suggest thatP. virginianum, P. filiforme, andP. neofiliforme are closely allied but distinct species.     

Spatial and temporal flowering patterns of the monocarpic dwarf bamboo Sasa veitchii var. hirsuta

In order to clarify spatial and temporal patterns in the flowering process of S. veitchii var. hirsuta, the authors investigated the flowering of the species at a regional level in a mountainous area of Kyoto City, and at population/individual culm levels at experimental forest stations within the same municipal region. S. veitchii var. hirsuta flowered in several separate areas spanning ranges between several and more than 10?km north?Csouth and east?Cwest from 2004 to 2007. Sporadic flowering occurred the year before mass flowering. In the mass-flowering year, 98% of the culms in an S. veitchii var. hirsuta population flowered, and the other 2% or so flowered in the years before and after mass flowering. These results show a spatiotemporal synchronous flowering pattern (with most culms of populations gregariously flowering in a year over a large area) and an asynchronous flowering pattern (with a small percentage of the culms of populations flowering before and after the mass-flowering year) for S. veitchii var. hirsuta. It was also clarified that the year and scale of mass flowering differ by area, and that mass flowering occurs in individual areas in successive years. The survey performed at the individual culm level revealed that the mean numbers of open florets and attached seeds per culm showed two peaks from April to July, suggesting that S. veitchii var. hirsuta flowering events occur twice a year.     

Marsileaceaephyllum, a new genus for marsileaceous macrofossils: leaf remains from the Early Cretaceous (Albian) of southern Gondwana

Three new taxa from Albian, Early Cretaceous assemblages in Gondwana (Australia and Antarctica) and two previously described fossils from the Late Cretaceous and Eocene of North America are attributable to the heterosporous semi-aquatic fern family Marsileaceae. They are assigned to Marsileaceaephyllum, a morphotaxon erected here for sterile remains (whole plants, and isolated leaves and leaflets) of Marsileaceae. The Gondwanan taxa, Marsileaceaephyllum lobatum and Marsileaceaephyllum spp. B-C, have either a cruciform leaflet arrangement or dichotomous and anastomosing venation characteristic of modern Marsileaceae. Two previously established taxa, Marsilea johnhallii and Marsilea sp., which represent sterile Marsileaceae, are also transferred to the new genus (now Marsileaceaephyllum johnhallii and Marsileaceaephyllum sp. A, respectively). Examination of all fossil venation patterns reveals four new venation types not present in extant taxa, suggesting that most fossil Marsileaceae (leaves) are distinct from extant genera, and are likely members of extinct lineages. This is further supported by the absence of modern megaspore types in the Early Cretaceous.     

Systematics ofJusticia sect.Pentaloba (Acanthaceae)

The transfer of the four taxa ofSiphonoglossa sect.Pentaloba toJusticia is proposed. It is shown that the taxa of this section were placed inSiphonoglossa primarily because of a single-character phenetic relationship and that they correctly belong inJusticia. In addition to morphology, data from cytology and flavonoid chemistry are also presented that support this intergeneric transfer. A key to the taxa and a detailed taxonomic treatment of the section are provided.     

A revision of Sullivantia (Saxifragaceae)

Sullivantia (Saxifragaceae) is restricted to the United States; its species occur in very specialized habitats such as wet cliffs and moist canyon walls in scattered localities from Washington and Oregon, eastward to Wyoming, Montana, Colorado, Missouri, Iowa, Illinois, Minnesota, Wisconsin, Indiana, Ohio, and Virginia. Relationships in the genus were reevaluated using data from morphology (including the use of greenhouse-grown plants), crossability, and flavonoid chemistry. The conclusion of this study is thatSullivantia comprises four morphologically similar allopatric taxa:S. oregana, S. hapemanii var.hapemanii, S. hapemannii var.purpusii, andS. sullivantii.     

STRUCTURE AND DYNAMICS OF BAMBOOS IN THE WOLONG NATURAL RESERVE,CHINA

Culm (ramet) demography of two bamboos, Fargesia robusta and Bashania fangiana, was studied over 7 yr in southwest China to determine if: 1) ramet numbers remain relatively constant; 2) ramet mortality was constant; 3) density response was greater in spreading vs. compact clones; 4) extrinsic factors (e.g., predation, climate) influenced ramet dynamics; and 5) ramets that die are smaller than ones that live. Culm numbers remained relatively constant (±20%) in most plots, but culm numbers increased (>;20%) on some sites. Mortality of culms >;6 mo old over 7 yr was low and constant, but mortality of younger culms (≤6 mo) was high and variable due to predation. Spreading clones had greater recruitment response to density than compact clones, but predation and climate may have diminished or amplified the observed demographic patterns. Predators and perhaps climate influenced culm demography, and culms that died were smaller than remaining live culms. The demography of the two bamboos we studied is compared to other clonal plants, including bamboos, to identify common processes controlling ramet demography.