Introduction

Willamette Valley wetland prairies are some of the most endangered ecosystems in the United States, and provide habitat for many federally listed species (Wilson et al. 1993, Noss et al. 1995, USFWS 2000, Schultz et al. 2003). A rigorous program of ecosystem restoration can protect and enhance wetland plants, animals, and services (Pywell and Putwain 1996). The restoration projects of the West Eugene Wetlands Program (WEWP) comprise one of the few large-scale and long-term integrated restoration programs in the world.

Successful ecosystem restoration requires establishing and maintaining native plants. In turn, plant establishment hinges on having suitable environmental conditions, using species with adequate germination and growth rates, and reducing competitive pressure from non-native plants (Figure 1). In year one of this project, we synthesized the wealth of plant establishment data during wetland restoration in the West Eugene Wetlands Program (Wilson 2004). In year two, our objectives were to expand on these results in several important ways: • Generalize these results through the investigation of plant traits (Table 1) that consistently correspond to the patterns of establishment and vigor (Figure 1). • Systematically compile the results into a trait database. This database includes findings from similar ecosystems, both in the Willamette Valley and elsewhere. • Testing the effect of habitat variation on the relationship between traits and seedling establishment patterns.

Our goal is to predict key aspects of prairie restoration performance, in this case establishment rates, based on species traits and habitat. These predictions can then be applied as management recommendations, such as which species to sow to maximize native plant abundance at a given site, even if the species have not been field tested.

The two components of our project–plant traits and the database–are crucial to this goal. • Without the generalization that traits allow, understanding of wetland restoration increases slowly and expensively, one case study at a time. • The organization of the database will increase the power and efficiency of revealing the relationships between plant traits and plant performance. Perhaps even more important is the role of the database in developing a Web-based expert system for managers wishing to plan wetland restorations.

Species and establishment rates

For this project we expanded the Willamette Valley Prairie Plant Trait Database to include additional species important to the WEWP restoration projects and additional traits important to understanding and prediction establishment success.

Traits, Final Report Page 2 Wilson, Clark and Roberts We selected 31 species (Table 2) that represent the range of establishment rates found among all species in the WEWP restoration sowings (Tables 3-5), a range of sowing rates, and a variety of traits. Establishment rate for this project is defined as cover in the second year after sowing per weight of seed sown per area (Wilson 2004). The units of establishment rate are cover (%) per g seed sown per cm2.

The highly skewed establishment rates required transformation to meet the assumptions of statistical analyses. Establishment rates for all species within a data set (Vernal pool habitats, Deschampsia-dominated habitats, Emergent wetland habitats, all habitats combined) were rank- transformed. Higher ranks represent higher establishment rates. Because various number of species established in each data sets (Table 4), the value for the top rank varied. In practice, several species had zero establishment within a habitat, so tied for lowest establishment with a rank >1. The ranks for the 31 test species were extracted from this larger set of species ranks. Ranks for species with zero establishment were further differentiated into species that established in no habitats, which were given a rank of 0 (Table 6).

The WEWP has its own Seeding Assessment rating system based on perceived success (E. Wold, pers. comm.). We updated these ratings of species with current propagation information (WEWP 2005) to create an additional ranking of the test species (Table 6).

Traits

The Willamette Valley Prairie Plant Trait Database is a compilation of ecological information on species important in upland prairies, wetland prairies, vernal pools, and emergent wetlands. The information is directly pertinent to prairie and wetland restoration. The Database currently contains more than 3800 data points on 134 species. The Database provided information on many of the traits of interest to this project (Table 1), such as clonal spread and growth form. We greatly expanded information in the Database for the 31 test species by measuring seed characteristics and plant growth under standardized growth chamber conditions.

Methods for growth analysis

Growth chamber procedures followed the general recommendations of Hendry and Grime (1993). Standardized conditions include specifications for germination media, transfer of germinants, pot size, growing media, nutrient solutions, growing illumination and temperatures, and dates of harvest. The use of standardized conditions allows us to integrate our results with those in the scientific literature.

We used appropriate dormancy breaking strategies and optimal germination conditions to provide approximately 40 germinants for each test species. When the radical of the germinant was 1 mm long, we transferred the germinant to prepared pots filled with water-washed sand. Each pot was watered with 5 ml of Hoagland's solution three times per week. The growing

Traits, Final Report Page 3 Wilson, Clark and Roberts regime in the growth chamber (Table 7) followed the general recommendations of Hendry and Grime (1993) and was monitored continuously with automatic sensors.

Seedlings were harvested at 7 days or 21 days. Leaf area was measured from scanned images of fresh leaves taped flat. Root mass, shoot mass, and leaf-only mass were determined by weighing material that had been dried at 80 C for 48 hours.

Measurement of seed mass, dimensions, and laboratory germination rate

We measured seed mass in five replicate batches of 100 seeds, after drying at 80 C for at least 48 hr. Included in the mass measurements is the seed, proper, and any fruit structures that disperse with the seed, such as awns and perigynia. Thus mass measurements are of dispersules.

Seed (dispersule) dimensions were measured so that we could calculate sphericity. Length, width, and thickness of the dispersule were measured with a caliper or micrometer. Sphericity was calculated as the variance of these three measurements, after each had been divided by the largest of the three values (Thompson et al. 1993).

We determined laboratory germination rates under favorable conditions for each species. Seeds of each species were sowed into germination boxes that were lined with thick germination paper and filled half-way with moistened sand. For those species requiring stratification (Table 8), seeds were allowed to imbibe, then place at 2 C - 5 C for the required number of weeks. For those species requiring scarification, we removed small chips of testa by gently rubbing seeds between medium sand paper.

Prepared seeds were added to germination boxes that were half-filled with washed sand. Temperature and light conditions were set to those favorable to the species being tested (Table 8).

Trait values for species

The new measurements and information from the Database provided for the 31 test species values in seven categorical traits, seven growth-analysis traits, and six other quantitative traits (Tables 9 and 10). Notice that these traits are characteristic of the species and independent of the particular field conditions at the WEWP restoration sites.

Relationships between plant traits and field establishment rates

Our prime objective is to determine which plant traits consistently correspond to the patterns of establishment, and to test the effect of habitat variation on these relationships. We examined these relationships with a series of statistical models.

Traits, Final Report Page 4 Wilson, Clark and Roberts The standardized traits varied widely in their relationships with field establishment rates, both overall and in the three habitats (Table 11). The strongest relationships were with the categorical traits dealing with form (Life form and Growth form) and with the quantitative traits of Flowering peak month and Seed mass. Flowering peak month and Seed mass were also consistent in the direction of relationship across habitats: later flowering and smaller seeds were always associated with higher establishment rates. Growth analysis traits provided little explanatory power for establishment rates – a real puzzle considering that early growth should be crucial for establishment success..

Nature is multivariate, so we developed multivariable models of field establishment rates vs. standardized traits. Stepwise procedures and model comparisons were used to select the most

parsimonious model with highest explanatory power, using a Mallows Cp procedure.

The results models successfully explained 50%-70% of the variability in the four data sets (Table 12). This high level of explanatory power shows that standardized traits can explain patterns of field establishment. Moreover, the traits that explain variability were rather consistent across models.

Flowering peak month

Flowering peak month was significant as a single variable in all four data sets, and was part of the final model for three (All habitats, Vernal pool habitats, and Deschampsia-dominated habitats). In each case, as typical flowering occurs later in the year, field establishment in the WEWP restoration sites increased (Figure 2). Species that flowering one month later have, on average, a rank in establishment rate seven slots higher.

Why might this be happening? High rates of establishment, as measured here, involve not only germination and the establishment of the germinant. For perennials, high establishment rates required vegetative survival and growth until monitoring occurred in the second year after sowing. For annuals, second-year plants needed to grow quickly enough to flower and produce seeds. Flowering time is directly related to time of vegetative growth for most species. We speculate that species that flower early are also active earlier in the season and are more susceptible to environmental threats from erratic water levels, disease, and herbivory (especially molluscs).

Perenniality / Life form

Perenniality was a variable in the final model for All habitats and Life form was a variable in the final model for Vernal pool habitats. Both traits are reflecting a similar pattern (Table 13), with annuals/therophytes establishing at much higher rates than perennials or other life forms. This pattern is also seen in Figure 1, where the annuals are all above the regression line. In fact, annuals consistently had higher establishment rates than perennials that had the same Flowering peak month. Geophytes established consistently poorly (Tables 3-6, 13), a phenomenon widely recognized by frustrated practitioners.

Traits, Final Report Page 5 Wilson, Clark and Roberts Laboratory germination rate

The higher the germination rate under standard laboratory conditions, the higher the establishment rate under field conditions in Vernal pool habitats (Table 11, Figure 3). The interpretation of this pattern is straightforward: the higher the inherent seed viability and ability to germinate, the more likely seedings will occur and grow. It is perhaps more noteworthy that Laboratory germination rate is not related to establishment rates in Deschampsia-dominated and Emergent wetland habitats.

Growth form

Grasses had higher establishment rates than other growth forms in the Deschampsia-dominated and Emergent wetland habitats (Table 13). Grasses tend to have broad germination conditions, providing an advantage to establishment. The ability of many of these grasses to tiller soon after establishment further decreases their risk of mortality.

Dicot forbs had the second highest establishment rates in both habitats (Table 13). The four annuals, each of which established at relatively high rates, are all dicot forbs.

Seed mass

Seeds of lower mass tended to have higher establishment rates (negative slope, b, in Table 11). Seed mass was a key factor in the Emergent wetland habitats (Table 12, Figure 4), also with a negative slope. Ecological theory suggests that a well-provisioned seed will grow and survive better. But there is a trade-off between seed size and seed number. Sowing rates used in this study are by weight (g/cm2). So species with lighter seeds would have had more seeds for a given sowing rate. Therefore, in the WEWP restoration sites, seed number was more important in maintaining high establishment rates than was seed provisioning.

WEWP Seeding Assessment

The relationship of traits to the WEWP Seeding Assessment ratings differed a bit from the previous results, which used establishment rate as the response variable. This difference is expected, because the Seeding Assessment is a subjective rating of dominance, not a measure of establishment per sowing rate. The final, parsimonious model accounted for 66% of the variability in Seeding Assessment ratings (Table 14). Clonal spread was a key trait, with non- clonal plants having significantly higher establishment rates. Non-clonal plants include all the annuals, each of which established at relatively high rates, but also herbaceous perennials such as Grindelia integrifolia and Microseris laciniata.

Seed mass was also a significant factor (Table 14). Species with smaller seeds tended to have higher Seeding Assessment ratings. This pattern is similar to those seen with establishment rates, as is the explanation: smaller seeds means more seeds per weight of seeding.

Traits, Final Report Page 6 Wilson, Clark and Roberts Typical maximum height was the third factor in the final model. We suggest that because Seeding Assessment ratings include observations beyond the second year, the ability to grow tall and shade neighbors produces an advantage here when it does not for establishment rates.

Discussion and management implications

Are the results consistent across habitats?

The results are generally consistent across habitats. Key traits analyzed as single variables tend to show strong relationships with establishments rates in at least two of the three habitats (Vernal pool habitats, Deschampsia-dominated habitats, and Emergent wetland habitats). This is the case with the categorical traits of Life form, Life form (expanded), and Growth form and the quantitative traits of Flowering peak month and Seed mass. Moreover, the final models have broad overlap in structure (the factors involved) and predictions (the coefficients in the relationships). For example, Flowering peak month was in the final model of two of the three habitats, and it always had a positive coefficient.

But habitats are still unique. For example, Laboratory germination rate was only important in Vernal pool habitats, but it was the single best predictor of establishment rate in that habitat.

How can the results be used by management?

A key decision when selecting species for a seeding mix is getting high establishment rates. Few sources of information are available to make this decision, however, outside of experience and hearsay. This situation means that there is little basis for predicting the success of untested species. Given a new species under consideration for inclusion in a seeding mix, the only basis for predicting it establishment rate in the West Eugene Wetlands restoration sites would be the overall mean. For all habitats, this is 1.7% cover per g/cm2 seed sown, with 95% confidence interval of 0.0 to 411.6. Such a wide confidence interval (the entire range of establishment rates!) is of little use.

The trait models developed in this study can provide better predictions. Consider, for example, a new species of perennial and a new species of annual, both with peak flowering around June 30 and seed sphericities of 0.15. The model for All habitats (Table 9, Figure 1) predicts that the perennial will have a mean establishment rate of 0.5% cover per g/cm2 seed sown, with a 95% confidence rate of 0.3 to 0.7. The prediction for the annual is a mean establishment rate of 24.1% cover per g/cm2 seed sown, with a 95% confidence rate of 2.0 to 411.6. The annual is the better bet, with 50× higher establishment predicted over the perennial.

Do not be distracted by the large confidence interval for the new annual species. The larger confidence interval is a by-product of the highly skewed establishment rates in the original data set. The model has still been useful by distinguishing biologically significant differences in establishment rates (0.5 vs. 24.1). The model has increased the precision of the predictions by

Traits, Final Report Page 7 Wilson, Clark and Roberts partitioning the extreme range of the confidence interval for the naive estimate (0.0 to 411.6) into narrower confidence intervals for the separate predictions for perennials (0.3 to 0.7) and annuals (2.0 to 411.6). Understanding the relationship between easily measured, standardized traits and field performance yields more accurate predictions of actual establishment rates and more reliable (precise) predictions of establishment rates.

Next steps

The relationships between standardized traits and field establishment rates were strong. The next step is to validate these results, with species in the WEWP restoration sites not used to develop the relationships and with other data sets from similar habitats. Once the relationships have been validated, then these findings can be converted into an expert system for managers. Such a system would combine the trait information in the Willamette Valley Prairie Plant Trait Database with the equations from our statistical models. Managers and others wishing to select species for restoration projects would then be able to query the expert system, both to determine if a species was suitable for a given site or to determine which traits to look for when selecting new species.

Acknowledgments

We acknowledge the valuable contributions of Diana Wageman in the collection of the plant trait data. We appreciate the help of Sherry Harlan. We thank the Bureau of Land Management, Eugene District, in particular the assistance from Dharmika Henshel, for the contribution of seed. Amy Bartow of the Plant Materials Center kindly shared her extensive knowledge of germination requirements of wetland species. We thank Dale Brown and the Oregon State University Seed Laboratory for the use of their excellent facilities.

References

Drake, D. K., K. Ewing, and P. Dunn. 1998. Techniques to promote germination of seed from Puget Sound prairies. Restoration and Management Notes 16:33-40.

Flynn, S., R. M. Turner, and J. B. Dickie, J. B. 2004. Seed Information Database (release 6.0, Oct. 2004). http://www.kew.org/data/sid.

Guerrant, E. O. and A. Raven. 1995. Seed germination and storability studies of 69 plant taxa native to the Willamette Valley wet prairie.A publication produced under a cost share agreement with TNC, Lewis and Clark College, The Berry Botanic Garden, and BLM.

Hendry, G. A. F. and Grime, J. P. 1993. Methods in comparative plant ecology. Chapman & Hall, London.

Traits, Final Report Page 8 Wilson, Clark and Roberts Noss, R. F., E. T. LaRoe III, and J. M. Scott. 1995. Endangered ecosystems of the United States: A preliminary assessment of loss and degradation. National Biological Service, Biological Report 28.

Pywell, R. F. And P. D. Putwain. 1996. Restoration for conservation gain. Pages 203-221 in Conservation Biology, I. F. Spellerberg (ed.), Longman, London.

Schultz, C. B., P. C. Hammond, and M. V. Wilson. 2003. The biology of Fender's blue butterfly (Icaricia icarioides fenderi) an endangered species of western Oregon native prairies. Natural Areas Journal 23:61-71.

Thompson, K., S. R. Band, and J. G. Hodgson. 1993. Seed size and shape predict persistence in soil. Functional Ecology 7:236-241.

US Fish and Wildlife Service. 1988. National list of vascular plant species that occur in wetlands. US Fish & Wildlife Service Biological Report 88. Updated in 1993 and 1996.

US Fish and Wildlife Service. 2000. Endangered status for Erigeron decumbens (Willamette daisy) and Fender's blue butterfly (Icaricia icarioides fenderi) and threatened status for Lupinus sulphureus spp. kincaidii (Kincaid's lupine). Federal Register 50 CFR Part 17:3875-3890.

West Eugene Wetlands Program. 2005. Propagation strategy list for plants that grow in the West Eugene Wetlands. http://www.eugene-or.gov/.

Wilson, M. V., K. P. Connelly, and L. E. Lantz. 1993. Plant species, habitat, and site information for Fern Ridge Reservoir. Report to the Waterways Experiment Station, US Army Corps of Engineers.

Wilson, M. V. 2004. Patterns of establishment success in West Eugene Wetlands Program restoration sites. Final report to the Bureau of Land Management, Eugene District. 42 pages.

Traits, Final Report Page 9 Wilson, Clark and Roberts Table 1. Plant traits and the ecological functioning or behaviors they correspond to in the field. Trait Functioning or behavior in establishment Growth form and life form space-holding ability resource acquisition competitive ability Clonal spread space acquisition resource acquisition perennation Dormancy breaking mechanism establishment success longevity in seed bank Life history (including plant longevity perenniality) space-holding ability disturbance tolerance Wetland Indicator Status affinity for hydric conditions Seedling growth analysis establishment - relative growth rate growth - unit leaf rate fecundity - leaf mass ratio competitive ability - specific leaf area space-holding ability - root/shoot allometric coefficient plasticity - shoot biomass stress tolerance - leaf area leaf longevity Flowering phenology resource acquisition and availability seed provisioning strategies stress avoidance disturbance avoidance Germination rate establishment success Height competitive ability generation time Seed (dispersule) mass dispersal distance longevity in seed bank establishment success fecundity Seed (dispersule) sphericity incorporation and longevity in seed bank

Traits, Final Report Page 10 Wilson, Clark and Roberts Table 2. Species used to test the relationships between standardized traits and field establishment rates in West Eugene Wetlands Program restoration sites. Scientific name, authority (common name) Agrostis exarata Trin. (spike bentgrass) Allium amplectens Torr. (narrowleaf onion) Beckmannia syzigachne (Steud.) Fern. (American sloughgrass) Camassia leichtlinii (Baker) S. Wats. (large camas) Camassia quamash (Pursh) Greene (common camas) Carex densa (Bailey) Bailey (dense sedge) Carex unilateralis Mackenzie (lateral sedge) Danthonia californica Boland. (California oatgrass) Deschampsia caespitosa (L.) Beauv. (tufted hairgrass) Dichanthelium acuminatum var. fasciculatum (Sw.) Gould & C.A. Clark var. (Torr.) Frecmann (western panicgrass) Downingia elegans (Dougl. Ex Lindl.) Torr. (elegant calicoflower) Downingia yina Applegate (cascade calicoflower) Epilobium densiflorum (Lindl.) Hoch & Raven (denseflower willowherb) Eriophyllum lanatum (Pursh) Forbes (common woolly sunflower) Glyceria occidentalis (Piper) J.C. Nels. (northwestern mannagrass) Grindelia integrifolia DC. (Puget Sound gumweed) Hordeum brachyantherum Nevski (meadow barley) Juncus acuminatus Michx. (tapertip rush) Juncus ensifolius Wikstr. (swordleaf rush) Juncus tenuis Willd. (poverty rush) Microseris laciniata (Hook.) Schultz-Bip. (cutleaf silverpuffs) Plagiobothrys figuratus (Piper) I.M. Johnston ex M.E. Peck (fragrant popcornflower) Potentilla gracilis Dougl. ex Hook. (slender cinquefoil) Prunella vulgaris L. (common selfheal) Ranunculus orthorhynchus Hook. (straighbeak buttercup) Rumex salicifolius Weinm. (willow dock) Sidalcea cusickii Piper (Cusick’s checkermallow) Symphyotrichum hallii (Gray) Nesom (Hall's aster) Veronica scutellata L. (skullcap speedwell) Wyethia angustifolia (DC.) Nutt. (California compassplant) Zigadenus venenosus Wats. (meadow death camas)

Traits, Final Report Page 11 Wilson, Clark and Roberts Table 3. Overall species establishment rates (cover [%] per amount sown [g/cm2]) for those sown species that established somewhere in the six analysis sites. CV is the coefficient of variation (standard deviation ÷ mean). Frequency is the proportion of the 22 site × habitat combinations in which the species established. (Untransformed data.) Species Mean CV Freq. Species Mean CV Freq. Agrostis exarata 24.1 1.6 73% Juncus bolanderi 3.8 3.9 9% Beckmannia syzigachne 0.1 2.1 23% Juncus oxymeris 411.6 4.1 18% Carex ovalis 0.2 2.9 11% Juncus tenuis 1.8 4.1 23% Carex unilateralis 0.1 4.7 5% Lasthenia glaberrima 0.4 4.7 5% Castilleja tenuis 14.9 2.6 14% Lotus unifoliolatus var. 6.7 1.9 55% unifoliolatus Danthonia californica 2.1 3.4 15% Madia glomerata 19.9 2.2 41% Deschampsia caespitosa 3.1 1.4 59% Madia sativa 2.2 2.5 29% Dichanthelium 0.3 4.0 9% Microseris laciniata 0.1 2.2 23% acuminatum var. fasciculatum Downingia elegans 24.1 2.7 59% Navarretia intertexta 122.5 2.2 32% Downingia yina 0.4 4.4 5% Orthocarpus bracteosus 1.4 2.6 23% Eleocharis acicularis 6.2 4.1 6% Perideridia oregana 0.1 4.4 5% Eleocharis ovata 1.7 3.3 14% Plagiobothrys figuratus 19.9 4.5 55% Eleocharis palustris 1.0 3.4 13% Polygonum 0.6 2.6 14% hydropiperoides Epilobium ciliatum 0.7 2.0 25% Potentilla gracilis 0.0 4.7 5% Epilobium densiflorum 2.0 2.8 45% Prunella vulgaris 0.2 2.5 16% Eryngium petiolatum 4.0 2.9 32% Rorippa curvisiliqua 0.8 2.9 12% Glyceria occidentalis 0.6 2.1 23% Rumex salicifolius 0.5 4.1 6% Gnaphalium palustre 357.8 3.7 27% Sisyrinchium 0.4 4.2 6% angustifolium Gratiola ebracteata 76.2 2.2 27% Symphyotrichum hallii 0.5 4.7 5% Grindelia integrifolia 1.7 2.1 45% Veronica peregrina 0.6 4.1 6% Hordeum 0.7 3.7 13% Veronica scutellata 17.0 4.7 5% brachyantherum Juncus acuminatus 0.0 4.4 5%

Traits, Final Report Page 12 Wilson, Clark and Roberts Table 4. Establishment rates of species in the three habitats (cover [ %] per amount sown [g/cm2]). CV is coefficient of variation (std. dev. ÷ mean) and frequency is the proportion of the macroplots in which the species established. ns: not sown. (Untransformed data.) Vernal Pool Deschampsia dominated Emergent Mean CV Freq. Mean CV Freq. Mean CV Freq. Agrostis exarata 4.9 1.8 67% 49.3 1.0 100% 10.8 2.0 25% Beckmannia syzigachne 0.1 1.7 33% 0.1 2.2 22% -- -- 0% Carex ovalis -- -- 0% 0.3 2.6 14% 0.6 2.0 25% Carex unilateralis -- -- 0% -- -- 0% 0.4 2.0 25% Castilleja tenuis -- -- 0% 34.7 1.7 33% -- -- 0% Danthonia californica 0.1 2.8 13% 3.7 2.8 13% 2.7 2.0 25% Deschampsia caespitosa 0.4 1.5 44% 6.6 0.7 89% 1.1 2.0 25% Dichanthelium acuminatum -- -- 0% 0.8 2.5 22% -- -- 0% Downingia elegans 57.8 1.7 89% 1.1 2.6 44% 0.2 2.0 25% Downingia yina 1.0 2.8 13% -- -- 0% -- -- 0% Eleocharis acicularis -- -- 0% -- -- 0% 35.2 1.7 33% Eleocharis ovata 1.3 2.0 22% -- -- 0% 6.2 2.0 25% Eleocharis palustris 0.3 2.4 17% -- -- 0% 4.5 1.7 33% Epilobium ciliatum -- -- 0% 1.4 1.4 50% ns ns ns Epilobium densiflorum 3.0 2.9 33% 2.0 1.6 78% -- -- 0% Eryngium petiolatum 8.6 2.0 67% 1.1 3.0 11% -- -- 0% Glyceria occidentalis 0.8 2.0 22% 0.3 2.0 22% 0.6 2.0 25% Gnaphalium palustre 160.6 2.1 33% 19.5 2.2 22% 1562.5 2.0 25% Gratiola ebracteata 151.4 1.5 56% -- -- 0% 78.3 2.0 25% Grindelia integrifolia 3.0 1.7 44% 1.1 1.3 67% -- -- 0% Hordeum brachyantherum 0.1 2.4 17% 1.6 2.4 17% -- -- 0% Juncus acuminatus 0.1 2.8 13% -- -- 0% -- -- 0% Juncus bolanderi 1.8 3.0 11% 7.5 3.0 11% -- -- 0% Juncus oxymeris 902.1 3.0 22% 12.0 3.0 11% 206.9 2.0 25% Juncus tenuis -- -- 0% 4.3 2.6 56% -- -- 0% Lasthenia glaberrima 0.9 3.0 11% -- -- 0% -- -- 0% Lotus unifoliolatus 1.9 2.4 44% 14.5 1.2 89% -- -- 0% Madia glomerata 13.1 2.2 44% 35.5 1.7 56% -- -- 0% Madia sativa 0.2 2.6 14% 5.1 1.6 57% -- -- 0% Microseris laciniata 0.0 2.1 22% 0.1 1.7 33% -- -- 0% Navarretia intertexta 223.5 1.7 44% 75.9 1.7 33% -- -- 0% Orthocarpus bracteosus -- -- 0% 3.3 1.5 56% -- -- 0% Perideridia oregana 0.3 2.8 13% -- -- 0% -- -- 0% Plagiobothrys figuratus 48.0 2.9 78% 0.6 1.4 56% -- -- 0% Polygonum hydropiperoides -- -- 0% 1.4 1.7 33% -- -- 0% Potentilla gracilis -- -- 0% 0.0 3.0 11% -- -- 0% Prunella vulgaris -- -- 0% 0.5 1.5 38% -- -- 0% Rorippa curvisiliqua 0.8 2.6 14% 1.1 2.6 14% -- -- 0% Rumex salicifolius 1.2 2.6 14% -- -- 0% -- -- 0% Sisyrinchium angustifolium -- -- 0% 0.9 2.6 14% -- -- 0% Symphyotrichum hallii -- -- 0% 1.3 3.0 11% -- -- 0% Veronica peregrina 1.5 2.6 14% -- -- 0% -- -- 0% Veronica scutellata -- -- 0% -- -- 0% 93.4 2.0 25%

Traits, Final Report Page 13 Wilson, Clark and Roberts Table 5. Species that were sown into the six analysis sites but did not establish. Alisma gramineum Epilobium brachycarpum Montia linearis Alisma triviale Eriophyllum lanatum Myosotis laxa Allium amplectans Galium trifidum Perideridia gairdneri Asclepias speciosa Gentiana sceptrum Ranunculus alismaefolius Brodiaea coronaria Juncus effusus Ranunculus orthorhynchus Brodiaea hyacinthina Juncus ensifolius Saxifraga oregana Camassia leichtlinii Juncus nevadensis Scirpus validus Camassia quamash Juncus patens Sidalcea cusickii Cardamine penduliflora Leersia oryzoides Sisyrinchium californica Carex densa Lomatium nudicaule Sisyrinchium hitchcockii Carex feta Lotus formosissimus Sisyrinchium idahoense Carex lanuginosa Lotus pinnatus Sparganium emersum Carex obnupta Ludwigia palustris Typha latifolia Carex stipata Luzula campestris Wyethia angustifolia Cicendia quadrangularis Microsteris gracilis Zigadenus venenosus

Traits, Final Report Page 14 Wilson, Clark and Roberts Table 6. Ranked establishment rates (modified) of species used in the analysis. Ranks are based on the complete set of species and account for degrees of absence. Because the number of species differed in each data set, the rank for highest establishment varies. Also included are the ranked Seeding Assessment ratings, which are subjective assessments of dominance after seeding. Species All habitats Vernal Deschampsia- Emergent Seeding pool dominated wetland Assessment Agrostis exarata 45 41 48 44 28 Allium amplectens 0 0 0 0 2.5 Beckmannia syzigachne 11 25 20 20.5 28 Camassia leichtlinii 0 0 0 0 2.5 Camassia quamash 0 0 0 0 5.5 Carex densa 0 0 0 0 2.5 Carex unilateralis 9 13 12 37 18 Danthonia californica 33 24 38 41 9 Deschampsia cespitosa 35 29 41 40 28 Dichanthelium acuminatum 15 13 26 20.5 18 var. fasciculatum Downingia elegans 44 45 30 36 28 Downingia yina 18 33 12 20.5 18 Epilobium densiflorum 32 39 36 20.5 28 Eriophyllum lanatum 0 0 0 0 18 Glyceria occidentalis 21 31 23 39 18 Grindelia integrifolia 30 40 29 20.5 18 Hordeum brachyantherum 24 23 35 20.5 18 Juncus acuminatus 8 22 12 20.5 9 Juncus ensifolius 0 0 0 0 9 Juncus tenuis 31 13 39 20.5 18 Microseris laciniata 10 21 21 20.5 28 Plagiobothrys figuratus 42 44 25 20.5 28 Potentilla gracilis 7 13 19 20.5 18 Prunella vulgaris 13 13 24 20.5 18 Ranunculus orthorhynchus 0 0 0 0 18 Rumex salicifolius 19 34 12 20.5 18 Sidalcea cusickii 0 0 0 0 5.5 Symphyotrichum hallii 20 13 32 20.5 9 Veronica scutellata 41 13 12 47 18 Wyethia angustifolia 0 0 0 0 9 Zigadenus venenosus 0 0 0 0 2.5

Traits, Final Report Page 15 Wilson, Clark and Roberts Table 7. Growing conditions after the Integrated Standard Regime recommended by Hendry and Grime (1993) Light irradiance 125 ± 10 :mol m-2 s-1 equivalent to 29 W m-2 Red/far red ratio 1.4 Daylength 14 hr day, 10 hr night (the daily integral of PAR at 29 W m-2 over a 16 hr day is 1.46 MJ m-2, corresponding approximately to the daily mean of short-wave radiation receipt on a winter’s day at 53° N) Temperature 20-22C° day, 15-17C° night Humidity Not controlled Sand Water washed sand Watering Distilled water added to saucers of pots as required pH 4.5 ± 0.2 Nutrient addition 0.40 ml ± 0.05 ml nutrients per cm3 sand per week

Traits, Final Report Page 16 Wilson, Clark and Roberts Table 8. Favorable germination conditions for the test species, as reported in the literature. CS: cold stratification. WC: winter chill (ambient conditions for 6 wk, then alternating 8 hr at 5 C and 16 hr at 10 C). Species Pre-treatment Temper- Dark/light Reference ature hours Agrostic exarata None Allium amplectens CS: 90 days A. Barstow (pers. comm.) Beckmannia syzigachne WC 10/20 8/16 Guerrant and Raven (1995) Camassia leichtlinii CS: 5 C, 6 wk 10/20 8/16 Camassia quamash CS: 5 C, 6 wk 10/20 8/16 Guerrant and Raven (1995) Carex densa CS: 5 C, 6 wk 10/20 8/16 Guerrant and Raven (1995) Carex unilateralis CS: 5 C, 6 wk 10/20 8/16 Guerrant and Raven (1995) Danthonia californica CS: 5 C, 6 wk 10/20 8/16 Guerrant and Raven (1995) Deschampsia caespitosa CS or ambient conditions 10/20 8/16 Guerrant and Raven (1995) Dichanthelium None 19/33 12/12 Flynn et al. (2004) (based acuminatum var. on related spp.) fasciculatum Downingia elegans None 15 8/16 Flynn et al. (2004) (based on other Downingia spp.) Epilobium densiflorum None 21 12/12 Flynn et al. (2004) (based on other Epilobium spp.) Eriophyllum lanatum CS: 3-6 C, 6 wk 9-18 Winter Drake et al. (1998) None 20/20 photo- period 8/16 Glyceria occidentalis WC 10/20 8/16 Guerrant and Raven (1995) Hordeum CS: 5 C, 6 wk 10/20 8/16 Guerrant and Raven (1995) brachyantherum Juncus acuminatus None 21 12/12 Flynn et al. (2004) (based on other Juncus spp.) Juncus tenuis WC Microseris laciniata WC 10/20 8/16 Guerrant and Raven (1995) Plagiobothrys figuratus None 10 8/16 Flynn et al. (2004) Potentilla gracilis CS: 5 C, 6 wk 10/20 8/16 Guerrant and Raven (1995) Prunella vulgaris None 20/30 8/16 WC 10/20 8/16 Guerrant and Raven (1995) Ranunculus seeds subjected to ambient outdoor conditions Guerrant and Raven (1995) orthorhynchus Sidalcea cusickii CS: 5 C, 6 wk 10/20 8/16 Guerrant and Raven (1995) Symphyotrichum hallii None 15 8/16 Flynn et al. (2004) (data for A. chilensis)

Traits, Final Report Page 17 Wilson, Clark and Roberts Species Pre-treatment Temper- Dark/light Reference ature hours Veronica scutellata CS: 6 C, 4 wk 9/23 12/12 Flynn et al. (2004) Wyethia angustifolia seeds subjected to ambient outdoor conditions Guerrant and Raven (1995) Zigadenus venenosus CS: 3-6 C, 6 wk 9/18 Winter Drake et al. (1998) photo- period

Traits, Final Report Page 18 Wilson, Clark and Roberts Table 9a. Categorical traits for the analysis set of species from the West Eugene Wetlands Program. See Table 10 for an explanation of traits terms. Blanks: information not available. Species Growth form Clonal spread Dormancy breaking Life form Life form Perenniality Wetland (annual) mech. (expanded) Indicator Agrostis exarata grass below 5-25 cm/yr not stratification hemicryptophyte H mstem perennial FACW Allium amplectens monocot forb below < 5 cm/yr geophyte G bulb perennial FAC+* Beckmannia syzigachne grass non-clonal therophyte T basal facultative OBL perennial Camassia leichtlinii monocot forb below < 5 cm/yr geophyte G bulb perennial FACW- Camassia quamash monocot forb below < 5 cm/yr stratification geophyte G bulb perennial FACW Carex densa rush/sedge below < 5 cm/yr hemicryptophyte H mstem perennial OBL Carex unilateralis rush/sedge below < 5 cm/yr hemicryptophyte H mstem perennial FACW Danthonia californica grass below < 5 cm/yr not dormant hemicryptophyte H mstem perennial FAC- Deschampsia cespitosa grass non-clonal not stratification hemicryptophyte H mstem perennial FACW Dichanthelium acuminatum var. grass below < 5 cm/yr hemicryptophyte H mstem perennial FACW fasciculatum Downingia elegans dicot forb non-clonal therophyte T sstem annual OBL Downingia yina dicot forb non-clonal not stratification therophyte T sstem annual OBL Epilobium densiflorum dicot forb non-clonal not stratification therophyte T sstem annual FACW- Eriophyllum lanatum suffrutescent below 5-25 cm/yr stratification chamaephyte Ch suff perennial FACU+* Glyceria occidentalis grass below < 5 cm/yr hemicryptophyte H mstem perennial OBL Grindelia integrifolia dicot forb non-clonal hemicryptophyte H mstem perennial FACW Hordeum brachyantherum gras below < 5 cm/yr stratification hemicryptophyte H mstem perennial FACW- Juncus acuminatus rush/sedge below < 5 cm/yr not stratification hemicryptophyte H mstem perennial OBL Juncus ensifolius rush/sedge below < 5 cm/yr not stratification hemicryptophyte H mstem perennial FACW Juncus tenuis rush/sedge below < 5 cm/yr not dormant hemicryptophyte H mstem perennial FACW- Microseris laciniata dicot forb non-clonal not dormant hemicryptophyte H mstem perennial FACW-* Plagiobothrys figuratus dicot forb non-clonal not dormant therophyte T mstem annual FACW Potentilla gracilis suffrutescent non-clonal stratification hemicryptophyte H mstem perennial FAC Prunella vulgaris dicot forb below 5-25 cm/yr not stratification hemicryptophyte H sstem perennial FACU+ Ranunculus orthorhynchus dicot forb non-clonal hemicryptophyte H mstem perennial FACW Rumex salicifolius dicot forb non-clonal not dormant hemicryptophyte H mstem perennial FACW Sidalcea cusickii dicot forb below < 5 cm/yr not stratification hemicryptophyte H mstem perennial FACW-* Symphyotrichum hallii dicot forb below 5-25 cm/yr hemicryptophyte H mstem perennial FACW* Veronica scutellata dicot forb above > 25 cm/yr stratification hemicryptophyte H rept perennial OBL Wyethia angustifolia dicot forb non-clonal stratification hemicryptophyte H sstem perennial FACU Zigadenus venenosus monocot forb below < 5 cm/yr stratification geophyte G bulb perennial FACU Table 9b. Growth analysis traits for the analysis set of species from the West Eugene Wetlands Program. See Table 10 for an explanation of traits terms. Blanks: information not available. Species RGR Unit leaf rate Leaf mass Specific Root - Shoot mass Leaf area (seedling) (seedling) ratio leaf area shoot (seedling) (seedling) (1/day) (mg/cm2day) (seedling) (seedling) allometry (mg) (mm2) (cm2/mg) (seedling) Agrostis exarata 0.22 0.69 0.82 0.41 0.56 3.86 175.5 Allium amplectens Beckmannia syzigachne 0.27 0.92 0.83 0.38 0.72 7.29 321.2 Camassia leichtlinii 0.13 1.68 27.8 Camassia quamash 0.03 0.00 0.0 Carex densa 0.08 0.50 0.67 0.25 0.51 1.84 61.9 Carex unilateralis 0.08 0.60 0.59 0.26 0.20 1.00 31.3 Danthonia californica 0.13 0.62 0.71 0.30 0.68 4.57 151.9 Deschampsia cespitosa 0.14 0.85 0.69 0.24 0.57 2.96 94.2 Dichanthelium acuminatum var. fasciculatum Downingia elegans -0.01 -81 0.19 0.55 1.16 0.20 4.3 Downingia yina 0.06 1.15 0.27 5.3 Epilobium densiflorum 0.18 0.70 0.59 0.48 0.56 6.66 269.8 Eriophyllum lanatum 0.14 0.49 0.70 0.60 0.54 2.21 101.9 Glyceria occidentalis 0.23 0.90 0.72 0.36 0.65 23.45 690.3 Grindelia integrifolia 0.14 0.70 0.63 0.30 1.02 11.25 315.4 Hordeum brachyantherum 0.21 0.80 0.74 0.39 0.89 29.35 945.3 Juncus acuminatus 0.00 0.00 0.6 Juncus ensifolius 1.23 0.18 5.2 Juncus tenuis 0.62 0.13 6.7 Microseris laciniata 0.12 0.55 0.48 0.35 0.57 4.39 156.1 Plagiobothrys figuratus 0.22 0.76 0.82 0.42 0.42 5.57 180.3 Potentilla gracilis 0.12 0.31 Prunella vulgaris 0.15 0.60 0.65 0.44 0.59 6.48 306.5 Ranunculus orthorhynchus 0.17 0.63 0.48 0.58 0.77 8.48 382.6 Rumex salicifolius 0.26 0.66 0.67 0.56 0.62 23.73 1203.6 Sidalcea cusickii 0.12 0.50 0.66 0.41 0.73 16.60 535.5 Symphyotrichum hallii 0.20 1.05 0.55 0.33 0.53 4.13 206.4 Veronica scutellata 0.21 0.54 0.59 0.59 0.43 6.08 301.0 Wyethia angustifolia 0.07 0.65 0.66 0.21 1.85 13.68 340.3 Zigadenus venenosus 0.63 28.3

Traits, Final Report Page 20 Wilson, Clark and Roberts Table 9c. Other numerical traits for the analysis set of species from the West Eugene Wetlands Program. See Table 3 for an explanation of traits terms. Blanks: Information not available.

Species Wetland Flowering Laboratory Typical Seed mass Seed Indicator peak month germination maximum (mg) sphericity score rate (%) height (cm) Agrostis exarata 9 7 85 100 0.08 0.29 Allium amplectens 7* 5.5 68 50 0.91 0.03 Beckmannia syzigachne 11 6.5 72 100 0.59 0.16 Camassia leichtlinii 8 4.5 98 70 7.89 0.03 Camassia quamash 9 4.5 90 60 3.87 0.06 Carex densa 11 6.5 71 48 0.66 0.19 Carex unilateralis 9 6.5 24 100 0.40 0.22 Danthonia californica 3 6 46 80 4.22 0.25 Deschampsia cespitosa 9 7 77 100 0.34 0.23 Dichanthelium acuminatum 9 7 6 80 0.31 0.12 var. fasciculatum Downingia elegans 11 6.5 93 40 0.12 0.11 Downingia yina 11 5 96 30 0.05 0.08 Epilobium densiflorum 8 7.5 97 95 0.52 0.14 Eriophyllum lanatum 4 6 36 28 0.42 0.23 Glyceria occidentalis 11 7 91 140 2.00 0.20 Grindelia integrifolia 9 7.5 52 75 3.48 0.17 Hordeum brachyantherum 8 7 92 90 3.71 0.24 Juncus acuminatus 11 7 92 90 0.01 0.10 Juncus ensifolius 9 7 45 0.01 0.13 Juncus tenuis 8 7.5 40 60 0.02 0.09 Microseris laciniata 8* 5 76 75 1.37 0.26 Plagiobothrys figuratus 9 5.5 92 40 0.52 0.09 Potentilla gracilis 6 6.5 28 80 0.27 0.12 Prunella vulgaris 4 6 83 50 0.92 0.09 Ranunculus orthorhynchus 9 5 34 95 3.30 0.21 Rumex salicifolius 9 7 77 55 1.37 0.07 Sidalcea cusickii 8* 5.5 23 150 2.43 0.09 Symphyotrichum hallii 9* 8 57 50 0.24 0.24 Veronica scutellata 11 6 60 40 0.18 0.19 Wyethia angustifolia 3 5.5 50 90 11.19 0.20 Zigadenus venenosus 3 5 52 70 2.78 0.16 *: Assigned using local data.

Traits, Final Report Page 21 Wilson, Clark and Roberts Table 10. Explanation of specialized trait terms used in Table 9 and elsewhere. Clonal spread Below: belowground clonal growth organs; above: aboveground clonal growth organs Stratification A seed-dormancy-breaking treatment involving cold-moist conditions followed by warm conditions. Life form (expanded) Ch Chamaephyte, perennating buds above the soil but <25 cm G Geophyte, perennating buds buried within the soil H Hemicryptophyte, perennating buds at the soil surface T Therophyte, perennating structure is the seed (annuals) mstem Multiple leaf-bearing stems sstem One leaf-bearing stem basal All leaves growing basally (but not in a rosette) bulb Geophyte perennating from a bulb rept Creeping growth form Wetland indicator status terms for Region 9 (from US Fish and Wildlife Service 1988, 1993, 1996) OBL Occurs almost always (estimated probability 99%) under natural conditions in wetlands. FACW Usually occurs in wetlands (estimated probability 67%-99%), but occasionally found in non-wetlands. FAC Equally likely to occur in wetlands or non-wetlands (estimated probability 34%-66%). FACU Usually occurs in non-wetlands (estimated probability 67%-99%), but occasionally found on wetlands (estimated probability 1%-33%). +, – + indicates the wet side of an status term; – indicates the dry side of a status term. Growth analysis (all measured under standard, growth chamber conditions) RGR Relative growth rate: innate rate of increase in total dry mass per plant 7-21 days after germination Unit leaf rate Rate of dry mass production per leaf area over the period of 7-21 days after germination Leaf mass ratio Ratio of leaf dry mass to total dry mass, averaged over the period of 7-21 days after germination Specific leaf area Ratio of leaf area to leaf mass, averaged over the period of 7-21 days after germination Root-shoot Ratio of root relative growth rate to shoot relative growth rate allometry Shoot mass Dry shoot mass at 21 days after germination Leaf area Leaf area at 21 days after germination Wetland indicator score Wetland Indicator Status, converted to scale of 11 (OBL) to 1 (UPL) Flowering peak month Typical month of peak flowering; 6 = mid-June, 6.5 = June 30 Laboratory germination The proportion of seed that germinate under suitable conditions in the laboratory rate Seed sphericity Variance of seed length, width, and depth, normalized by the longest dimension

Traits, Final Report Page 22 Wilson, Clark and Roberts Table 11. Relationships between single explanatory variables and establishment rates (modified ranks). b: slope of the linear relationship; r2: proportion of variance in establishment rates among 2 2 species explained by that variable; r adj: r adjusted for degrees of freedom. Values in bold represent statistically significant relationships at " = 0.10. All habitats Vernal pool habitats Deschampsia- Emergent wetland dominated habitats habitats 2 2 2 2 2 2 2 2 b r r adj b r r adj b r r adj b r r adj Clonal spread 20% 11% 26% 18% 10% 0% 16% 7% Dormancy breaking 17% 7% 32% 24% 20% 11% 10% 0% mechanism Life form 31% 23% 49% 43% 28% 20% 29% 21% Life form (expanded) 49% 33% 55% 41% 32% 11% 43% 25% Growth form 36% 26% 38% 28% 45% 37% 41% 32% Perenniality 20% 14% 35% 30% 5% -2% 2% -5% Wetland Indicator 12% -10% 18% -3% 10% -13% 16% -4% Status Wetland Indicator 1.70 7% 4% 2.33 14% 11% 0.66 1% -2% 1.82 9% 6% score Flowering peak month 7.36 21% 18% 6.89 18% 15% 9.88 39% 36% 7.17 20% 17% Laboratory germination 0.17 9% 6% 0.27 22% 20% 0.10 3% 0% 0.09 2% -1% rate Maximum height -0.03 0% -3% 0.03 0% -3% 0.08 2% -1% 0.08 2% -1% Leaf mass ratio -2.42 0% -5% 1.36 0% -5% 11.50 1% -2% -0.52 0% -5% Seed mass -1.98 11% 7% -2.00 10% 7% -1.90 10% 7% -2.28 14% 11% Seed sphericity 52.95 6% 3% 28.39 2% -2% 79.94 14% 11% 86.18 16% 14% Relative growth rate 62.53 8% 4% 62.57 8% 4% 69.50 11% 7% 59.07 7% 3% Unit leaf rate -7.10 3% -1% -6.23 2% -2% -0.19 0% -4% -5.35 2% -2% Specific leaf area 20.47 2% -2% 9.92 1% -4% -21.81 3% -2% -4.86 0% -5% Root:shoot allometry -6.27 2% -3% -3.03 0% -5% -6.07 2% -3% -14.63 10% 5% Aboveground mass 0.11 0% -4% 0.36 3% -1% 0.19 1% -3% 0.13 0% -3% (21 d) Leaf area (21 day) 0.00 1% -3% 0.01 4% 0% 0.00 1% -3% 0.00 1% -3%

Traits, Final Report Page 23 Wilson, Clark and Roberts Table 12. Results of stepwise regression of species traits in explaining species establishment rates in the West Eugene Wetlands Program restoration sites. The response variable was the modified rank of the average establishment rates. (See text for explanation.) Factors were added only if they contributed significant explanatory power, using a

Mallows Cp procedure. Factor d.f. bF P All habitats (R2 = 50%) Flowering peak month 1 6.6 8.0 <0.01 Perenniality 2 7.2 <0.01 Seed sphericity 1 61.1 3.6 0.07 Vernal pool habitats (R2 = 70%) Laboratory germination rate 1 0.2 7.9 <0.01 Life form 3 5.7 <0.01 Flowering peak month 1 5.3 5.7 0.03 Deschampsia-dominated habitats (R2 = 62%) Flowering peak month 1 8.5 10.7 <0.01 Growth form 4 3.8 0.02 Emergent habitats (R2 = 52%) Growth form 4 4.9 <0.01 Seed mass (sqrt) 1 -7.3 5.5 0.03

Traits, Final Report Page 24 Wilson, Clark and Roberts Table 13. Categorical variables with significant influence on establishment rates in the final, multivariate models. Means that do not share a letter differ statistically.

All habitats Establishment rate (modified rank) by Perenniality Facultative Perennial Annnual perennial Count 1 26 4 Mean 11.0 13.9a 34.0b SE – 2.9 5.9

Vernal pool habitats Establishment rate (modified rank) by Life form Geophytes Chamaephytes Hemi- Therophytes cryptophytes Count 4 1 21 5 Mean 0.0a 0.0 17.0ab 37.2b SE 0.0 – 2.9 3.7

Deschampsia-dominated habitats Establishment rate (modified rank) by Growth form Rush/sedge Suffrutescent Monocot forb Dicot forb Grass Count 5 2 4 13 7 Mean1 12.6a 9.5ab 0.0ab 17.9ab 33.0b SE 7.1 9.5 0 3.5 3.9

Emergent wetland habitats Establishment rate (modified rank) by Growth form Monocot forb Suffrutescent Rush/sedge Dicot forb Grass Count 4 2 5 13 7 Mean 0.0a 10.25ab 15.6ab 19.0ab 32.2b SE 0.0 10.25 7.0 3.7 4.2

1The apparently disparity between means and the statistical comparison arises because of small sample sizes and high variance.

Traits, Final Report Page 25 Wilson, Clark and Roberts Table 14. Results of stepwise regression of species traits in explaining Seeding Assessment ratings in the West Eugene Wetlands restoration sites. The response variable was the rank of the ratings, which are subjective assessments of dominance after seeding. (See text for explanation.) Factors were added only if they contributed significant explanatory power, using a Mallows Cp procedure. Categories within the trait of Clonal spread that do not share a letter differ statistically. Factor d.f. mean bFP R2 = 66% Clonal spread 3 11.5 <0.01 below, <5 cm/yr 9.9a below, 5-25 cm/yr 18.3ab above, >25 cm/yr 18.0ab non-clonal 22.4b Seed mass 1 -1.49 12.4 <0.01 Maximum height 1 0.08 5.1 0.03

Traits, Final Report Page 26 Wilson, Clark and Roberts

All habitats

perennial annual annual perennial 40

perennial perennial annual perennial 30 perennial

perennial

perennial perennial 20 perennial annual

perennial perennial facultative perennial perennial 10 perennial Establishmentrank) rate (modified perennial perennial

0 perennialperennial perennial perennial perennial perennial perennial

4.5 5.5 6.5 7.5 Flowering peak month

Figure 2. Relationship between establishment rate and Flowering peak month for All habitats, with Perenniality shown. For Flowering peak month, 6 = mid-June.

Traits, Final Report Page 28 Wilson, Clark and Roberts Vernal pool habitats

therophyte hemicryptophytetherophyte 40 hemicryptophyte therophyte hemicryptophyte

therophyte 30 hemicryptophyte hemicryptophyte

hemicryptophyte therophyte hemicryptophyte hemicryptophyte hemicryptophyte 20

hemicryptophyte hemicryptophyte hemicryptophytehemicryptophyte hemicryptophyte hemicryptophyte hemicryptophyte 10 Establishmentrank) rate (modified

hemicryptophytehemicryptophytechamaephyte geophyte geophyte hemicryptophyte 0 geophyte hemicryptophyte geophyte

10 30 50 70 90 Laboratory germination rate (%)

Figure 3. Relationship between establishment rate and Laboratory germination rate in Vernal pool habitats, with Life form shown.

Traits, Final Report Page 29 Wilson, Clark and Roberts Emergent wetland habitats

50 dicot forb

grass grass grass grass 40 rush/sedge

dicot forb

30

rush/sedge dicot forb dicot forb dicot forb grass rush/sedged t forb dicot forbdicot forb 20 dicot forb dicot forb grass suffrutescentgrass

Establishmentrank) rate (modified 10

monocot forbmonocot forb suffrutescent monocot forb 0 rush/sedge dicot forb rush/sedge dicot forb dicot forb monocot forb

0.0 0.5 1.0 1.5 2.0 2.5 3.0 3.5 Seed mass (sqrt mg)

Figure 4. Relationship of establishment rate and seed mass (square-root transformed), with growth form shown.

Traits, Final Report Page 30 Wilson, Clark and Roberts