Morphological variation and female reproductive success in two sympatric Trillium species: evidence for phenotypic selection in Trillium erectum and Trillium grandiflorum (Liliaceae)

I investigated the mating systems and phenotypic variation of two sympatric spring ephemerals, Trillium erectum and T. grandiflorum (Liliaceae), and phenotypic selection acting through female reproductive success for 11 morphological characters in five sympatric populations of the two species. I examined the degree of self-compatibility, pollinator-visitation rates, and pollen limitation of fruit and seed production in both species. Both Trillium species were self-compatible, but outcrossed flowers produced more successful fruits and seeds than self-pollinated flowers. Pollinator-visitation rates to the two species were low compared to other insect-pollinated spring ephemerals. In addition, both T. erectum and T. grandiflorum experienced pollen limitation in fruit and/or seed production; however, levels of fecundity in both species may be influenced by resource availability as well. I found significant phenotypic variation in 11 morphological characters within and among the five study populations. The sizes of all morphological characters were positively correlated. In general, larger T. erectum and T. grandiflorum produced more seeds. Phenotypic selection analysis revealed that direct and indirect selection acted on the size of morphological characters for both species. But there was no detectable selection acting on plant shape. This study reveals that variation in plant size exists within and among populations of both species, and this variation is associated with variance in female reproductive success. Spatial and temporal variation in pollinator and/or resource abundance may play a role in the phenotypic variation exhibited by both Trillium species.     

Revision of Trillium subgenus Phyllantherum (Liliaceae)

A taxonomic revision, based mainly on morphological and distributional criteria, is presented for all taxa ofTrillium subgen.Phyllantherum Raf., which is characterized by sessile flowers. The 22 species recognized as valid include 17 morpho-geographical species in the eastern United States and five in the northwestern United States and California. These species form three morphological alliances that are characterized informally as “species groups.” All the taxa are described, and a key to species and varieties is provided. Five new species are described:Trillium decipiens, T. reliquum, T. foetidissimum, T. albidum, and T. kurabayashii; and a new status,T. angustipetalum (Torrey) Freeman, based onT. sessile var.angustipetalum Torrey, is proposed. Six new flower color forms are described:T. reliquum f. luteum, T. stamineum f. Iuteum, T. maculatum F. luteum, T. maculatum f. simulans, T. foetidissimum f. luteum, and T. cuneatum f. luteum. County distribution maps are included for all species and varieties, and an annotated synonymy is presented for each taxon.     

Stigmatic Self-Incompatibility and Mating Patterns in Trillium grandiflorum and Trillium erectum(Melanthiaceae)

Post-pollination processes governing mating patterns in Trillium,a well-known genus of insect-pollinated woodland herbs, arepoorly understood. Mechanisms influencing outcrossing were investigatedin T. grandiflorum and T. erectum, two widespread species nativeto eastern North America. In southern Ontario, Canada, the twospecies are often sympatric; they flower in early May, and arepollinated by different assemblages of insects. Controlled cross-and self-pollinations and structural observations of pollengermination and pollen tube growth were conducted to determinewhether the two species possess a self-incompatibility (SI)system and, if so, the specific site(s) of self-rejection. Controlledpollinations indicated that both species set significantly moreseeds from cross-pollination than self-pollination, implicatingthe action of SI. This was confirmed by structural studies whichdemonstrated that self-recognition and rejection reactions occurredon dry-type stigmatic papillae. Observations of pollen hydrationrevealed that self-rejection was rapid, being initiated within10 min of pollination and prior to pollen tube emergence. Finalself-rejection resulted in failure of pollen tube growth atthe base of stigmatic papillae. SI was expressed more weaklyin T. erectum and thereby resulted in considerable self-seedset in some individuals . Estimates of outcrossing rates usingallozyme markers indicated that T. erectum displayed a mixed-matingsystem whereas T. grandiflorum was more highly outcrossed. Structuralstudies of pollen traits indicated that the two species differedwith respect to the size of grains and their aggregation withimplications for pollen dispersal and mating. The ecologicaland evolutionary implications of the variable expression ofSI in Trillium are discussed. Copyright 2001 Annals of BotanyCompany Trillium grandiflorum, Trillium erectum, self-incompatibility, mating     

Seed dispersal in Erythronium grandiflorum (Liliaceae)

Primary and secondary seed dispersal was investigated for the glacier lily Erythronium grandiflorum in the Colorado Rocky Mountains. These heavy seeds have no obvious adaptations for biotic or abiotic dispersal, but can be thrown short distances when the dehiscent fruits are shaken by wind. We used sticky traps to measure primary transport of seeds up to 1 m away from individual plants. A seed cafeteria experiment examined the role of ants and rodents in secondary seed transport. Primary dispersal by wind was positively skewed and median transport distances were influenced by variation in plant height. Secondary dispersal was negligible compared to Viola nuttallii, an elaiosome-bearing species. Thus, seed dispersal was highly restricted in E. grandiflorum, and a 1 m radius encompassed the modal section of the seed dispersal curve. The seed dispersal component of gene flow was quantified and combined with previous measurements of pollen flow to yield a more complete estimate of Wright's neighborhood size, N _e, for E. grandiflorum. The lack of a special seed dispersal mechanism in E. grandiflorum is discussed in terms of a source-sink model for seedling establishment with respect to distance from the parental plants.     

A new Trillium (Liliaceae) from the northwestern United States

A new species,Trillium parviflorum, is described inTrillium subgen.Phyllantherum Raf.Trillium parviflorum is easily distinguished fromT. albidum Freeman, which it superficially resembles, by its overall smaller size, much smaller linear-lanceolate petals, clove-like fragrance, and dark maroon fruit.     

Genetic Structure in a Heterocyanic Population of Trillium sessile (Liliaceae)

Abstract Species of Trillium in the subgenus Phyllantherum are either polymorphic for flower color, or monomorphic for flower color and related to a polymorphic species. This leads to the suggestion that polymorphic species may be the progenitors for monomorphic ones. For this to be true, it must be demonstrated that genetic divergence among flower morphs can occur within polymorphic populations. Genetic structure was assessed in a population of T. sessile that contains a polymorphism for flower color. A survey of 11 enzyme systems using starch gel electrophoresis revealed three polymorphic loci: 6PGD-1, AAT-1 and AAT-2. Analysis of large and small scale spatial structure, stage classes, and flower color classes revealed significant genetic divergence in all instances. Spatial structure in the population is likely a result of genetic neighborhoods which can maintain populational variation via random genetic drift. Genetic divergence of the yellow flower color morph was probably initiated through genetic drift since the morph occurs in low frequencies. The results imply that the initial genetic divergence of species in the subgenus can arise within polymorphic populations.     

Pollen viability,vigor, and competitive ability in Erythronium grandiflorum (Liliaceae)

To determine how the capability of pollen is affected by lengthy exposure to field conditions, we obtained pollen samples (from Erythronium grandiflorum) that were fresh (from newly dehisced anthers) or aged, from 6 to 24 hr. Without pretreating the collected grains, we compared their abilities 1) to retain viability, as indicated by the Heslop-Harrison fluorochromatic reaction (FCR) test; 2) to retain germinability, as indicated by pollen-tube growth in vitro on Brewbaker-Kwack medium; 3) to sire seeds when used as a pollen donor on recipient flowers in the field; and 4) to sire seeds competitively when mixed with another donor pollen in competitive pollinations. FCR scores declined drastically with pollen age. Germinability typically declined with age, but less drastically; the correlation between FCR and germinability ranged from strong to nil in different trials. The sharp decline in FCR was probably due to nonlethal, reversible desiccation. Seed-siring success, both in competitive and noncompetitive trials, bore no relation to pollen age or to germinability. Any loss in seed-siring ability in E. grandiflorum pollen over 24 hr is apparently overwhelmed by other sources of variation. The lack of correlation between seed-siring ability and FCR or germinability scores means that those tests should not be used to estimate overall pollen competence in this species. They should be so used in other species only if experiments have demonstrated their greater reliability.     

Trillium sulcatum (Liliaceae), a new species of the southern Appalachians

Trillium sulcatum, a new species with two petal color forms, was first circumscribed by Barksdale as a variety in 1938, but without a Latin diagnosis. It superficially resembles the mainly northernT. erectum and appears to be closely related to the primarily midwesternT. flexipes. Its distinguishing features are ringent flowers, short sepals, and long pedicels; additional characters are presented in a chart contrasting it with species with which it has been confused. Distribution suggesting relictual significance is documented. Except that intermediates withT. flexipes are occasional,T. sulcatum maintains its distinctiveness throughout its southern Appalachian range.     

Post-glacial history of Trillium grandiflorum (Melanthiaceae) in eastern North America: inferences from phylogeography

Dispersal and migration are important processes affecting the evolutionary history and genetics of species. Here we investigate post-glacial migration and gene flow in Trillium grandiflorum (Melanthiaceae), a wide-ranging, forest herb from eastern North America. Using phylogeographic approaches, we examined cpDNA and allozyme diversity in 35 populations of T. grandiflorum sampled from throughout the geographic range of the species. Nested clade analysis (NCA) of cpDNA haplotypes indicated that T. grandiflorum likely survived in two refugia in the southeastern US during the last glaciation and that long-distance dispersal characterized the post-glacial recolonization of northern areas. There was no evidence for reduced allozyme diversity in populations from glaciated compared to ice-free regions, probably because of the greater abundance and larger effective size of populations in the north. An analysis of isolation-by-distance based on the allozyme data suggested a pattern of population differentiation consistent with restricted gene flow. Notwithstanding the significance of rare seed dispersal events for migration, a comparison of allozyme and cpDNA genetic structure indicates that pollen flow between populations is more likely than seed dispersal. These results for T. grandiflorum represent the first phylogeographic analysis of a temperate woodland herb in eastern North America and support the importance of occasional long-distance dispersal events in the post-glacial migration of plants.     

A demographic study of deer browsing impacts on Trillium grandiflorum

When white-tailed deer populations reach high densities, they have negative and often dramatic effects on forest herb populations. However, it is not clear how deer affect the demographic processes of plant populations. We examined how the structure and dynamics of Trillium grandiflorum (Michx.) Salisb. populations are affected by deer browsing in the Upper Great Lakes region by sampling populations from nine study sites in a forested landscape in 1998 and 1999. We constructed a stage-based matrix population model for the regional population. Our model indicated that the long-term growth rate of the population to be –3.56% per year ( = 0.965). Mortality rates were highest for seeds (97.5%) followed by seedlings (29.1%), and lower for all remaining stage classes (4.9 to 8.5%). The observed stage distribution significantly differed from the stable stage distribution, and the damping ratio ( = 1.103) indicated the population would not reach its stable stage distribution anytime soon. In the absence of deer browsing, the long-term growth rate would improve to between –3.46% and –1.61% per year. A moderate drought during the study could account for the negative population growth rate, but deer browsing accelerates the rate of decline. Population growth is most sensitive to the proportion of plants remaining in the nonflowering stage, and deer browsing reduces this proportion. Browsing damage was relatively low in this study (5.4% of stems in 1998, 11.5% in 1999) compared to another study of browsing impacts on T. grandiflorum, indicating deer could have far more severe demographic consequences in populations subject to higher levels of browsing.     

Life History Studies on the Genus Trillium (Liliaceae) I. Reproductive Biology of Four Japanese Species

Abstract Four Japanese Trillium (Liliaceae) species which are representative perennial herbs of the temperate deciduous forests— Trillium kamtschaticum (2x), T. apetalon (4x), T. tschonoskii (4x) and T. smallii (6x)—were studied for their reproductive characteristics, e.g., patterns of reproductive resource allocation and reproductive output.
In spite of the differences in ploidy levels (from 2x to 6x), all four species showed very similar reproductive traits. It became evident that in response to the increase in reproductive allocation to total reproductive organs (RA), the number of seeds produced per plant (P_N) clearly increased. This trend is well in accord with the relationship found in four North American species and also several temperate woodland perennial herbs which occur in closed, stable and predictable environments, and possess typical xenogamous breeding systems. Although there is no conspicuous trend between ploidy levels and P_N, one of the significant differences noted in this study was in seed weight, which ranged from 2.93 mg in diploids, to 3.42–3.45 mg in tetraploids, and to 4.47 mg in hexaploids.     

Life History Studies on the Genus Trillium (Liliaceae) V. Reproductive Biology and Survivorship of Three Declinate-flowered Species

Absract Three pedicellate-declinate-flowered species of Trillium (Liliaceae), T. vaseyi and T. flexipes , were studied for their life history characteristics, e.g., stage class structures of natural populations and reproductive features, including energy allocation to reproductive activities. The populations structures of all three species showed similar depletion structures characterized by a conspicuous decrease of individuals in the small juvenile stages, as was also observed in pedicellate-erect-flowered Trillium species However, with respect to reproductive characteristics, these three declinate-flowered species showed different features from erect-flowered species, although they belong to the same pedicellate-flowered group. That is, these declinate-flowered Trillium species exhibited low seed setting rates of 30% in T. catesbaei , 45% in T. vaseyi and 34% in T. flexipes , suggesting that they possess different mating systems from erect-flowered species which showed high seed setting rates of 50–90%.     

Impact of tree leaf phenology on growth rates and reproduction in the spring flowering species Trillium erectum (Liliaceae)

We investigated the impact of overstory tree leaf phenology on growth rates, carbon allocation pattern, and fruit characteristics in the spring flowering species, Trillium erectum (Liliaceae). Air temperature, overstory canopy closure, and T. erectum phenology were monitored at three locations following a latitudinal gradient in Québec, Canada. Northern sugar maple trees leaf out at cooler temperatures than more southern populations, while Trillium development was initiated at the same soil temperature irrespective of the latitude. Therefore, in northern areas, the time between initiation of T. erectum leaf expansion and canopy closure was shorter than in southern areas, which left less time for northern plants to accumulate reserves before canopy closure. Differences in growth patterns were noted between T. erectum populations. From a south-north gradient, investment to reproduction, total plant biomass, and annual growth rate decreased, while specific leaf area and stem height increased, indicating shade acclimation. The length of the high light period in early spring seems to be a determinant for spring flowering plants' growth and reproduction and may explain the northern distribution limit of some of these species.     

Postfertilization causes of differential success of pollen donors in Erythronium grandiflorum (Liliaceae): nonrandom ovule abortion

Seed paternity in Erythronium grandiflorum does not fully reflect the proportion of pollen on the stigma. When two types of pollen are simultaneously applied to the stigma, outbred seeds are produced over inbred, and seeds from more distant donors are produced over seeds from donors nearby. I looked for postfertilization causes of these previously reported patterns of differential success of pollen donors. I simultaneously pollinated stigmas with pollen from two donors and observed ovule development through a window sliced in the ovary. Pollen donor pairs were self and cross, donors 1 and 100 m from the recipient, and two donors each 100 m from the recipient. Since one donor was always the alternate homozygote from the recipient at the malate dehydrogenase locus, I could determine the paternity of developing seeds. When it appeared that ovules were aborting, I removed them and determined their paternity using starch gel electrophoresis. Ovules fertilized by self pollen were more likely to abort than ovules fertilized by cross pollen, and ovules fertilized by nearby donors were more likely to abort than ovules fertilized by distant donors. Ovules fertilized by donors 100 m from the recipient were equally likely to abort. There was not a significant relationship between the proportion of ovules fertilized by a pollen donor and the probability of those fertilized ovules developing into seeds. There was no relationship between ovule position within a fruit and ovule abortion. I manipulated available resources by removing leaves and by permitting only one fruit to develop per plant. Decreasing the amount of resources increased the proportion of aborted ovules. Abortion of ovules of lesser quality appears to release resources that can then be used to develop other offspring.     

Effects of chronic herbivory and historic land use on population structure of a forest perennial,Trillium catesbaei

Question: How have long-term herbivory and past land use impacted the population structure of Trillium catesbaei, a long-lived rhizomatous herb? Location: Western Great Smoky Mountains National Park, Tennessee, USA. Methods: We examined T. catesbaei populations at three sites: (1) Cades Cove (CC), an area of intensive historic land use that has been maintained as open fields and woodlots with a history of chronic deer herbivory, (2) Whiteoak Sink (WOS), a reference area with similar land-use history, geology, and soils that has succeeded to closed-canopy forest with relatively low levels of deer herbivory, and (3) Leadbetter Ridge (LBR), an area of primary forest that has never received significant anthropogenic disturbance. Trillium catesbaei is the most common Trillium species at the three study sites, but smaller in stature, shorter lived, and more of a habitat generalist. Results: Chronic herbivory in CC has created a highly-truncated age structure with no plants older than 9 years, while plant ages at the other sites were more evenly distributed. Compared to WOS, plants in CC were younger at a given height and more likely to flower when younger. Across all life stages, populations at CC contained 68 × fewer plants than WOS. The age structures of WOS and LBR were similar. Compared to published age estimates for other Trillium species, our results suggest that T. catesbaei is relatively short lived within the genus. Conclusions: Chronic herbivory had pronounced effects on the population structure of a perennial herb. Other long-lived herbaceous species may exhibit similar truncated age structures and flowering by younger and smaller plants. Habitat generalist species within a genus, such as T. catesbaei, that are able to reproduce more quickly may persist longer under chronic herbivory. However, chronic herbivory has likely caused the loss of herbaceous species from CC and may eventually cause the local extirpation of T. catesbaei populations.     

Structural variability and chromosome numbers variation in natural populations ofScilla autumnalis (Liliaceae)

Chromosome variability within three populations ofScilla autumnalis was analysed. Among a total of 167 individuals we have detected one triploid structural heterozygote, 26 B-containing (with two different B-types), 11 structural variants (in the fifth chromosome pair), and one asynaptic. The possible relation between both types of chromosome variability (structural and chromosome numbers) is discussed.     

Pollen morphology and its evolutionary significance in Hemerocallis (Liliaceae)

Pollen morphology of 10 species of Hemerocallis was investigated with LM and SEM. In addition to pollen with reticulate sexine, a new verrucate sexine pattern is found in three Chinese endemic species, H. plicata, H. forrestii and H. nana . Between the two typical sexine patterns, there are transitional variants. Also, a new pollen shape, boat-shaped-elliptic, is observed exclusively in H. nana , besides-oblong shape in other species. Phylogenetically, the common reticulate sexine pattern would be more primitive than the verrucate one. The typical reticulate sexine pattern would be basic, from which transitional variants would have derived and finally led to the typical verrucate pattern. Similarly, the common boat-shaped-oblong pollen would be more primitive than the -elliptic one, from which the latter would have derived during phylogenetic development.