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.     

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.     

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.     

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.     

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.     

Natural hybridization between an outcrossing and a selfingPhlox (Polemoniaceae): The maternal species of F1 hybrids

Natural F₁ hybrids between the outcrossingPhlox drummondii and the predominantly selfingP. cuspidata were examined to ascertain the proportion of hybrid individuals mothered by each species. Species-specific restriction fragment patterns (both nrDNA and cpDNA) were established as markers, and synthetic hybrids of known parentage were utilized to determine that the chloroplast genome is maternally inherited. Of 89 mature natural hybrids examined, approximately two thirds were mothered byP. drummondii, the outcrosser. That the outcrosser should mother most hybrids is expected since it is dependent upon incoming pollen for fertilization, and hybrids may result when heterospecific pollen is received. The fact that the highly selfingP. cuspidata mothered nearly one third of the hybrids is surprising, and may be related to both pre-zygotic and post-zygotic factors. Which species mothers hybrids has important implications for the potential for introgression as well as its direction.     

RESPONSE TO SELECTION ON AUTOGAMY IN PHLOX

Two cycles of artificial selection were performed to increase autogamous fruiting in two wild populations of the self-incompatible Phlox drummondii, to decrease autogamous fruiting in two wild populations of the self-compatible Phlox cuspidata, and to both increase and decrease autogamous fruiting in a cultivar of P. drummondii which is pseudo-self-compatible. The breeding systems were determined to be genetically quite flexible, independent of inbreeding depression and other genetic phenomena which could hinder a breeding system shift. This is especially true for increasing autogamy. Self-pollen-pistil compatibility seems to be the single character affected by selection. Based on the continuous variation in both autogamy and self-compatibility, we suggest that the change has been due to genes which modify the self-incompatibility reaction rather than to the simple segregation of alleles at the S-locus.     

SEGREGATION AND RECOMBINATION OF CHEMICAL CONSTITUENTS IN A HYBRID SWARM OF BAPTISIA LAEVICAULIS × B. VIRIDIS AND THEIR TAXONOMIC IMPLICATIONS

Turner , B. L. (U. Texas, Austin), and Ralph Alston . Segregation and recombination of chemical constituents in a hybrid swarm of Baptisia laevicaulis × B. viridis and their taxonomic implications. Amer. Jour. Bot. 46(9): 678–686. lllus. 1959.—Selected plants from a hybrid population of Baptisia laevicaulis × B. viridis and individuals from “pure” populations of the parental species were examined chromatographically. The resulting patterns were then compared with the respective phenotypes by the use of hybrid indices. Although the parental type individuals yielded nearly uniform patterns, hybrid-type plants showed a striking recombination of chemical components and there was an excellent correlation between chromatographic patterns and hybrid expression. Of particular interest was the detection of a new and distinctive compound in 2 of the hybrid-type plants which was not observed in either parent. Chromatograms of 3 other Baptisia species also showed distinctive patterns, and their relationship to each other, as indicated by morphological features, was accompanied by similarities in the biochemical pattern. The segregation and recombination of biochemical constituents are discussed with respect to their evolutionary and taxonomic implications. It is suggested that an extension of the techniques utilized might provide for the establishment of “biochemical profiles” which should prove of considerable value to the systematist.     

A comparative study of the North AmericanPyrola asarifolia and its Asian vicariad,P. incarnata (Ericaceae)

Samples of North AmericanPyrola asarifolia and its Asian vicariad,P. incarnata, were compared in morphology and leaf flavonol constituents. No qualitatively distinct attributes separate the plants from these two regions. In addition, a broad overlap exists in the ranges of the 11 morphological characteristics examined. Both Asian and North American plants exhibit some regional differentiation, but no simple clinal pattern is evident. Sepal width is the only morphological characteristic that exhibits a bimodal tendency in sample means. Asian plants have significantly narrower sepals compared with the North American samples. The chromatographic pattern of leaf flavonols in the Asian plants is distinct from the North American because of the reduced number of compounds present and in having one large, highly concentrated quercetin glycoside as the major constituent. The Asian vicariad is recognized asP. asarifolia Michaux subsp.incarnata (DC.) Haber & Takahashi, comb. nov. It is also considered to have a close affinity with the EurasianP. rotundifolia.     

INTERSPECIFIC HYBRIDIZATION AND TAXON UNIFORMITY IN ARABIS (CRUCIFERAE)

Intermixed populations of Arabis holboellii Hornem. var. pendulocarpa (A. Nels.) Roll., A. williamsii Roll., and their putative hybrids; and of A. holboellii var. retrofracta (Grah.) Rydb., A. drummondii Gray and A. divaricarpa A. Nels., the latter also considered to be a probable hybrid taxon, were studied in the field in Montana and Wyoming. In spite of the “intermediacy” of the putative hybrids in both cases, hybrid swarms were not found. In each instance, the “intermediate” as well as the taxa regarded as parental were remarkably uniform. The existence of three relatively uniform morphotypes in each of the several populations is contrary to the usual situation involving hybridizing taxa. A possible explanation is that seed production, at least in the “intermediate” morphotype individuals, involves apomixis, thus circumventing the usual sexual process that would result in a hybrid swarm. Amphiploidy as a possible mechanism for producing A. divaricarpa as a stable hybrid was ruled out in one population where the chromosome number 2n = 14 was found in all three taxa present in the population. Arabis divaricarpa, widely distributed in northern North America, often occurs as a uniform morphotype where no other species of Arabis grows. If it is of hybrid origin, as the evidence suggests, a mechanism other than sexual reproduction must be present to keep up uniformity and at the same time perpetuate the taxon. It is suggested that facultative apomixis is involved in the reproductive process and that it controls the genetic variability that would otherwise be present by insuring maternal inheritance in each successive seed crop.     

Dioecy derived from distyly and pollination in Psychotria rubra (Rubiaceae) occurring in the Ryukyu Islands,Japan

Floral morphology and breeding system were examined in natural populations of Psychotria rubra (Rubiaceae) occurring in the Ryukyu Islands in southwest Japan. This species is morphologically dimorphic with long‐ and short‐styled morphs; however, these morphs are functionally dioecious. All long‐styled morphs set fruits, but their short anthers completely lack pollen grains; thus, they function as pistillate flowers. Alternately, short‐styled morphs usually have fertile pollen and never set fruit, whether after open or obligate pollinations; thus, they function as staminate flowers. However, some short‐styled morphs in Mounts Katsuu‐dake and Oppa‐dake populations have no fertile pollens. No fruit was obtained from bagging experiments, indicating that P. rubra requires pollinators for its reproduction. In our pollinator observations, flies and short‐tongued wasps were the main visitors of the flowers of P. rubra. Thus, these insects are probably effective pollinators for P. rubra in the Ryukyu Islands. Considering the morphological features, the dioecy found in P. rubra is probably derived from distyly; however, no sufficient evidence is available showing that dioecy has evolved “directly” from distyly.     

MORPHOLOGICAL VARIABILITY IN GENETICALLY DEFINED CATEGORIES OF ANURAN HYBRIDS

Hybridization phenomena in anurans have traditionally been studied through morphological comparisons, under the assumption that various hybrids (e.g., F₁‘s, backcrosses) are predictably intermediate to parental species. We critically evaluate this assumption by examining morphology in genetically categorized hybrids between the treefrogs Hyla cinerea and H. gratiosa. A total of 202 frogs from a hybridizing population in Alabama were assayed for allozyme and mitochondrial DNA genotype and for a large suite of osteological characters. Discriminant analyses demonstrated distinct morphological separation between the genetically “pure” parental species. Morphometric analyses of genetically identified hybrids showed: 1) an extreme range of phenotypic expression within F₁ and backcross classes, and 2) no apparent directional parental bias on the F₁ phenotype. Had morphology alone been used as a guide, over 40 percent of the individuals with known hybrid ancestry would have been misclassified as “pure” parental species, and about 25 percent of the backcross individuals would not have been distinguished from F₁‘s. These results exemplify the utility of joint comparisons of morphology and genotypic constitution in studies of natural hybridization, and they emphasize the limitations inherent in describing hybrid classes solely by morphological criteria.     

MORPHOLOGICAL VARIATION WITHIN AND BETWEEN NATURAL POPULATIONS OF NON-TIDE POOL FUCUS DISTICHUS (PHAEOPHYTA) IN NEW ENGLAND1,2

The sources of morphological variation in natural populations of Fucus distichus L. ssp. edentatus (De la Pyiaie) Powell and F. distichus L. ssp, evanescens (C. Agardh) Powell from New England were evaluated. Individuals from different populations were morphologically distinct as judged by population means and analysis of variance. A correlation between the plant's form and its habitat was established by field observation. The broadest material grows in calm estuarine habitats, while the narrowest plants occur in exposed, open coastal areas. Major differences in morphology also appear seasonally. In addition, microhabitat factors such as exposure to wave action and elevation explain some morphological variation. Cultured germlings from distinct populations of “evanescens” and “edentatus” type plants were outplanted to an experimental garden in order to ascertain whether the variation was heritable or environmentally induced. The latter experiments indicate that major differences in morphology are genetically determined.     

Morphological clines across a karyotypic zone of house mice in Central Italy

Multivariate morphometric differentiation between chromosomal races of the mouse Mus domesticus in Central Italy was investigated using a population of 2n = 22 “CB” karyotype, three populations of standard 2n = 40 karyotype, five populations of 2n = 22 “CD” karyotype and three populations from the hybrid zone between the latter two karyotypes. Whilst populations of different karyotype generally have significantly different morphometry, canonical analysis does not reveal that the populations ordinate into distinct aggregations based on karyotype, largely because the 2n = 22 “CD” populations are so diverse. Nevertheless, canonical analysis does reveal a significant cline in morphology across the contact zone between the 2n = 40 and 2n = 22 “CD” mice. The nature of this transition, i. e. a cline 1. within the 2n = 40 range, 2. within the hybrid range (but unrelated to chromosome number) and 3. within the 2n = 22 “CD” range, tends to indicate that the morphometric divergence is due to adaptation to the different ecological regimes across which these mice are distributed rather than the phylogenetic divergence of the karyotypic races.     

THE GEOGRAPHICAL DISTRIBUTION OF B-CHROMOSOMES OF XANTHISMA TEXANUM DC. (ASTERACEAE). I. SURVEY OF THE RANGE

The distribution of B-chromosomes throughout the range of Xanthisma texanum has been determined from approximately 750 individuals from 80 wild populations in Texas, Oklahoma, and New Mexico. Counts of meiotic chromosome figures of n = 4_II + 1 B_I and n = 5_II (4_II + IB_II) are reported for the first time for subsp. texanum var. texanum and for subsp. texanum var. orientalis. The wide occurrence of B-chromosomes in natural populations of subsp. drummondii is also documented. B-chromosomes are present in 22 (27.2%) randomly distributed populations whose characteristics are in no way distinct from populations without B-chromosomes. In populations with B-chromosomes present, 19.8% of the plants contained one or two B-chromosomes, but in total only 5.8 % of all plants contained one or two B-chromosomes. All populations in one isolated population group of subsp. drummondii lack B-chromosomes.     

Bryophyte County Floras

Abstract

Experimental study of Pohlia populations that regularly produce axillary bulbils in Britain has shown that there are seven species present. The distinguishing bulbil characters of field populations of each species are retained when the plants are cultivated under uniform conditions, and can be used as a valid taxonomic criterion. Variation in bulbil morphology within and between populations of P. proligera is due to a combination of genetic, environmental and developmental variation. Bulbils show considerable phenotypic plasticity in P. drummondii but in the other five species studied variation is largely or entirely developmen tal.     

EXPERIMENTAL HYBRIDIZATION OF XANTHIUM STRUMARIUM (COMPOSITAE) FROM ASIA AND AMERICA. II. SESQUITERPENE LACTONES OF F1 HYBRIDS

Effects of hybridization on sesquiterpene lactones in Xanthium strumarium were studied for clues to the relationship of Old and New World populations. In crosses between indigenous Asiatic plants in the “strumarium” morphological complex and various American complexes that produce xanthinin as a major sesquiterpene lactone, the F₁ hybrids contained xanthinin and the related compounds, xanthanol, xanthatin, and xanthinosin. In other crosses with various American complexes that produce xanthumin, the stereoisomer of xanthinin, the F₁ hybrids contained xanthinin and xanthumin as well as xanthinin-related compounds and their stereoisomers, xanthumanol, deacetoxylxanthumin, and tomentosin. The Asiatic plants of “strumarium” from Hong Kong involved in the crosses produce approximately equal percentages of xanthinin, xanthatin and xanthinosin, but those from India contain only xanthinosin. The putative introduction of the American morphological complex, “chinense,” contains xanthumin as the major component but shows chemical diversity that indicates genes derived from the indigenous “strumarium” complex.     

Field bioassays show heterospecific mating preference asymmetry between hybridizing North American Papilio butterfly species (Lepidoptera: Papilionidae)

Few studies of interspecific mating preferences of naturally hybridizing species have been done in the field. Yet this is the only way potentially critical habitat-specific factors can be included in mating behavior evaluations. We conducted mate selection (male preference) studies using tethered pairs of heterospecific, size-matched virgin yellow females of Papilio glaucus and P. canadensis with free-flying male P. glaucus populations in south-central Florida, and with free-flying male P. canadensis populations in northern Michigan. Florida males clearly preferred the conspecific P. glaucus females of the pairs, as indicated by the touches/attempted matings and by the actual copulations observed. In 1997, 66.7% of the total attempts (n=168) and 94.2% of all copulations (n=69) were with the conspecific. In 1998 the same pattern of conspecific male preference was observed, with 67.3% of the touches/attempts (n=49) and 100% of the copulations (n=30) being with the P. glaucus females. In contrast, the natural populations of P. canadensis males did not show the expected conspecific preference. In fact, with wild P. canadensis males in 1997 a very strong preference was observed for the heterospecific P. glaucus females of the pairs, accounting for 75.8% of all touches/attempted matings (n=483) and 81.7% of all copulations (n=476). The strong asymmetry in interspecific mating preferences among these hybridizing Papilio species may reflect an ancestral trait in P. glaucus females that elicits strong preference in P. canadensis males, due perhaps to previously possessed male sensory bias or "good genes" indicators in the females. However, reinforcement of reproductive isolation in or near the hybrid zone of species overlap (sympatry) cannot be ruled out. Electronic Publication