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.     

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.     

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.     

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.     

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.     

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.     

Do annual and perennial populations of an insect-pollinated plant species differ in mating system?

Background and AimsTheory predicts that outcrossing should be more prevalent among perennials than annuals, a pattern confirmed by comparative evidence from diverse angiosperm families. However, intraspecific comparisons between annual and perennial populations are few because such variation is uncommon among flowering plants. Here, we test the hypothesis that perennial populations outcross more than annual populations by investigating Incarvillea sinensis, a wide-ranging insect-pollinated herb native to China. The occurrence of both allopatric and sympatric populations allows us to examine the stability of mating system differences between life histories under varying ecological conditions.MethodsWe estimated outcrossing rates and biparental inbreeding in 16 allopatric and five sympatric populations in which both life histories coexisted using 20 microsatellite loci. In each population we measured height, branch number, corolla size, tube length and herkogamy for ~30 individuals. In a sympatric population, we recorded daily flower number, pollinator visitation and the fruit and seed set of annual and perennial plants.Key ResultsAs predicted, outcrossing rates (t) were considerably higher in perennial (mean = 0.76) than annual (mean = 0.09) populations. This difference in mating system was also maintained at sympatric sites where plants grew intermixed. In both allopatric and sympatric populations the degree of herkogamy was consistently larger in outcrossing than selfing plants. Perennials were more branched, with more and larger flowers than in annuals. In a sympatric population, annuals had a significantly higher fruit and seed set than perennials.ConclusionsGenetically based differences in herkogamy between annuals and perennials appear to play a key role in governing outcrossing rates in populations, regardless of variation in local ecological conditions. The maintenance of mating system and life history trait differentiation between perennial and annual populations of I. sinensis probably results from correlated evolution in response to local environmental conditions.     

DYNAMICS OF A HYBRID ZONE IN PHLOX: AN EXPERIMENTAL DEMOGRAPHIC INVESTIGATION

The dynamics of a narrow hybrid zone between Phlox drummondii ssp. drummondii and ssp. mcallisteri was investigated using an experimental demographic approach. Three populations of each subspecies as well as those from the hybrid zone were involved. Experimental plots containing seeds from each of the nine populations were established in each of the eight sites. Germination, survivorship to reproduction, and fecundity were monitored at 4–6 wk intervals. Finite rates of increase were estimated from these data. There were significant site effects for each demographic variable and rate of increase as well as significant population effects for germination and finite rate of increase. Although there were population effects, no significant differences were found between plant types in sites of the subspecies or the hybrid zone. We reject the hypothesis that the narrow width of the hybrid zone is the result of hybrid disadvantage. We propose that the narrow width is a result of restricted gene flow and discuss the supporting evidence.     

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.     

Breeding systems and hybridization inPetrorhagia sect.Kohlrauschia (Caryophyllaceae)

In the diploid speciesP. prolifera, the evolution of autogamy has resulted in the presence of distinct selfing and outcrossing races. The change in breeding system toward autogamy is associated with a reduction in the size of floral features and a lower pollen production. The outbreeding populations have only been found in Greece, while the selfing populations have colonized much of Europe. The two races appear to be reproductively isolated from one another. —Petrorhagia sect.Kohlrauschia contains four species.P. glumacea is outbreeding and shows unilateral incompatibility with the small-flowered race ofP. prolifera and interfertility with the large-flowered race; althoughP. glumacea is sympatric with the large-flowered race, they do not appear to hybridize in the wild. Amongst the allopatric species, internal breeding barriers are also found. These may take the form of hybrid sterility or seed incompatibility.P. velutina is autogamous and reproductively isolated from all the other species of the section regardless of whether they are sympatric or allopatric. It is suggested that the evolution of autogamy has been an important factor in the spread ofP. velutina, the tetraploidP. nanteuilii, and the small-flowered race ofP. prolifera in Europe.     

Shifts to earlier selfing in sympatry may reduce costs of pollinator sharing

Coexisting plant congeners often experience strong competition for resources. Competition for pollinators can result in direct fitness costs via reduced seed set or indirect costs via heterospecific pollen transfer (HPT), causing subsequent gamete loss and unfit hybrid offspring production. Autonomous selfing may alleviate these costs, but to preempt HPT, selfing should occur early, before opportunities for HPT occur (i.e., “preemptive selfing hypothesis”). We evaluated conditions for this hypothesis in Collinsia sister species, C. linearis and C. rattanii. In field studies, we found virtually identical flowering times and pollinator sharing between congeners in sympatric populations. Compared to allopatric populations, sympatric C. linearis populations enjoyed higher pollinator visitation rates, whereas visitation to C. rattanii did not differ in sympatry. Importantly, the risk of HPT to each species in sympatry was strongly asymmetrical; interspecies visits comprised 40% of all flower‐to‐flower visits involving C. rattanii compared to just 4% involving C. linearis. Additionally, our greenhouse experiment demonstrated a strong cost of hybridization when C. rattanii was the pollen donor. Together, these results suggest that C. rattanii pays the greatest cost of pollinator sharing. Matching predictions of the preemptive selfing hypothesis, C. rattanii exhibit significantly earlier selfing in sympatric relative to allopatric populations.     

Floral traits and mating systems in sister species of <Emphasis Type="Italic">Nicotiana</Emphasis>: interpopulational variability and sympatry effects

Mating systems of Angiosperms are important determinants of population genetic structure and evolutionary potential. Nicotiana longiflora and N. plumbaginifolia are self-compatible, sister species, with contrasting floral morphology and can be found in allopatry and sympatry in North Argentina. In two sympatric and 10 nearby allopatric populations we studied their natural interpopulational variability and sympatry effects on corolla length and anther–stigma distance. We also estimated seed set by selfing and via pollinators using pollination treatments. Both corolla length and anther–stigma distance varied significantly among N. longiflora, but not among N. plumbaginifolia populations. We did not detect an effect of sympatry in either species for any of the floral traits studied. Pollination treatments suggest that N. longiflora is mainly an outcrosser, although selfing occurs at some extent. Seed set attributed to pollinators was significantly higher in N. longiflora whereas most seeds in N. plumbaginifolia were sired through self-pollination. In N. plumbaginifolia, selfing seems to assure reproduction in sympatric populations, where floral visitors have a strong preference for N. longiflora. Corolla length was significantly negatively correlated with an increase in the percentage of self-seeds estimated by pollination treatments. Within N. longiflora, anther–stigma distance showed a positive correlation with selfing.     

Does sympatry predict life history and morphological diversification in the Mexican livebearing fish Poeciliopsis baenschi?

Understanding why some species coexist and others do not remains one of the fundamental challenges of ecology. Although there is evidence to suggest that closely‐related species are unlikely to occupy the same habitat because of competitive exclusion, there are many cases where closely‐related species do co‐occur. Research comparing sympatric and allopatric populations of co‐occurring species provides a framework for understanding the role of phenotypic diversification in species coexistence. In the present study, we compare phenotypic divergence between sympatric and allopatric populations of the livebearing fish, Poeciliopsis baenschi. We focus on life‐history traits and body shape, comprising two sets of integrated traits likely to diverge in response to varying selective pressures. Given that males and females can express different phenotypic traits, we also test for patterns of divergence among sexes by comparing size at maturity and sexual dimorphism in body shape between males and females in each population type. We take advantage of a natural experiment in western Mexico where, in some locations, P. baenschi co‐occur with a closely‐related species, Poeciliopsis turneri (sympatric populations) and, in other locations, they occur in isolation (allopatric populations). The results obtained in the present study show that sympatric populations of P. baenschi differed significantly in life‐history traits and in body shape compared to their allopatric counterparts. Additionally, males and females showed different responses for size at maturity in sympatric conditions versus allopatric conditions. However, the amount of sexual dimorphism did not differ between sympatric and allopatric populations of P. baenschi. Hence, we conclude that not all traits show similar levels of phenotypic divergence in response to sympatric conditions. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100 , 608–618.     

Post-pollination reproductive isolation between diurnally and nocturnally flowering daylilies, Hemerocallis fulva and Hemerocallis citrina

To examine whether floral and post-pollination isolation develops independently or not, we conducted a crossing experiment between Hemerocallis fulva and Hemerocallis citrina that shows large floral divergence adapted for diurnal and nocturnal pollinators that have been believed to be fully cross-fertile. Flowers of the two species from sympatric populations were hand-pollinated with conspecific pollen from the same population (control), interspecific pollen from the same area (sympatric cross), and interspecific pollen from the different area (allopatric cross). After capsule dehiscence, the fruit set, seed set per fruit and seed set per flower were determined among three cross categories. The seed sets per flower were 32 and 77% lower in sympatric and allopatric crosses than in the control when H. fulva was the pollen recipient. There was no difference in three reproductive measures among the cross categories when H. citrina was the pollen recipient. This finding indicates that post-pollination isolation does exist between H. fulva and H. citrina, although it is partial, asymmetric, and weakened in sympatry. Our result suggests that floral and post-pollination isolation may develop independently, and reinforcement may not be a general phenomenon in plants.     

Calling song and selective phonotaxis in the field crickets,Gryllus firmus andG. pennsylvanicus (Orthoptera: Gryllidae)

The field cricket species, Gryllus firmusand G. pennsylvanicus,occur in a mosaic hybrid zone that roughly parallels the eastern slope of the Appalachian mountains in the northeastern United States. It is important to know what role, if any, the calling song plays in mate choice in sympatric and allopatric populations. In this report, we present results on the variability of calling song properties along transects across this hybrid zone. We also present the results of experiments on phonotactic selectivity of females from an allopatric population of G. firmus.The male calling song of allopatric G. firmuswas significantly slower in temporal rhythm (i. e., chirp and pulse repetition rates) and lower in pitch (i.e., dominant frequency) than that of allopatric G. pennsylvanicus.Calling song properties of males recorded in the hybrid zone varied considerably in temporal and spectral properties. In two-stimulus (choice) phonotaxis experiments, allopatric females of G. firmuspreferred synthetic calling songs with conspecific pulse repetition rates over songs that had lower and higher pulse rates. This preference persisted even when the sound pressure levels of alternative stimuli were unequal. Therefore, allopatric females of G. firmuscan discriminate between conspecific and heterospecific calling songs. Whether or not this same selectivity is present in sympatric populations remains unclear. Investigations of phonotactic selectivity in other allopatric and sympatric populations of both species are currently under way.     

Novel chloroplast DNA polymorphism in a sympatric region of two pines1

Nine hundred and two individuals from two populations in a Pinus banksiana — P. contora sympatric region were classified by restriction fragment length polymorphisms of two polymorphic chloroplast DNA markers. A large number of novel chloroplast DNA variants were identified which have not been observed in the allopatric ranges of the two parental species. Three apparently recombinant chloroplast DNA genotypes were discovered, in each of which one marker was typical of P. banksiana and the other was typical of P. contorta. These unusual sympatric chloroplast DNA's are evidence that the genetic complexity and rare variants of hybrid zones are not limited to the nuclear genome.     

Temperature Effects on the Temporal Properties of Calling Songs in the Crickets <Emphasis Type="Italic">Gryllus fultoni</Emphasis> and <Emphasis Type="Italic">G. vernalis</Emphasis>: Implications for Reproductive Isolation in Sympatric Populations

Two closely related wood-cricket species, Gryllus fultoni (Orthoptera: Gryllidae) and G. vernalis, produce similar calling songs, consisting of 3-pulse chirps. Analysis of field and laboratory recordings of calling songs showed that, after correction to a common temperature, there was a divergence in chirp and pulse rates between far allopatric populations of G. fultoni and populations sympatric with G. vernalis. To determine whether the divergence in calling songs potentially provides reproductive isolation between G. fultoni and G. vernalis throughout the temperature range over which these insects sing, we recorded calling songs of lab-reared G. fultoni and G. vernalis populations between 18 and 28°C. Mean chirp rate significantly differed between sympatric and far allopatric G. fultoni populations as well as between sympatric G. fultoni and G. vernalis populations. Although there was a significant difference in mean pulse rate between sympatric G. fultoni and G. vernalis populations, pulse rate did not differ between sympatric and far allopatric G. fultoni populations in the laboratory study. Considering the magnitudes of differences in calling song characters discriminated by females of G. fultoni and the mean differences and the variability in calling song characters between the two species, the joint difference in chirp and pulse rates may be adequate for species discrimination over most of the range at which these crickets breed.     

Effects of pollination and pollen source on the seed set ofPedicularis palustris

Habitat alteration can deteriorate plant-pollinator interactions and thereby increase the risk of population extinction. As part of a larger study on the effects of changes in land use on fen grassland vegetation, factors influencing the seed set of a short-lived, endangered wetland plant,Pedicularis palustris, were studied. We conducted field pollination experiments in one large and one small population. To investigate the effect of pollen source on seed set, individual flowers of caged plants were left unpollinated or were pollinated with pollen from the same flower, the same population or another population. To study pollen limitation and flower display, whole plants were subjected to pollinator exclosure, hand pollination or natural pollination. Self-compatibility was high, but differed between populations (61% and 97% of seed set after cross-pollination within populations). Cross-pollination between populations did not significantly alter seed number per capsule. Pollinator exclosure resulted in a very low seed set (<15% of natural seed set), despite high self-compatibility. The most likely explanations for high self-compatibility in combination with low autofertility are geitonogamy as reproductive assurance, selective neutrality of self-compatibility and phylogenetic constraints. Because of low autofertility, the seed set inP. palustris depends on pollinators. In the study populations, natural pollination was clearly sufficient for maximum seed production per plant, but seed set per capsule was significantly pollen-limited in the smaller population. Plants in this population also had a higher maximum percentage of simultaneously open flowers than those of the large population (31% vs. 13%), while flower longevity was generally extended without pollination. It is concluded thatP. palustris may influence pollinator behaviour and therefore the risk of pollen limitation by flower display.P. palustris showed a flexible reaction to differing pollination regimes without losses in overall seed set in the study populations.     

Character displacement in sailfin mollies, <Emphasis Type="Italic">Poecilia latipinna</Emphasis>: allozymes and behavior

Synopsis We analyzed variation in allozymes and mating preferences in 12 populations across much of the range of the sailfin molly, Poecilia latipinna. Sailfin mollies can be sympatric with its sexual parasite Amazon mollies, P. formosa. Amazon mollies must co-exist and mate with bisexual males of closely related species (including sailfin mollies) to induce embryogenesis but inheritance is strictly maternal. Where sailfin and Amazon mollies are sympatric there is evidence of reproductive character displacement as males show a significantly stronger mating preference for sailfin molly females over Amazon mollies compared to preferences of males from allopatric populations. From the allozyme data we found a moderate amount of genetic variation across all populations but this variation did not reveal significant partitioning between sympatric and allopatric populations. Additionally, we found no evidence for isolation by distance as genetic distance was not significantly correlated with geographic distance. While allozyme variation also did not significantly correlate with male mating preferences, there was a significant correlation between male mating preferences and geographic distance. This correlation between mating preferences and geographic distance may have arisen from coevolution with Amazon mollies resulting in reproductive character displacement. Taken together, the distribution of genetic and behavioral variation among sympatric and allopatric populations suggests that behavioral evolution has outpaced evolution at the allozyme loci we examined in P. latipinna.