A HACKELIA TAYLORI (BORAGINACEAE), NEW A FROM NORTH CENTRAL WASHINGTON STATE SPECIES (U.S.A.) Richy Harrod Malmquist* Lauri A. J. USDA USDA Forest Service Forest Service Okanogan-Wenatchee National Forest Wenatchee River Ranger District 21 5 Melody Lane 600Sherbourne Wenatchee, Washington 98801, U.S Leavenworth, Washington 98826, U.SA. . [email protected] [email protected] Robert Carr L. Department ofBiology Eastern Washington University Cheney, Washington 99004, U.S.A. ABSTRACT RESUMEN Hackelia Opiz. includes 45 species of perennial plants distributed within Northern Temperate region, Central and South America combined (Mabberley 1987). In North America, 28 species are recognized, comprising 34 & many taxa, of which are narrow endemics (Gentry Carr 1976). Species of Hackelia can be found in a wide range of habitats, including sagebrush steppe, steep talus slopes, moist rock crevices, open deciduous forests, many and Abies or Pinos forests; and the distribution of species are narrowly restricted based on habitat, geog- & raphy, or elevation (Carr 1974; Gentry Carr 1976). Gentry and Carr (1976) completed a comprehensive study of Hackelia and taxonomic relationships of the species and subspecies in North America. However, the clarified (Gamon taxonomic status of H. venusta (Piper) St. John has remained in question 1988) and recently has been the focus of taxonomic research (Harrod et al. 1999; Hipkins et al. 2003). Hackelia venusta, as originally described, includes a white-flowered form found at one low elevation site km (488 m) 9.6 northwest of Leavenworth, Washington, and a blue-flowered form found at four currently km known high elevation 2050 m) alpine locations about 18 northwest and southwest of Leavenworth, (ca. & Washington (Carr 1974; Gentry Carr 1976; Hitchcock et al. 1959) (Fig. 1). The two forms were shown to be distinct from each other based on morphological traits (Harrod et al. 1999). However, enzyme band pattern analyses did not provide evidence for taxonomic separation of the two color forms (Hipkins et 2003). Al- al. though at first this may seem to be a dilemma, taxonomy is often the result of synthesis across many lines of evidence, some of which may to support the taxonomic hypothesis (Grant 1992; Winston 1999; Hipkins fail et 650 Journal of the Botanical Research Institute of Texas 7(2) Taxonomic 2003). al. entities are defined on a combination of morphology, genotypic data isozymes), (e.g., ecology, reproductive and isolation, geographic distribution. There are compelling reasons to consider the two ™ flower color forms be to separate species based on ™ morphological and ecological distinctions He dC £ Pr 3 techniCal descri " »*** Ptio for the form at the rank of named species, here Hackelia , Although taylori. H. was shown tayl,ori be to morphologically from distinct H. venusta in our previous work va.ie.KS of H. (Doug iffusa ex Lehmann) Johnston which „e did no. study previously, but which were in- eluded in the enzyme work of Hipkins et al. (2003). MATERIALS AND METHODS Study Sites m 7 ° fr 10 Ukti°nS US£d by Harr°d WUh H 31 Ct (1999) 2 ° additi nal Populations ^ V3r COtt(mii (Piper) Carr on located Private land in tthhee RRaattttlleessnnaakkee HilL fRfn km WA m ’ Hills (RH), 25 north of Sunnyside, 150 and elevation; H. 2) diffusa var diffusa lo- ca e at Oneota Gorge (CXI) near Multnomah km 50 OR m Falls, east of Portland, 75 elevation. Morphological Measurements and Analyses Statistical We followed the same methods and used most of the same data (individual plants from original 10 populations with missing morphological data were dropped from analysis) as described in Harrod et (1999). Nineteen al. morphological characters from three categories (vegetative, floral, and fruit) were scored for statistical analysis and an additional 11 descriptive characters leaf shape, leaf surface, color) were recorded (Table These (e.g., 1). data were collected from 25 randomly selected individuals from each population except the Cashmere Moun- population, which consisted of data from 14 individuals, and Crystal Cirque with only 10 individuals. tain Both principal components and discriminant analyses were performed on the quantitative morphological data using SPSS 16.0 (SPSS, Chicago, Illinois). Principal components analysis (PCA) was used to evaluate the natu- groupings among each sampling unit or operational taxonomic unit (population). Discriminant analysis ral was used to establish the non-arbitrariness of group assignments. This analysis places each case (plant) within the group (population) with which shares discriminating characters (Anderson and Taylor 1983). The analy- it biased in that positions cases within the ordination based on discriminating characters to achieve sis is it A maximum separation of the defined groups. plot of the cases based on the two discriminating functions first can assist in visualizing distinction among groups and species. The data for these analyses involved a 237 * 19 morpho- Descriptive characters were not subjected to statistical analyses but were used to further detail new logical characteristics of the taxon. The addition of H. diffusa var. cottonii and H. diffusa var. diffusa in both the principal components and discrimi- nant analyses lead to tighter groupings of cases as compared to the results of Harrod et al. (1999). Of the 19 com- ponents that accounted for all the variance in the PCA, the first three accounted for 65.1% (32.5%, 20.7%, and Characters highly correlated with the component were lower cauline leaf width and 11.9%, respectively). first and length, upper cauline leaf width, and radical leaf width (Table 2). Hackelia diffusa var. cottonii H. diffusa var. component separated from other taxa along the (Fig. 2A). Plant height, radical leaf petiole length, diffusa first component and upper cauline length were characters highly correlated with the second radical leaf length, leaf CM) (Table Along this second component, H. taylori (CC and separated from the H. diffusa var. arida popu- 2). lations forming a distinct group that was somewhat overlapping with the H. venusta population (TC) (Fig. 2A). The third component separated H. venusta from H. taylori and characters highly correlated with the third com- ponent were upper cauline leaf length, limb width, fomice protuberance, and calyx length (Table Fig. 2B). 2, The discriminant analysis showed H. taylori was more clearly distinct from H. venusta and almost the en- tire H. diffusa complex (Fig. 3). The first two discriminant functions accounted for 81.8% of the ability to dis- tinguish among groups (51.2% and 30.6%, respectively). Total predictability that a case from a certain popula- tion was correctly classified to that population was 95.8%. Individual cases (plants) had high predicted group CM membership with the sampled population from which they were sampled. Hackelia taylori plants from CC 100% and from were 95.8% with 4.2% with were those correctly classified, classifying correctly classified = new Harrod et aL, A species of Hackelia from north central Washington 653 Factor 2 Fig. 2. A. Ordination of populations of Hackelia examined in this study based on scores of principal components 1 and 2. The first two components accounted for 53.2% of the total variance, 32.5% and 20.7%, respectively. Hackelia taylori populations (CC and CM) are highlighted with heavy black line. B. Ordination of populations of Hackelia venusta and H. taylori examined in this study based on scores of principal components 2 and 3. The third = BM TW component accounted for an additional 1 1 .9% of the variance for a total of 65.1% for the first three. H. diffusa var. arida Burch Mountain, : = = MC = = = Tumwater Canyon, SC Swakane Canyon, PE Ponderosa Estates, Moses Coulee, DE Derby Canyon, DC Douglas Creek. H. diffusa var. OG diffusa-. = = = CM = = Oneota Gorge. H. diffusa var. cottonir. RH Rattlesnake Hills. H. venusta: TC Tumwater Canyon. H. taylori: Cashmere Mountain, CC Crystal Creek. Journal of the Botanical Research Institute of Texas 7(2) Score 1 = = Tumwater CM Canyon. H. taylori: Cashmere Mountain, CC Crystal Creek. dm many Moderately short perennial, 1-2 tall; stems often from a slender taproot, erect or spreading from branched caudex. Leaves ciliate, antrorsely appressed strigose (Fig. 4A); radical leaves 3.7-10.4 cm long, 0.8- cm 2.9 wide, appressed linear-elliptical, strigose, petiolate for 1/3 to 1/2 their length; cauline leaves linear to cm linear-lanceolate, sessile; lowermost cauline leaves 0.6-3.1 long, 0.4-1.2 cm wide; upper cauline leaves cm cm 1 .9-5.3 long, 0.5-1.3 wide, reducing upward to the inflorescence, becoming small 2.3- bracts. Pedicel mm mm and mm, 5.1 in flower (Fig. 3B) 5.0-7.0 in fruit. Calyx length 2.4-3.4 linear-lanceolate, strigose. Co- mm cm rolla limb blue, 1.0-1.7 wide with five lobes, lobes 3.0-5.0 long 4D, Fomices with appendages (Fig. F). mm showy, white, sometimes tinged pink, slightly emarginate, papillate-pubescent, rising 0.8-1.0 above the mm mm protuberances throat; yellow, pandurate, 0.6-1.0 long 4D, Anthers (Fig. E). 0.8-1.0 long. Nutlets mm 1.8-3.6 long, lanceolate-ovate; dorsal surface verrucose-hispidulous, intramarginal mar- prickles 7-13; mm ginal prickles connate for up to 1/2 their length, forming a flange 1.2-2.4 wide around the nutlet; distinct mm, prickle length 0.7-1.4 a long prickle alternating with one or two shorter ones (Fig. 4C). .— “ Etymology The epithet ” honors Ronald who taylori Dr. Taylor, taught botany Western Washington at J. University, Bellingham, Washington. Dr. Taylor co-authored the first status report for H. venusta in 1979 and was actively involved in native plant conservation in the Pacific Northwest for nearly 40 years. . Harrod et at, A new species of Hackelia from north central Washington 657 taylori flowers are always blue and H. venusta flowers are white but sometimes tinged blue suggesting perhaps some they share genes for color. new Like H. venusta, this species would benefit from well-developed conservation strategies. Populations are at risk from loss due to stochastic events, such as rock slides, which were the cause of the loss of most of one known population of H. taylori. Conservation strategies might include long-term seed banking so that popula- tions could be re-established in the event of a stochastic loss. ACKNOWLEDGMENTS We would like to thank John Gamon, Loyal Mehrhoff, and Kali Robson for their work early-on suggesting a new species description. Dottie Knecht, Mark Cedar Drake, Ellen Kuhlmann, and Shelly Benson pro- Ellis, We Pam Camp vided field assistance. Brandy Reed assisted with data analysis. thank for help in locating popu- We BLM lations of H. diffusa var. arida on land. thank Guy Nesom for providing the Latin description. This USDA project was cooperatively funded by the Forest Service, USDI Fish and Wildlife Service, and the Wash- ington Natural Heritage Program. Figures and 2 were developed by Dan O’Connor, Okanogan-Wenatchee 1 We National Forest. Illustrations of H. taylori in Figure 3 were drawn by Eve Ponder. also thank two anony- mous and reviewers for detailed constructive reviews. REFERENCES Anderson, A.V. and RJ. Taylor. 1983. Patterns of morphological variation in a population of mixed species of Castilleja (Scrophulariaceae). Syst. Bot. 8:225-232. Carr, R.L 1974. A taxonomic study in the genus Hackelia in western North America. Ph.D. Dissertation, Oregon State Gamon, 988. Report on the status of Hackelia Heritage Program, Olym- vt J. 1 il Washington. pia, A and 1976. of the genus Hackelia (Borac Gentry, R.L. Carr. revisit n Jr., J.L. Mem. New York Bot. Gard. 26:1 21-227. Grant, 1992. Systematics and phylogeny of the Ipomopsis aggregata group (Polemoniaceae): traditional and molecu- V. lar approaches. Syst. Bot. 1 7:683-691 A Harrod, RJ., LA. Malmquist, and R.L Carr. 1999. review of the taxonomic status of Hackelia venusta (Boraginaceae). Rhodora 101(905):16-27. Wilson, RJ. Harrod, and C. Aubry. 2003. Isozyme variation in showy stickseed, a Washington endemic Hipkins, V.D., B.L. plant, and relatives. N.W. Sci. 77:1 70-1 77. Hitchcock, A. Cronquist, M. Ownbey, and J.W. Thompson. 959. Vascular plants of the Pacific Northwest Part 4: Ericaceae C.L., 1 Washington WA. through Campanulaceae. University of Press, Seattle, The Cambridge Boston MA. Mabberley, DJ. 1987. plant-book. Press, New Columbia Winston 1999. Describing species. University Press, York. J.E.