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Hawkweeds in the Northwestern United States

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Hawkweeds in the Northwestern United States 18 RANGELANDS 19(4), August 1997 Hawkweeds in the Northwestern United States Linda M. Wilson, Joseph P. McCaftrey, Paul C. Quimby, Jr., and Jennifer L. Birdsall everal species of hawkweed (Hieracium spp.) are re- being genetic forms of a "related" species (Hitchcock et al. cent additions to the weedy flora of the Inland and 1955) S Pacific Northwest, the region encompassing The genus is divided into 4 subgenera, 3 of which are Washington, northern Idaho and northwestern Montana. found in north-temperateflora and are distinguished by the Over 60,000 acres are currently estimated to be infested. following key (adaptedfrom LePage 1971): Private landownerconcern about the rapid rate of spread of 1a. South Americanflora Mandonia hawkweeds is shared land wildlife by public managers, lb. North temperateflora managers, timbercompanies, livestock farmers, and recre- 2a. Plants normally stoloniferous (except. ationists. Lack of control of these weeds has become a se- H. Leaves rious land piloselloides Vill.): always managementconcern in the last decade. Various basal; involucreformed of two or three chemical and cultural methods have successfully controlled series of imbricatedbracts small Piosella infestations, but larger, less accessible populations, 2b. Plants not stoloniferous: comprising the bulk of the infestation, are more difficult to 3a. Involucreformed of more than two control. series Biological control is being investigated as a tool of bracts Hieracium with which to unequal length combat the increasing hawkweed problem. 3b. Involucre formed of two series of The of this is to summarize purpose paper whatis currently unequal length bracts Stenotheca known about the introduced hawkweeds in the United There are considered to be about 36 of States in order to facilitate future research and manage- generally species ment of hawkweeds. Hieracium belonging to 3 subgenera in America north of Mexico. About 25 species are native to the United States and Canada and fall into 2 Taxonomy subgenera, Hieracium (3 species) and Stenotheca (22 species) (Table 1). Subgenus The hawkweeds (Hieracium spp.) are members of the Hieracium is circumboreal in distribution, and is represent- ed in North tribe Lactuceae (= Cichorieae) in the family Asteraceae. America by H. umbellatum L., H. robinsonii The name Hieracium is derived from the Greek hierax, Zahn and H. canadense Michx. Species in this subgenus meaning hawk; allegedly keen-sighted hawks of yore ate reproduce sexually and apomictically. Subgenus the sap of the brightly colored plants to sharpen their Stenotheca is restricted to the New World and contains eye- most of our native sight (Fernald 1950). Hieracium is an extremely diverse taxa. Species in this subgenus are strict- genus morphologically.It includes a huge number of taxa in ly sexual. Conflicting characters used to distinguish species which apomixis, hybridization, and are all com- have generated considerable debate regarding classifica- polyploidy tion. For mon (Stebbins 1950). Apomixis, or asexual reproduction, example, Guppy (1978) suggestedthat H. scouleri includes both agamospermy (production of seeds without Hook., H. albertinum Farr., and H. cynoglossoides L. are too pollen) and vegetative reproduction. These reproductive closely correlated in characters to be distinguished into and strategies generate enormous morphological and cytologi- separate species, instead form a complex of species. cal variation within and among species, making Hieracium Similarly, Guppy (1978) suggested that H. umbellatum L., one of the most taxonomically difficult genera. Problems and its close relative H. canadense Michx., may be 2 forms of the same with taxonomy are worsened by the lack of clearly defined, species. Deardorff (1977) added H. longiberbe stable characters that are important in classifying species Howell and H. nudicale Heller to the H. scouleri complex. The hawkweed within the genus. For example, species are broadly distin- remaining species found in the United guished by the character and amount of leaf States and Canada are introduced from Europe and belong pubescence, to 2 venation, flower color, stolon morphology, and the size, subgenera (Table 2). Four species belong to the sub- Hieracium: H. murorum H. shape, and color of the inflorescence (Bailey and Bailey genus L., argillaceum Jordan 1976, Voss and Bählke 1978). The result has been the de- group, H. sabaudum L., and H. groenlandicum Arv.-Touv. Plants in this scription of a huge number of species, subspecies, vari- subgenus have not become weedy in North America. The eties, and forms that differ slightly (Sell 1974). Zahn (1921- remaining 7 species belong to the subgenus 22) conducted the most comprehensive treatment of the Pilosella.As in other subgenera,these plants are difficult to genus in which he described 756 species world-wide. It is distinguish morphologically. From a practical standpoint, in estimated that while over 10,000 Hieracium species, sub- however, plants subgenus Pilosella fall into 3 groups. species, varieties and forms have been described world- The first group is represented by H. pilosella, mouse-ear wide, only about 700 species are valid, with the remainder hawkweed, easily distinguished by its single, yellow flower. RANGELANDS 19(4), August1997 19 Table 1. List of Hieracium native to America north of species Species Subgenus Mexico,by subgenus (H=Hleraclum,S= Stenothoca). = H. idahoense Gand. H. nudicaleHeller Species Subgenus = H. paddoense Gand. H. canadense Michx. (Canada h.) H = H. washingtonense Gand. = H. columbianum Rydb. = H. cascadorum Zahn. = H. umbellatum var. scathisculum Farw. = H. chelannense Zahn. = H. laevigatum ssp. canadense Zahn. = H. chapacanum Zahn. H. robinsonji (Zahn) Fern (Robinson h.) H = H. scoulen var. scouleri = H. scoulen var. nudicale H. umbellatum L. (narrow-leaved h.) H (Gray) = H. scabrisculum Schwein. H. traillii(Green's h.) S = H. canadense var. scabrum = H. greenei Porter & Britton (? Gray) = H. macranthum Nutt. = H. howellhiGray. = H. canadense var. angustifolium 1. & G. = H. bathigerum Greene = H. kalmii = H. oregonicum Zahn. = H. suksdorfiiGand. = H. cinereum Howell. H. albiflorum Hook. (whitef lowered h.) S H. triste Wilid.(alpine h.) S = H. vancouvenanum Arv.-Touv. H. venosum L. (rattlesnake weed) S = H. hefleriGand. = H. leptopodanthum Gand. = H. candelabrum Gand. While predominantly an eastern weed, H. piosella has a = H. albiflorum subvar. rosendahlii Zahn. limited western distribution in coastal Washingtonand = H. albiflonimf. lyalliiZahn. Oregon (Roche 1992). The second group includes H. au- H. albertinum Farr (western h.) s rantiacum L., orange hawkweed, the only introduced = H. absonumMacbr. species with orange flowers. The remaining group consists = H. scoulenvar. albertinum of species with multiple, yellow flowers and includes H. H. argutum Nutt.(S. Californiah.) s pratense Tausch. (= H. caespitosumDumort), H. praealtum H. bolanden Gray. (Bolander'sh.) Vill., H. piloseioides (= H. florentinum All.), H. floribundum = H. siskiyouense Peck. Wimm. & Grab., and H. flagellare Wilid. (= H. auricula L.). H. cameum Greene s Known variously as yellow hawkweed,meadow hawkweed, field devil and devil H. cynoglossoides L. (houndstongue h.) hawkweed, king hawkweed, yellow = H. griseumRydb. hawkweed, this group represents most of the taxonomic = H. cusickilGand. complexities. = H. flettliSt. John & Warren Until recently, it was thought that only H. pratense (Figure = H. panyiZahn. 1 occurred in the Inland Northwest, however, H. flonbun- = ) H. iydbeipiiZahn. dum and H. pioselloides have also been identified in the = H. scoulen var. gnseum (Rydberg) past 6 years. Other populationscontain plants that are mor- H. fendleriSchultz (Fendler h.) S phologically similar to H. pratense and H. pilose/loides. H. gracile Hook. (slender h.) S Lepage (1967) suggested that H. pioseioides is a hybrid = H. hookenSteud. between H. praealtum and H. flagel/are. Similarly, he treat- = H. tnstevar. detonsum ed H. flage/lareas a hybrid between H. pratense and H. p1- H. gronoviiL. (hairy h.) S lose/Ia (Lepage 1967, 1971). Guppy (1976) suggestedthat H. homdum Fries. (shaggy h.) 5 H. floribundumand H. pratense, distinguishableonly by the = H. brewenGray degree of pubescence, may be different phases of a single H. lemmoniiGray species that exhibits a broad morphologicalrange. Lepage H. longiberbe Howell (long-beaked h.) S (1967) recognized the largest number of hybrids among the = H. piperi St. John &Warren introduced hawkweeds, particularly between H. auranti- = H. scoulen var. longiberbe acum, H. pilose/la, H. floribundum and H. pratense. It is H. longipilum Tori. (long-bearded h.) 5 likely that the combined influences of morphologicalvaria- H. mananum Willd. (Maryland h.) tion, polyploidy, frequent apomixis, and widespread hy- dridization the introduced has led to a H. paniculatum L. (panicled h.) among species huge of intermediate and that distinctions H. array types, species pringlelGray. (woolly h.) can no longer be made in North America based solely on H. rusbyiGreene 5 descriptionsfrom Europeanflora. H. scabrum Michx. (rough h.) S H. scouleri Hook. (Scouler's h.) s Biogeography = Pilosella scoulen Schu. & Schu.-Bip. = H. amplumGreene H. was introduced into the United States in = pratense likely H. absonum Macbride & Payson 1828 (Britton and Brown 1970), and is now commonly = H. cinereum Howell found from Québec and Ontario (Frankton and Mulligan 20 RANGELANDS 19(4), August1997 Table 2. List of Hieracium species Introduced into America
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