EXPERIMENTAL HYBRIDIZATION,CHROMOSOMAL DIVERSITY,AND PHYLOGENY WITHIN GAURA (ONAGRACEAE)

Patterns of interfertility for intersectional hybridizations in Gaura are generally consistent with current taxonomic hypotheses regarding the sectional division of the genus. Crossability is generally lower for intersectional crosses than for intrasectional, interspecific crosses. Average intersectional cross-success ranged from 0–14% among the eight sections, and a total of 54 intersectional hybrids were produced from 1,872 intersectional pollinations. Low rates of capsule-set and no intersectional hybrids were produced by G. mutabilis (section Gauridium), an early offshoot of a primitive ancestor of Gaura. Two species of section Xerogaura that are regarded as relicts of the primitive ancestor of all other Gaura showed low averages for cross-success, but did produce hybrids in combination with three relatively advanced sections (Campogaura, Stipogaura, Pterogaura). Low capsule-set and no intersectional hybrids were produced by G. parviflora (section Schizocarya). Extreme morphological specialization for autogamous reproduction has been accompanied by reproductive isolation of G. parviflora. Only three hybrids resulted from crosses using G. coccinea (section Campogaura), a polyploid complex. Each successful cross paired G. coccinea with a putative progenitor, either a species of section Xerogaura or G. villosa of section Stipogaura. Hybrids of G. coccinea with odd ploidy number showed reduced pollen staining. Species of Stipogaura produced only three hybrids, one each in combination with species of sections Xerogaura, Campogaura, and Pterogaura. Gaura drummondii (section Xenogaura) registered the highest average cross-success despite being a tetraploid. All 25 hybrids of G. drummondii resulted from crosses with G. suffulta, and were triploids that showed reduced pollen staining. Intersectional crosses fail to support the hypothesized relationships of G. drummondii either to G. coccinea or to species of section Stipogaura. Section Gaura produced moderate averages of cross-success, but all 20 hybrids resulted from crosses with G. suffulta. Fifty of the 54 intersectional hybrids were descended from a species of section Pterogaura. Crosses that used G. suffulta as one parent produced 49 of these hybrids. Four hybrids were from crosses with G. macrocarpa (section Xerogaura), and the remaining 45 were due to the recurrent success of crosses that paired G. suffulta as maternal parent with either G. drummondii or a species of section Gaura.     

QUANTITATIVE VARIATION IN PHLOX: COMPARISON OF SELFING AND OUTCROSSING SPECIES

Variation of 20 quantitative characters was examined within and among 10 populations of the predominantly outcrossing Phlox drummondii and 4 populations of the predominantly selfing P. cuspidata grown in a greenhouse. Multivariate analysis of variance, considering all characters simultaneously, indicated that there were significant differences among populations in both species while analysis of individual characters demonstrated that there were significant population differences for 19 characters in P. drummondii and 13 characters in P. cuspidata. On average, 16% of the total phenotypic variation in P. drummondii occurred among populations compared to less than 4% of the total variation in P. cuspidata. In addition, P. drummondii exhibited significant differences among families within populations more frequently than P. cuspidata. Most observed variation in both species occurred within families where environmental and genetic sources of variation could not be partitioned. There was a trend for P. drummondii to have higher heritabilities than P. cuspidata for most characters even when assumptions about breeding systems were relaxed. Thus, the outbreeding species exhibited larger genetic differences among populations and among families within populations than the selfing species in the greenhouse environment. These data suggest that P. drummondii has the greater evolutionary potential of the two species and are consistent with the hypothesis that differences in population structure result from differences in the breeding systems of the two species.     

HYBRIDIZATION BETWEEN ANNUAL SPECIES OF PHLOX: POPULATION STRUCTURE

Phlox cuspidata (n = 7) and P. drummondii subsp. drummondii (n = 7) are closely related annuals which are indigenous to eastern and central Texas. The species typically occupy different ecological niches but may form contiguous or confluent populations in disturbed habitats and hybridize therein. On the basis of correlative interpretations of exomorphic, chromatographic and fertility information, hybridizing populations can be segregated into three distinct classes: (1) highly fertile plants with the morphological and phenolic attributes of P. drummondii; (2) highly fertile plants with the morphological and chemical attributes of P. cuspidata; (3) sterile plants with manifestly intermediate morphology and complementary chromatographic patterns. These data strongly suggest that hybridizing populations of P. drummondii and P. cuspidata are tritypic, being composed of “pure” or essentially “pure” parental species and a group of plants which has all of the attributes characteristic of an F₁ hybrid.     

A PENNSYLVANIAN CALAMITEAN CONE WITH ELATERITES TRIFERENS SPORES

In 1943 L. R. Wilson described some scattered elater-bearing spores found in a Middle Pennsylvanian coal ball from What Cheer, Iowa, as Elaterites triferens. The spores, averaging 60 μ in diameter, with a trilete scar on the proximal surface and three conspicuous elaters attached to their distal surface, have now been found in a fragment of a cone from the same locality. The cone is similar to Calamostachys in having whorls of sporangiophores, each with four adaxial sporangia, and some secondary xylem at the nodes, but it possibly differs in lacking sterile bracts.     

Two new species of the liverwort genus Riccia L. from tropical Africa: R. somaliensis and R. erubescens

Abstract

Two new Riccia species, R. somaliensis and R. erubescens from Africa are described and illustrated. They are both classified in the subgenus Riccia, section Riccia. R. somaliensis has hyaline, somewhat frilly scales and apolar, wingless spores; R. erubescens has shiny dark, purple-red to bright cherry-red scales and polar, winged spores with numerous small areolae; its tissues are sometimes distorted by gelatinous globules that are gradually extruded on the dorsal surface.     

SPORE MORPHOLOGY IN THE GENUS BRUCHIA SCHWAEGR. (MUSCI)

Scanning electron microscopy observations of the spores of Bruchia have resulted in the recognition of four spore types based on the ornamentation of the distal spore surface: warty or verrucate, pitted, reticulate, and spinose. The proximal surface of the spores of all species, except B. brevipes, is characterized by a central aperture region surrounded by a triangular murus or rows of spinae forming a triangle and 1 or 2 smooth or verrucate collars. The ornamentation patterns observed are considered to be characteristic for the genus. Spore morphology alone can rarely be used to distinguish species but in conjunction with certain other characteristics, it is an important taxonomic feature. Spore morphology is a major characteristic used to define the limits of the highly variable species B. flexuosa (spinose spores) and to distinguish it from the closely related species B. texana (reticulate spores). Variations in ornamentation patterns within the genus support the recognition of two subgenera (subgenus Bruchia and subgenus Sporledera). Spore morphology also supports the close relationship between Bruchia and Trematodon and is sufficient to eliminate several questionable taxa from the genus.     

Pleospora drummondii spec. nov. und ihr Anamorph Stemphylium drummondii spec. nov.

Pleospora drammondii spec. nov. and its anamorph Stemphylium drammondii spec. nov. The causal organism of a leaf spot disease on Phlox drummondii is described as Pleospora drummondii spec. nov. and its anamorph Stempbylium drummondii spec, nov. — in contrast to earlier reports from the USA, in which the fungus was identified as S. botryosum.     

Scanning electron microscopy observation of host entry by two brown algae endophytic in Laminaria saccharina (Laminariales, Phaeophyceae)

The germination of spores of the endophytic brown algae Laminariocolax aecidioides and Laminarionema elsbetiae (Ectocarpales sensu lato) on their host Laminaria saccharina was studied using scanning electron microscopy. Zoids were the infective units in both taxa. Despite a large difference in size between spores of the species studied, the initial steps of infection were similar. Zoids settled on the host surface, and during secretion of adhesive material from their anterior end, the spores became elongated to rod-shaped and stood erect on the host. A single germ tube developed at the proximal end and penetrated the intact surface of the host. Sharp edges around the entrance hole and the absence of inward deformation of the host surface around the settled zoids suggest an enzymatic rather than a mechanical penetration mechanism.     

Simultaneous evolution of competitiveness and defense: induced switching in Arabis drummondii

Optimality theory for plant defense against herbivores predicts an evolutionary tradeoff between the abilities to compete and defend. We tested this hypothesis by studying the effects of genetic variation in competitiveness on defense expression. Two closely related and differentially competitive congeners were compared for levels of resistance, tolerance, and secondary metabolite production. In a growth room experiment, plants of Arabis drummondii and A. holboellii were grown in the presence and absence of the common bunch grass Boutelloua gracilis, the specialist herbivore Plutella xylostella, and generalist herbivore Trichoplusia ni. Tolerance to competition, measured as growth next to the grass relative to controls in the absence of grass, was greatest for A. drummondii, the species that occurred in communities with higher densities of inter-specific neighbors. Measures of defense (resistance to herbivores, tolerance to damage, and concentrations of glucosinolates) varied inconsistently between the Arabis, species, depending on type of herbivore, competition level, and type of defense. The better competitor A. drummondii was more resistant to specialist herbivores, as in the field, and exhibited greater herbivore- and competition-induced changes in glucosinolate profiles. Further, when plants of A. drummondii were fed upon in competitive environments, the induced glucosinolate response was reduced while tolerance levels increased in an apparent switching of induced strategies. We suggest that competitiveness and defense responses are sometimes positively correlated because some defensive traits also function as competitive traits. A competitive function for defenses may also explain why defenses were affected by competition. Alternatively, since the induced response did not increase estimates of total glucosinolate content significantly, minimal defense costs might also allow the simultaneous evolution of competitiveness and defense. Finally, when faced with both herbivory and competition, some competitive species, such as A. drummondii, may switch to growth-based rather than toxin-based strategies as recent theoretical models predict.     

REPRODUCTIVE CHARACTER DISPLACEMENT IN PHLOX

Plants of the red- and pink-flowered P. drummondii were introduced into a natural population of P. cuspidata. Their seeds were progeny-tested, and the percentage hybrid seed determined for each variant. Thirty-eight percent of the seed from the pink variant were hybrid compared to 13% of the red. The difference in hybrid production is best explained by corolla color. The self- and cross-compatibility of nine sympatric and nine allopatric populations were studied in the greenhouse. On the average, sympatric populations are more self-compatible than allopatric populations. Pollen germination is 13.7% vs. 7.3%, the difference being statistically significant. The self-compatibility differential was accompanied by a self seedset differential. On the average, sympatric populations are more cross-compatible with P. cuspidata than are allopatric populations. The germination of P. cuspidata pollen on sympatric P. drummondii stigmas was 13.3% vs. 9.8% on allopatric stigmas. However, the difference is not statistically significant. The presence of P. cuspidata has promoted reproductive character divergence in P. drummondii. The shift in corolla color and the increase in self-compatibility reduce the potential for gametic wastage and interspecific hybridization.     

Consequences of shrub expansion in mesic grassland: Resource alterations and graminoid responses

Abstract. In the mesic grasslands of the central United States, the shrub Cornus drummondii has undergone widespread expansion in the absence of recurrent fire. We quantified alterations in light, water and N caused by C. drummondii expansion in tall‐grass prairie and assessed the hypothesis that these alterations are consistent with models of resource enrichment by woody plants. Responses in graminoid species, particularly the dominant C₄ grass Andropogon gerardii, were concurrently evaluated. We also removed established shrub islands to quantify their legacy effect on resource availability and assess the capability of this grassland to recover in sites formerly dominated by woody plants. The primary effect of shrub expansion on resource availability was an 87% reduction in light available to the herbaceous understorey. This reduced C uptake and N use efficiency in A. gerardii and lowered graminoid cover and ANPP at the grass‐shrub ecotone relative to undisturbed grassland. Shrub removal created a pulse in light and N availability, eliciting high C gain in A. gerardii in the first year after removal. By year two, light and N availability within shrub removal areas returned to levels typical of grassland, as had graminoid cover and ANPP were similar to those in open grassland. Recovery within central areas of shrub removal sites lagged behind that at the former grass‐shrub ecotone. These results indicate that the apparent alternative stable state of C. drummondii dominance in tall‐grass prairie is biotically maintained and driven by reductions in light, rather than resource enrichment. Within areas of shrub removal, the legacy effect of C. drummondii dominance is manifest primarily through the loss of rhizomes of the dominant grasses, rather than any long‐term changes in resource availability. C. drummondii removal facilitates grassland recovery, but the effort required to initiate this transition is a significant cost of woody plant expansion in mesic grasslands. Prevention of woody plant expansion in remnant tall‐grass prairies is, therefore, a preferred management option.     

INBREEDING DEPRESSION IN PARTIALLY SELF-FERTILIZING PHLOX

Inbreeding depression was measured in terms of the relative survivorship of self versus cross seed in 14 Phlox drummondii cultivars and 10 populations of P. cuspidata. The cultivars are predominantly outcrossing; P. cuspidata is predominantly self-fertilizing. The relative survivorship of self versus cross seed averages 0.84 in the cultivars and 0.99 in P. cuspidata. In wild P. drummondii, the average is 0.83. The mean number of lethal equivalents per zygote averages 0.69 in the cultivars and 0.05 in P. cuspidata, versus 0.79 in the wild P. drummondii. There is a significant negative correlation between the amount of automatic self-seed produced by a cultivar and the number of lethal equivalents it carries (r = ?0.60). Phlox cuspidata has a lower level of cross-seed abortion than P. drummondii, which probably is not due to a difference in the incidence of lethal genes in their populations.     

蜈蚣草孢壁发育及其系统学意义

利用光镜、扫描电镜和透射电镜对凤尾蕨科(Pteridaceae)蜈蚣草(Pteris vittata L.)孢壁的形成和发育进行研究。结果表明:蜈蚣草孢子四面体型,极面观钝三角圆形,赤道面观半圆形或超半圆形,近极面具瘤状纹饰和近极脊,远极面具脊并连成网状,具赤道环;孢子具乌毛蕨型外壁,由外壁外层构成纹饰的轮廓;实心型周壁由2层构成,且内层薄、外层具小球体。结合孢子外壁和周壁的发育特征,认为凤尾蕨科与裸子蕨科和水蕨科的亲缘关系较近,支持将裸子蕨科和水蕨科置于凤尾蕨科。     

Pollen competition and environmental effects on hybridization dynamics between Phlox drummondii and Phlox cuspidata

Pollen competition between species strongly influences hybridization dynamics in plants. By performing single- and mixed-donor pollinations, we show that soil Ca alters the outcome of interspecific pollen competition in the annual Phlox hybrid system of Phlox cuspidata and P. drummondii. In the absence of interspecific pollen competition, heterospecific pollen siring success of both species was influenced most strongly by the maternal growth environment, such that hybridization was facilitated when heterospecific pollen was deposited on stigmas of maternal plants growing in high Ca soils. When heterospecific pollen was forced to compete against conspecific pollen, however, the maternal growth environment did not influence hybridization, but the environmental origin of heterospecific pollen did, and this effect depended on the maternal species. Pollen of P. drummondii was more effective at outcompeting P. cuspidata pollen and preventing hybridization in P. drummondii dams when P. cuspidata pollen was derived from low Ca. Pollen competition within pistils of P. cuspidata was unaffected by pollen Ca environment. In situations in which P. cuspidata grows in lower soil Ca than P. drummondii, as has been documented in one population, these results suggest that the competitive ability of heterospecific pollen will be diminished by environmental effects of soil Ca. Thus, the environment in which pollen develops can influence interspecific pollen competition and hybridization frequency.     

Mating system and the hybridization between self-compatible Phlox cuspidata and self-incompatible Phlox drummondii

Mating system can impact the frequency of hybridization and therefore the maintenance of species diversity. I evaluate the effects of weak self-incompatibility (SI) in Phlox cuspidata and SI in Phlox drummondii on mating success within species and on hybridization dynamics between species under controlled conditions. The effects of SI on hybridization frequency were assessed by manipulating the relatedness of conspecific pollen and the relative timing of pollen deposition in mixed-donor interspecific pollinations. Selfing as opposed to outcrossing increased hybridization by 16% in P. cuspidata maternal plants and by 48% in P. drummondii maternal plants because self pollen did not compete as well against heterospecific pollen. The relative timing of conspecific versus heterospecific pollen deposition also impacted hybridization. In self-compatible P. cuspidata, the deposition of self pollen 5 h earlier than heterospecific pollen decreased hybridization by 28%. In self-incompatible P. drummondii, a 5 h delay in the deposition of compatible conspecific pollen increased hybridization by 32%. In this hybrid system, early self-pollination can decrease hybridization (but increase inbreeding) by P. cuspidata maternal plants, and SI may increase hybridization by P. drummondii maternal plants.     

A COMPARATIVE STUDY OF CELL DIVISION PATTERNS DURING GERMINATION OF SPORES OF ANEMIA,LYGODIUM AND MOHRIA (SCHIZAEACEAE)

Cell division patterns during germination of spores of Anemia (A. hirsuta, A. munchii, A. phyllitidis), Lygodium (L. circinatum, L. flexuosum, L. japonicum, L. salicifolium) and Mohria caffrorum have been examined by light microscopy of glycol methacrylate embedded materials. Spores of all species in a genus exhibited a constant pattern of division under different conditions of germination. In spores of species of Anemia, following an asymmetrical division, the proximal cell differentiated into the protonemal cell while the distal cell divided to produce the rhizoid. A similar division sequence was found in spores of Mohria caffrorum, but the fate of cells formed was reversed. In Lygodium spores, a proximal cell formed by an initial division of the spore cut off a protonemal cell, a rhizoid and a wedge-shaped cell by walls parallel to the polar axis. Our results contradict earlier observations on cell division sequence during germination of spores of these genera based on whole mount preparations.     

Paramagnetism in Bacillus spores: Opportunities for novel biotechnological applications

Spores of Bacillus megaterium, Bacillus cereus, and Bacillus subtilis were found to exhibit intrinsic paramagnetic properties as a result of the accumulation of manganese ions. All three Bacillus species displayed strong yet distinctive magnetic properties arising from differences in manganese quantity and valency. Manganese ions were found to accumulate both within the spore core as well as being associated with the surface of the spore. Bacillus megaterium spores accumulated up to 1 wt.% manganese (II) within, with a further 0.6 wt.% adsorbed onto the surface. At room temperature, Bacillus spores possess average magnetic susceptibilities in the range of 10⁻⁶ to 10⁻⁵. Three spore‐related biotechnological applications—magnetic sensing, magnetic separation and metal ion adsorption—were assessed subsequently, with the latter two considered as having the most potential for development.

     

DEVELOPMENT AND DISTRIBUTION OF MUCILAGE CANALS IN LYCOPODIUM

Two distinct types of mucilage canals are found in Lycopodium. One type, the veinal canal, is found in both sporophylls and in vegetative leaves, and is always in close proximity to the leaf trace. The other, the basal canal, is restricted to the strobilus where it forms a complex and extensive mucilaginous cylinder in the outer cortex and extends into the base of the sporophylls. Protruding secretory cells are formed in both types during a lysigenous developmental process. The occurrence of these two types of canals correlates with the three subgenera. Urostachys which lacks canals, Lepidotis which possesses both veinal and basal canals, and Lycopodium which possesses only basal canals.