The Scarlet-Flowered Species of Echinocereus in Utah

Home , Echinocereus, Echinocereus arizonicus

Sego Lily July 2010 33 (4)

July 2010 (volume 33 number 4)

In this issue:

The Scarlet-Flowered Species of Echinocereus in Utah ...... 1, 4 Chapter News ...... 2 Annual Field Trip Highlights . . . 3 Bulletin Board ...... 3 New San Juan/Four Corners Native Plant Society Global Garlic Mustard Survey Cedar Breaks Wildflowers A Comparison of Utah Echino- cereus Species ...... 7 Dodder Doesn‟t Dodder Around 8 Utah Botanica: Zion Vegetation Program: In Bloom Year-Round ...... 9 Botanist‟s Bookshelf: Summer Reading Special ...... 10

Cover: A small clump of Claret cup (now called Echinocereus mojavensis) in the San Rafael Swell. Photo by Dorde W. Woodruff

The Scarlet-Flowered Species of Echinocereus in Utah

By Dorde W. Woodruff fairly common sight in Utah‟s can- have not been studied extensively or yons, foothills, plateaus, and the thoroughly enough. The scarlet-flowered higher parts of desert valleys. The Unfortunately Utah is a backwa- clumps commonly grow in rocky ter for cactus research, so members Echinocerei are predomi- of the cactus family are less well un- places and usually on slopes. derstood here than in better- nantly hummingbird-pollinated and Botanists have often sought to researched states such as Arizona. have large, showy, bright red flowers identify cactus specimens from To my knowledge we have never had in cespitose clumps that can reach Utah on the basis of keys derived a professor at one of our Utah col- an impressive size of a hundred or from other states that don‟t work leges or universities interested in more heads. Sizeable clumps with well here or are not valid. Many Cactaceae, with a stable of eager their bright flowers are a striking cactus populations in our state graduate students to do the work. feature of late spring. They are a In this some- [continued on page 4] Copyright 2010 Utah Native Plant Society. All Rights Reserved. Utah Native Plant Society

Education: Ty Harrison Sego Lily Editor: Walter Fertig Horticulture: Maggie Wolf ([email protected]). The deadline for Invasive Weeds: Susan Fitts the September 2010 Sego Lily is 15 Rare Plants: Walter Fertig August 2010. Scholarship: Bill Gray Copyright 2010 Utah Native Plant So- Chapters and Chapter Presidents ciety. All Rights Reserved Cache: Michael Piep Cedar City: Marguerite Smith The Sego Lily is a publication of the Officers Escalante: Harriet Priska Utah Native Plant Society, a 501(c)(3) President: Walter Fertig (Kane Co) Fremont: Lisa White not-for-profit organization dedicated Vice President: Kipp Lee (Salt Lake Co) Manzanita: Walter Fertig to conserving and promoting steward- Treasurer: Charlene Homan (Salt Lake Mountain: Mindy Wheeler ship of our native plants. Use of con- Co) Salt Lake: Marni Ambrose tent material is encouraged but re- Secretary: Mindy Wheeler (Summit Southwestern/Bearclaw poppy: Mar- quires permission (except where ex- Co) garet Malm empted by statute) and must be cor- Board Co-Chairs: Bill King (Salt Lake Utah Valley: Celeste Kennard rectly credited and cited. Articles, Co) and Dave Wallace (Cache Co) photographs and illustrations submit- Website: For late-breaking news, the ted to us remain the property of the UNPS Board: Loreen Allphin (Utah UNPS store, the Sego Lily archives, submitting individuals or organiza- Co), Robert Fitts (Utah Co), Susan Fitts Chapter events, links to other websites tions. Submit permission requests to (Utah Co), Ty Harrison (Salt Lake Co), (including sources of native plants and [email protected]. We encourage read- Celeste Kennard (Utah Co), Margaret the digital Utah Rare Plant Field ers to submit articles for potential Malm (Washington Co), Larry Meyer Guide), and more, go to unps.org. publication. By submitting an article, (Salt Lake Co), Therese Meyer (Salt Many thanks to Xmission for an implicit license is granted to print Lake Co), Leila Shultz (Cache Co), sponsoring our website. the article in the newsletter or other Maggie Wolf (Salt Lake Co). For more information on UNPS: UNPS publications for reprint without Contact Bill King (582-0432) or Susan permission (in print and electronic Committees Fitts (801-756-6177), or write to media). When submitting an article, Communications: Larry Meyer UNPS, PO Box 520041, Salt Lake City, please indicate whether it has been Conservation: Bill King and Tony UT, 84152-0041 or email previously published or submitted for Frates [email protected] consideration to other publications.

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yon Botanical Area on Dixie Na- Chapter News tional Forest, off UT Hwy 12 west of Bryce Canyon National Park. We Manzanita (Kane County): On will carpool from the Grand Stair- May 28, a dozen Kanab area plant case-Escalante NM visitor center enthusiasts embarked on an expedi- parking lot in Kanab at 8 AM and tion to the East Rim of Zion Canyon plan to arrive at the botanical area to view the spring wildflowers of by 9:30 AM for a leisurely half day Zion National Park. Part of our mis- of searching for Claron endemics sion was also to confirm the pres- among the Bristlecone pines. For ence of Dwarf cryptanth (Crypt- more info, contact me antha humilis, below), a species ([email protected]).—W. Fertig. which had been previously reported for Zion NP, but not photographed Salt Lake: There are three remain- or documented with a museum ing field trips scheduled for summer voucher. We successfully located a 2010: small patch of the low-growing, Saturday, July 17: Upper Lambs white-flowered, bristly plants along the trail on Carmel Limestone. Several other plants of interest Canyon with Bill Stockdale and were found with the cryptantha, Mindy Wheeler including the yellow-flowered Saturday, August 7: Brighton to Charleston Mountain violet (Viola see Wood nymph (Moneses purpurea var. charlestonensis) uniflora) with Bill Nelsen and Rose‟s spring-parsley Saturday, August 14: Hiking and (Cymopterus purpureus var. ro- potluck in Upper City Creek Canyon sei). In all, we found 132 vascular with Marni Ambrose. plant species in bloom. For more information on these Our next field trip will be on trips, contact Bill Gray at cyber- Saturday, 17 July to the Red Can- [email protected] or 801-532- 3486.—Bill Gray 2 Sego Lily July 2010 33 (4) Annual Field Trip Bulletin Board Highlights New San Juan/Four Corners Native Plant Society: Many of you On June 12, the Manzanita Chap- may already be aware of the „Southwest Colorado Wildflowers‟ website cre- ter hosted the UNPS state board for ated by Al Schneider (www.swcoloradowildflowers.com/) - it is a real treas- its yearly foray to southern Utah. As ure trove of information and photos about the plants of the Four Corners part of the festivities, the chapter area. Al has now started a new society dedicated to the native plants of the sponsored the first botanical foray of area called the San Juan/Four Corners Native Plant Society. He invites all the Best Friends Animal Sanctuary, UNPS members and especially those from the southeast part of Utah to par- located 7 miles north of Kanab. Best ticipate in his field trips and other events. Follow the link above to get more Friends is a no-kill animal shelter information or contact Al directly if you would like to be included in email focusing on dogs, cats, horses, rab- notification. - Bill Gray bits, and exotic birds. The grounds of the sanctuary include several Global Garlic Mustard Field Survey: Do you want to be part of the hundred acres of unspoiled redrock world‟s largest scientific research project on invasive species? The „Global canyons, pinyon-juniper forest, sand Garlic Mustard Field Survey‟ is an international collaboration to obtain dunes, and riparian woodlands. Our much-needed data on the abundance and distribution of Garlic mustard goal was to develop a plant species (Alliaria petiolata) across its native and introduced ranges. In our first list for the sanctuary. field season last year, we received measurements and seed samples from 65 Our small but enthusiastic team populations, with a majority from Europe. Our goal for this summer is 150 of botanists and naturalists spent 6 or more, with a stronger emphasis on the southern and mid-west to western hours scouring the area in search of United States. new plant species. Entomologist This year we are hoping to increase participation among educators, as Ken Kingsley (below) provided ex- well as land managers and citizen scientists who may not have much formal pert commentary on a number of science training. The survey involves a simple protocol that can be followed unusual insects in the area, includ- directly or incorporated into field courses and nature surveys. A population ing the yucca moth and the parasi- takes two people about 2-4 hours to measure. We are also planning to de- toid wasp that preys on its develop- velop internet-based teaching modules and tools to aid with monitoring and managing this invasive plant. The sampling protocol, along with contact information is available at the Global Garlic Mustard Field Survey website: www.GarlicMustard.org. Ideal sampling time is 2-4 weeks after flowering finishes (mid to late July in Utah). Please contact me if you would like to participate—Dr. Robert Colautti, Biology Dept., Duke University ([email protected]).

Cedar Breaks Wildflower Festival: The 5th annual Cedar Breaks Wild- flower Festival will be held from Friday, July 2 to Sunday, July 18 at Cedar Breaks National Monument in scenic southeastern Iron County, Utah (23 miles east of Cedar City). Volunteer wildflower specialists will be on hand for guided hikes at 10 AM and 1 PM daily. A Junior Ranger “wildflower scaven- ger hunt” will be held on Fridays and Saturdays dur- ing larvae within the flower of the ing the festival at 3 PM. All Narrow-leaved yucca (Yucca angus- activities begin at the Fee tissima). Booth area (adjacent to the Prior to the foray, I had collected main parking area, about 1.5 or observed 179 vascular plant spe- miles inside the monument‟s cies in the Best Friends area. By south entrance). The visitor day‟s end our group had observed center will also be hosting 129 plant species, of which 36 were an ongoing electronic dis- new to the sanctuary. Our day‟s play of wildflower images work increased the flora of the area and offer free wildflower by nearly 17%, bringing the total photography tip sheets, and discounts on wildflower-themed books. For number to 215 species. more information on the festival and associated activities, go to the Cedar After the board meeting, our Breaks National Monument website, www.nps.gov/cebr or call 435-586- group reconvened at the home of 0787. There is an entrance fee of $4 for Cedar Breaks National Monument Best Friends co-founder Jana de for persons 16 and older and visitors in July should be prepared for poten- Peyer for a potluck dinner and gen- tial afternoon showers and temperatures dropping to the 60s. eral merriment.—Walter Fertig 3 Utah Native Plant Society The Scarlet-Flowered Species of Echinocereus in Utah (continued from page 1) gloomy picture the large collection In 1985 Nigel Paul Taylor‟s of cactus specimens at the Brigham monograph “The Genus Echinocer- Young University (BYU) is a great eus” was published. Taylor is a bota- asset. nist specializing in the Cactaceae at Many botanists don‟t like to deal the famous Royal Botanic Gardens with cacti and they are underrepre- at Kew, England. This book covers sented in herbaria. Because of their the whole genus (not just those of succulence and also their armor, the U. S. and Canada), and includes making decent dried specimens out three species in Sect. Triglochidia- of them is difficult, and the dried tus: E. scheeri, a complex Mexican specimens are more changed in species, E. triglochidiatus, and E. form from their live state than are polyacanthus (which is now consid- those of non-succulent plants. The ered a synonym of E. coccineus). study of live plants in their environ- The treatment of Echinocereus ment is more important in this fam- for the 2003 volume of Flora of ily. Their morphology is different, North America (FNA) was written and botanists who don‟t deal with by Allan Zimmerman and Bruce them much don‟t have an eye for Parfitt, both one-time students of them. Consequently, the study of Donald Pinkava‟s at Arizona State cacti lags behind that of most other University in Phoenix. They in- plant families, though more ad- cluded three U.S. species in Sect. vanced in the southern states where Triglochidiatus: E. coccineus, E. there is more academic interest than triglochidiatus and E. arizonicus. in Utah. Although recognizing E. coccineus, These red-flowered hedgehog they did not include Utah in its dis- cacti have been placed in a sub- tribution. genus, section Triglochidiatus. Go- Like Benson, Pinkava is one of ing through the literature on Echi- the Grand Old Men of cactus re- nocereus shows that this section has search; unlike Benson, he is still not been at all well understood. with us and still active. His special- Various authorities have subdivided ties are counting chromosomes and the basal species E. triglochidiatus studying polyploidy, and also the into a confusing array of species or genus Opuntia. Now retired from varieties. Above: Hedgehog in the city: A clump Arizona State University, he is still Quoting the late Lyman Benson, of Echinocereus mojavensis perches consulting with botanists of the De- the largest single figure in cactus high above Salt Lake City in quartzite. sert Botanical Garden in Phoenix research, “Twenty-nine years of this Photo by Tony Frates. and others. The work of Pinkava and [research] has not produced the ulti- his students in studying chromo- mate answer” to resolve “…the insta- Most of Benson‟s work was fin- some numbers has been important bility of the populations of the pro- ished well in advance of publica- in sorting out Echinocereus. posed species and their extensive tion of his 1982 major work The Last fall, in one of my periodic and bewildering intergradation with Cacti of the United States and tours of herbaria, I visited the each other.” In recent years most Canada. Not only must so compre- Garrett Herbarium at the University experts have used Benson‟s classifi- hensive a work necessarily be done of Utah (U of U) and found that in cation for this section, in which he over a period of years, but also be- preparation for his treatment of the reduced the various scarlet-flowered cause it is a large and expensive genus for Intermountain Flora species to varieties of E. triglochi- volume, the Stanford University (IMF), Marc A. Baker (also once a diatus. He did produce some order Press had to wait several years af- student of Pinkava‟s) had visited the for these plants in his varieties, ter completion of the manuscript three major Utah herbaria: Utah which others have built on. It was until it had accumulated funds to State University and BYU, in addi- not a perfect analysis, but was the print it, so in spite of Benson‟s long tion to the U of U, and annotated the best available. I have followed Ben- and patient work on the cacti, it specimens of Echinocereus. Speci- son in determining all the scarlet- was outdated even when it first was mens formerly determined as E. flowered hedgehogs in Utah as E. published. Nevertheless, it was the triglochidiatus or one of its varieties triglochidiatus but without the va- most comprehensive work on the were annotated by Baker mostly as rieties, which didn‟t seem to work family for a long time. E. mojavensis and a few as E. well in Utah. 4 Sego Lily July 2010 33 (4) coccineus. Before this E. mojavensis was usually recognized as a variety of E. triglochidiatus, and in previous years E. coccineus was not gen- erally recognized as occurring in Utah. Preparation for major works like the various volumes of the continu- ing series of IMF and FNA offers a great opportunity in making results of research accessible, as the experts who were chosen to write up the genera review data and visit herbaria. Scientific papers on the cacti don‟t always appear in the most ob- vious place, the Cactus and Succu- lent Journal of the Cactus and Suc- culent Society of America, or its more technical offspring Haseltonia but are scattered in many journals. The Internet makes it easier, but even so, it takes a formidable amount of time spent searching to keep up with current botanical literature, even in this one family. coccineus, E. mojavensis, E. san- Above: Claret cup (Echinocereus coc- The forthcoming volume of IMF taritensis, E. triglochidiatus, and cineus) near the Petrified Forest Trail in that includes the Cactaceae will E. yavapaiensis. And he recog- Zion National Park. Photo by Tony bring us a much better understand- nized some Utah specimens as E Frates. ing of the species of section coccineus. Our limited population Triglochidiatus. So how did this of E. coccineus, as far as is known, new understanding occur? Baker, is only in the eastern part of Wash- cineus in IMF, while placing E. can- now an environmental consultant in ington County and mostly in and yonensis in synonomy. It is possible Arizona, has been working on Echi- near Zion. The Utah plants are that further study will change this. nocereus for years. Baker deter- different than typical E. coccineus. The theory of its difference is that mines the boundaries of taxa on the Ours are part of the group named these plants grow in areas that are basis of morphology, geographic by Michigan botanists Elzada Clo- hotter and drier, where humming- distribution, polyploidy, and floral ver and Lois Jotter (professor and birds are not abundant, and there- dimorphism (in some species male student), the first women to run fore are becoming adapted to bees and female flowers are produced on the entire Colorado River, as E. rather than hummers. Flowers separate plants). For morphology, canyonensis, with the type speci- adapted to bees are smaller than he measures sizes in a set of 22 men collected down in the Grand hummingbird flowers. Bees don‟t characters of flowers and stems, and Canyon at Hermit Falls. need as much nectar as the larger analyzes them using advanced sta- They are characterized by birds with their high metabolisms, tistical techniques: analysis of vari- smaller flowers, by some (but not and often seek pollen rather than ance (ANOVA), discriminant analy- all) plants displaying one of a se- nectar. Though bees visit the scarlet- sis (DA), multivariate analysis of ries of flower colors trending to- colored hummingbird flowers, they variance (MANOVA), principle com- wards yellow and away from the prefer yellow flowers. The predomi- ponent analysis (PCA), and un- scarlet color of the hummingbird nant pigments in the Cactaceae are weighted pair group method with pollination syndrome, and by be- betalains, the chemistry of which arithmetic mean (UPGMA). He also ing visited by hordes of bees rather precludes anthocyanins with their has extensive field experience and than hummers as the predominant blue hues, which bees favor. The has counted many a chromosome pollinators. Our plants are north- scarlet hues of the hummingbird set. As good botanists do, he studies ern outliers, with plants like ours hedgehogs come from flavonoids, type specimens and the occurrence on the Arizona Strip connecting to and these flavonoids are different of the plant at the type locality to the type locality in the Grand Can- pigments not found in most cacti. understand what the author in- yon. The recognition that E. coccineus tended, and does not recognize taxa Because there is some intergra- is found in Utah solved a longtime until he has enough data to do so. dation, and because this group is puzzle for me. In May of 1973 I came So what does Baker recognize? U. not sufficiently well known, Baker upon a curious population of hedge- S. species recognized by him in sect. includes these plants with E. coc- hog cacti on the Kolob road. These Triglochidiatus are E. arizonicus, E. 5 Utah Native Plant Society plants had a somewhat different open and chunky look, and has on color of flower in orangish hues, average shorter stems, fewer ribs, moving away from the usual pure areoles farther apart, and shorter scarlet, and were attracting swarms and fewer spines. of bees. I made specimens of it but E. coccineus in Utah may be dis- did not know what to call it, putting tinguished not only by spine struc- it under the umbrella of E. triglochi- ture, but also by its location in diatus but stating that it was differ- Washington County (as far as is ent and being visited by bees. known). E. mojavensis also occurs At that time, the best and most there, but in more mesic locations. If complete reference for keying out flowering, E. coccineus will display Utah cacti was Benson‟s 1969 3rd smaller flowers with perhaps differ- edition of The Cacti of Arizona; I ent colors, and is predominantly also referred to his other 1969 book, visited by bees rather than hum- The Native Cacti of California. Both mingbirds. Each plant will have only of these carried E. coccineus and E. male or only female flowers. Stem canyonensis in synonomy (along and spine measurements overlap with many other synonyms) but with E. mojavensis but the ones I‟ve with no descriptions. Another refer- seen seem to be taller and more ence was Boissevain and Davidson‟s slender with finer spines on better- 1940 Colorado Cacti, which did in- separated areoles. This is tentative, however, and spine surface and clude E. coccineus. But their concept Above: True Echinocereus triglochi- of the species was not the tetraploid, diatus from the Sandia Mountains in flower characters are more useful dimorphic entity as it is understood New Mexico. Photo by Bob Sivinski. until we know this species in Utah today, on the basis of the type, and better. So far I‟ve accumulated about their circumscription of it did not fit 11 locations from various sources, the environment of my specimen. group, a western group repre- most in or near the south end of This is not to dismiss their work; sented by the type and an eastern Zion, but also extending from Pah they did well for that time when so group, perhaps represented by the Tempe Hot Springs near Hurricane much less was known about the Cac- basionym Cereus mojavensis to the Little Creek Mountains. taceae. Engelm. & J. M. Bigelow var. According to specimens in the Baker‟s annotation of E. mo- zuniensis J. M. Bigelow & Engelm, major Utah herbaria, E. mojavensis javensis for all the other Utah speci- for which the type locality is in is found in all Utah counties except mens of sect. Triglochidiatus was Canyon Diablo, east of Flagstaff, the far northern tier of Box Elder, more puzzling. For one thing, speci- Arizona.” To resolve this, an Weber, Cache, Rich, Morgan, Davis, mens in our East Desert are some- analysis such as that employed by and Summit. what different from those in our Baker needs to be done. Baker has annotated the speci- West Desert. For another, because As to the second issue, is E. mo- mens in our three major herbaria, of all the wildly different treatments javensis the only sect. Triglochi- but most of the smaller Utah her- of this section in cactus literature diatus taxon in Utah other than E. baria have probably not yet caught though the years, until now, E. mo- coccineus? Baker‟s herbarium vis- up with this recent work. javensis, which is based on a speci- its showed that although E. men from the Mojave River in Cali- triglochidiatus as now understood References: fornia, had not been clearly and con- occurs near the Four Corners, it sistently differentiated from E. has not appeared in herbaria as Baker, M.A. 2006 A new florally dimor- triglochidiatus, based on a speci- occurring in Utah. To best differ- phic hexaploid, Echinocereus yavapaiensis sp. nov. (section Triglochidia- men from Wolf Creek in New Mex- entiate between these three species ico. tus, Cactaceae) from central Arizona in the field or in herbarium speci- Plant Systematics and Evolution 258: Addressing the first issue are two mens, Baker uses spine characters. 63-83 statements, the first from FNA: Viewed under 30x magnification, “That taxon [mojavensis] includes E. coccineus has spines round in Baker, M.A. 2006 Circumscription of curly-spined plants (mainly in Cali- cross section and mostly smooth- Echinocereus arizonicus subsp. arizoni- fornia) and straight-spined plants surfaced with little trend toward a cus: Phenetic analysis of morphological (including most populations in Ari- papillate surface. E. mojavensis characters in section Triglochidiatus zona, Utah, and western Colorado).” also has round spines but they are (Cactaceae), Part II Madroño 53(4): 388-399 And from Baker, “…UPGMA sug- papillate. And E. triglochidiatus gests that E. triglochidiatus subsp. has spines that are smooth but mojavensis [this was written before angled in cross section, fewer, and he had enough data to feel satisfied stouter. In the field E. triglochi- in recognizing this taxon in specific diatus is the most noticeably dif- rank] may be composed of more ferent vegetatively, with a more than one taxonomically definable 6 Sego Lily July 2010 33 (4)

A Comparison of Utah Echinocereus Species

By Tony Frates and Dorde Woodruff

Species Synonyms and notes Location in Utah Description (none limited to UT) Echinocereus A confusing array of synonyms Only known from eastern Stems in closely set clumps; coccineus have been used; this taxon was Washington County flowers dimorphic, with a not formerly well understood slightly constricted “waist”; Scarlet claret cup, Scarlet 7-19 smooth spines per ar- hedgehog Utah plants are part of the pro- eole, uniform in color posed E. canyonensis; more though gray in age, round in study is needed for acceptance cross-section; flowers The specific epithet coc- of this as a valid taxon smaller, narrower, < 5 cm cineus=scarlet is some- long, may be more yellowish what a misnomer for than in E. mojavensis ours since they are much Visited by bees more than visited by bees and thus hummingbirds drifting from the scarlet hummingbird-attracting Tetraploid, n=22 color Echinocereus Vars. chrysocentrus, variegatus, A southern Utah, Great Stems single to several in engelmannii & purpureus have been recog- Basin, and Colorado Pla- open clumps but not mound nized in the past, but differences teau species in Beaver, forming; central spines with Engelmann's hedgehog are inconsistent or insufficiently Garfield, Iron, Kane, contrasting light and dark documented and vars are no Juab, Millard, Tooele, colors, flat or angled; flow- longer accepted San Juan, and Washing- ers perfect, pale to dark ton cos. rose-pink, to 9 cm tall and wide

Bee pollinated; easily distinguished from other Utah Echinocereus

Tetraploid, n=22 Echinocereus Most often known as E. The most widespread Plants in closely set clumps; mojavensis triglochidiatus var. mojavensis, Echinocereus in Utah, flowers perfect, scarlet, usu- with a confusing number of occurring in all counties ally >5 cm long; 5-11 spines Claret cup; Hedgehog; other synonyms. Sometimes except the northern tier per areole, uniform in color Hummingbird hedgehog misspelled as “mohavensis”. The of Box Elder, Cache, though gray in age, mostly specific epithet is unfortunately Davis, Morgan, Rich, and round in cross-section, a misnomer since it is not re- Summit cos. papillate (use 30x lens) stricted to the Mohave Desert, but named for the Mojave River. Hummingbird pollinated. There may be differences between western and eastern Diploid, n=11 plants.

The name var. inermis has been applied to the spineless form of far east-central Utah, but this taxon is no longer accepted.

7 Utah Native Plant Society

Dodder Doesn’t Dodder Around

By Peter Lesica (Adapted from Kelseya, the newsletter of the Montana Native Plant Society)

Dodders (Cuscuta spp.) are surely among the world‟s most unusual plants. They are parasitic on other flowering plants and lack leaves or any photosynthetic tis- sues. This is odd enough, but unlike most other parasitic plants, such as broomrapes (Orobanche spp.), paintbrushes (Castilleja spp.) or even mistletoes (Arceuthobium spp.), dodders have no roots. They can be annu- als arising from seed each year or perennials arising from overwin- tering stem segments. Our species have yellow or orange, twining stems. Dodders are closely related to morning glories, but the flowers are inconspicuous though sometimes with intricately ornamented corollas. There are about 150 species of dodder worldwide, most Above: 1796 print of Cuscuta europaea dodder develops two different exten- by the painter Jacob Sturm from sions from its haustoria: a hand-like common in subtropical and tropi- Deutschlands Flora in Abbildungen. cal America. Eleven species are form that surrounds the phloem and a straw-like form that pierces the reported for Utah, and based on the small number of herbarium choose a host is also reminiscent of xylem. In this way dodder can ob- collections and my own experi- an insect herbivore or parasite. tain everything it needs (water, min- ence, most (with the exception of Researchers have demonstrated eral nutrients and food) from its C. pentagona) are not common in that dodder stems will orient to- host. the state. In Utah they have often ward a tomato plant or even a vial Most species of dodder can para- been collected parasitizing native of tomato extract as long as an sitize many different host plants, but and introduced legumes as well as odor can be detected. Other stud- studies have shown that dodder spotted knapweed and other ies have shown that dodder can grows better on some hosts com- members of the Aster Family. locate a host by the quality of light pared to others. Furthermore sev- In some ways, dodder acts reflected off the host‟s leaves and eral studies have shown that dodder more like an animal than a plant! will even preferentially move to- is able to preferentially infect those Other parasitic plants, such as ward hosts with higher chlorophyll hosts that provide the most benefit. broomrape, have seeds that germi- content, in other words, those that Colleen Kelly, from Oxford Univer- nate only when they are contacted are greener, and presumably with sity, demonstrated this and found by host root exudates, but not more sugars. that the choice is made based on dodder. Dodder seeds germinate As soon as a dodder stem twines chemicals in the bark that dodder on the surface of the ground and around its host it begins to form can detect before forming haustoria. then forage for their host. The haustoria, specialized short stems Kelly also found that having two stems grow outward while at the that tap into the host vascular tis- different hosts was better than one, same time waving around until sue. Flowering plants have two although a second independent they reach a host plant. The juve- kinds of vascular tissue. Xylem study failed to confirm this result. nile stems can reach up to 6 cm carries water and mineral nutri- The effects of dodder go beyond (2.4 inches), but none are able to ents from the roots up the stem, just individual host plants. By the reach twice that far, and they must and phloem moves energy-rich end of a growing season a single find a host within a few days or die carbohydrates to the roots or other dodder plant may form thousands of trying. The way dodder is able to areas of need. To match this, haustorial connections with many different host species and cover an 8 Sego Lily July 2010 33 (4) area the size of a small house. Of course this can have significant ef- Utah Botanica fects on the plant community. Since Odds and Ends from Utah Botany they are somewhat host-specific, dodders can alter community struc- Zion Vegetation Program: In Bloom Year-Round! ture by preferentially damaging some species more than others. For In 2007, Zion NP received funding to put in seed increase fields at its Na- example, University of Montana‟s tive Plant Nursery, with the goal of growing and harvesting large amounts of Ray Callaway and his collaborators seed and using that seed for large-scale restoration projects such as fire re- found that dodder reduced the habilitation, annual brome competition, and campground revegetation. dominance of glasswort (Salicornia) Part of this funding included the purchase of a new seed storage cooler, in favor of sea-lavender (Limonium) which would be used to store seed from Zion and Cedar Breaks and Pipe in California coastal marshes. Dod- Spring National Monuments at a consistent temperature year-round. The ders also may damage commercial Zion Veg Program has long needed a dependable unit for long-term seed crops such as tomato, pumpkin and storage, and we got it in the form of a 10x12 walk-in refrigerator with light- alfalfa. Indeed, this is why we know ing, a floor drain, and adjustable temperatures that can be set as low as 35°F so much about dodder ecology. (below right). Dodders‟ negative impacts also ex- tend beyond simple parasitism. For example, they can be conduits between host plants for viruses, including disease-causing pathogens. Some diseases can spread more quickly through a crop field infested with dodder than one without. On the positive side, a native Chi- nese dodder has been used to con- trol bittervine (Mikania) a serious invasive weed in China as well as Puerto Rico. The native dodder causes a decline in the invader re- sulting in greater nutrient availabil- ity to native members of the community. Although it might seem like host plants are defenseless against the wiley dodder, this may not always be To celebrate the new cooler and do away with unnecessary “white space,” the case. Recently researchers have the Veg Program hired fellow plant nerd Shannon Eberhard (below left), an found that some host plants transfer aspiring plant illustrator and a former employee of the Fire Effects Monitor- messenger RNA (mRNA) into their ing Team, to paint the cooler with native plant illustrations (above). Shan- dodder parasites. Some of these non did an amazing job, and that means that the Veg employees are lucky mRNAs can incapacitate dodder‟s enough to look at beautiful, accurate native plant paintings on a daily basis. genetic machinery, thereby reducing Right awn!! -Rebecca Lieberg, Zion Lead Reveg Bio Tech its ability to make proteins and grow. This discovery has spawned an interest in genetically engineer- ing crop plants that produce dodder- destroying mRNA. With luck agri- culturalists may be able to turn dodder‟s voracious appetite against it.

References: Koch, A. M., C. Binder, & I. R. Sand- ers. 2004. Does the generalist parasitic plant Cuscuta campestris selectively forage in heterogeneous plant communi- ties? New Phytologist 162: 147-155. Runyon, J. B., M. C. Mescher & C. M. DeMoraes. 2006. Volatile chemical cues guide host location and host selection by parasitic plants. Science 313: 1964-67.

9 Utah Native Plant Society Botanist’s Bookshelf: Summer Reading Special

By Walter Fertig and Dan Norris, 2009, Micro-Optics Press, 430 pp. One of my annual Ahh summer … time to unroll the New Year‟s resolutions is to become hammock, pour a cool drink, and more competent identifying mosses relax with a good botany book … or and bryophytes. California Mosses two. The following are some recent may just be the book I‟ve been wait- titles you might consider for your ing for to help me. The authors have summertime reading pleasure. amassed an amazing array of color Wasatch Wildflowers: A photos of 600 moss species of Cali- Field Guide, by Steve Hegji, 2010, fornia that illustrate growth form, Cedar Fort, Inc, Springville, UT. leaf shape, and important cellular 207 pp. While this slim guide does details. Many of the photos are not cover every plant species in Above: Oenothera howardii by quite beautiful and their subjects northern Utah‟s Wasatch Range, it Steve Hegji. resemble pieces of stained glass art. does include 200 of the most com- Accompanying the brief descriptions mon or showy wildflowers. Each for identifying anything but the are black-and-white drawings of color photo is accompanied by a most showy species, but is an ex- stylized leaves that emphasize diag- brief, often entertaining caption. cellent introduction to the overall nostic differences between genera For example, American bistort diversity of genera and species and species. A table at the end of (Polygonum bistortoides) is de- found in North America. It would the book compares and contrasts scribed as being “… a bit like a giant be an excellent companion to a these leaf shapes to help the profes- Q-tip sticking up above all the other more formal taxonomic textbook sional or novice bryologist place plants.” A series of thumbnail sym- in teaching students about floristic their unknown specimen into its bols is used to organize brief discus- diversity. proper sequence. Though the book sions of key flower, leaf, and habitat The Sibley Guide to Trees, covers California specifically, my traits and other information. Hegji by David Allen Sibley, 2009, Al- bryologist buddy John Spence as- is a gifted photographer with a real fred A. Knopf, New York. 426 pp. sures me that most of the species eye for composition and interesting Over the past decade Sibley has from Utah and western North Amer- perspectives. His book is geared to a created a small publishing empire ica are included. non-technical audience, but the based on his series of bird guides, Flora of North America: vol- photos will be appreciated by all but now has branched into trees of umes 7 and 8, by the Flora of plant and nature lovers. North America. Done in much the North America Editorial Committee, National Wildlife Federa- same style as his original bird field 2009-2010. The Flora of North tion Field Guide to Wildflow- guide, The Sibley Guide to Trees America (FNA) project was con- ers of North America, by David covers nearly 600 native and natu- ceived nearly 20 years ago and 16 of Brandenburg, 2010, Sterling Pub- ralized tree species with watercolor the planned 28 volumes have now lishing, New York. 673 pp. Com- paintings of leaves, flowers, fruits, been published. The ambitious goal pact but hefty, this field guide has twigs, bark, and other identifying of the project is to produce a taxo- more than 4000 color photos of characteristics. The book has brief nomic treatment of the entire flora 2200 native and introduced wild- descriptions of each species and a of the North American continent, flowers found across North America. detailed range map, as well as es- including bryophytes, ferns, gymno- The book is organized taxonomically says on families and genera. Un- sperms, and flowering plants. Vol- rather than by flower color and like many recent tree guides, this umes include keys to all native and shape as in many other popular field book is organized phylogenetically naturalized species, range maps, guides. Descriptions of species are and not by leaf arrangement brief taxonomic descriptions, and necessarily brief, and only a few of (opposite vs. alternate) or shape. often a line drawing (at least one per the more common or showy species I noticed some minor errors, such genus). The two most recent vol- in each genus are featured, along as the range map of Utah juniper umes cover 30 families of flowering with color range maps. An innova- omitting most of the Colorado Pla- plants, including the Saxifragaceae, tive feature is the use of thumbnail teau, but no more than one might Primulaceae, Ericaceae, Salicaceae, images of different flower types ar- expect from an ornithologist writ- and Brassicaceae. ranged by color and shape located at ing about botany. The book is Because it covers all of North the beginning of the book which probably too large for field use, but America, the taxonomic keys in the helps orient the user to the correct would be a valuable reference for FNA are often difficult as they must family or genus without resorting to keying out specimens at home or focus on obscure characters to split difficult keys. This book covers too for general information. out so many taxa, or species tend to wide of a geographic area and too California Mosses, by Bill come out in the key at multiple few species to be especially useful and Nancy Malcolm, Jim Shevock, points. Those interested in keying 10 Sego Lily July 2010 33 (4) out their local species are better Major Taxonomic Changes limestone fellfields of Beartooth served by good state or regional flo- in FNA volumes 7 & 8 Butte in Wyoming (but not in Mon- ras which only have to deal with a tana as reported). Likewise, Draba subset of all the taxa in the FNA. Brassicaceae (Cruciferae) calcifuga is a cryptic but distinct The real value of the FNA comes in Arabis: split into three genera - species from Montana and Wyoming its monographic treatment of fami- Arabis (sensu stricto), Boechera, that can be readily distinguished lies and genera. The taxonomic con- & Turritis from D. oligosperma but has been cepts and nomenclature used in the Cardaria: lumped with Lepidium synonymized without comment. FNA should guide the treatment of Cusickiella: new genus (formerly The value of FNA would increase species in state and regional floras included in Draba) greatly if more emphasis was placed in the future. Lesquerella: lumped with Phy- on discussions of taxonomic prob- Numerous taxonomic changes saria lems associated with species (like have been made in volumes 7 and 8, Schoenocrambe: lumped with the Drabas mentioned above). In especially in the Brassicaceae, Cap- Sisymbrium or moved to new the interest of brevity, such discus- paraceae (now Cleomaceae), and Genus Hesperidanthus sions are frequently excluded or among genera formerly included in Sinapis: formerly included in kept too short to be meaningful. the Primulaceae, Pyrolaceae, and Brassica This is unfortunate, because without Monotropaceae. Such changes can adequate explanation taxonomic be upsetting to those of us more Cleomaceae (formerly decisions can appear arbitrary or comfortable with the names we Capparaceae) weakly supported. I would also like learned in our youth, but are not a Peritoma: new genus for Cleome to see the new volumes return to the bad thing if they represent advances lutea and C. serrulata. original practice of having range in taxonomic concepts (taxonomy is maps depict the approximate area a science after all, and not just Crassulaceae inhabited by a species, and not just stamp collecting). The accompany- Rhodiola: formerly included in have one dot per state. This is re- ing table summarizes the more sig- Sedum dundant anyway, given that states nificant changes that affect species, Tillaea: lumped with Crassula and provinces are listed in the text genus, and family concepts in Utah. for each species. The best maps in Many of the changes seem rea- Ericaceae the FNA series were those done for sonable. A lot of morphological and Family expanded to include the the two grass volumes which had genetic data support merging the Monotropaceae and Pyrolaceae. county level distribution. There is mustard genera Lesquerella and Ledum transferred to Rhododen- no good excuse for excluding such Physaria. Likewise, no one will lose dron information as county-level distri- sleep over combining Cardaria with bution maps are now readily avail- Myrsinaceae Lepidium. Other changes may be able through the work of John New family for UT flora, includes more controversial, such as splitting Kartesz and the BONAP program. Anagallis and Lysimachia (from Arabis into Boechera and Turritis Plant Endemism and Geoen- Primulaceae). Genus Glaux (though there is good evidence for demic Areas of Utah. By Stanley transferred to Lysimachia this), or recognizing new taxa of Welsh and N. Duane Atwood. 2009. Boechera that are of complex apo- Resedaceae Self-published, Orem, UT, 97 pp. mictic or hybrid origin. Oligomeris linifolia—new for UT Only a handful of states have more Personally, I‟m more concerned based on an 1877 Palmer collec- species of vascular plants than Utah. with species treatments that don‟t tion from “Southern Utah” The size of the state flora is caused, seem to match reality in the field. in part, by the high number of en- Elimination of all varieties of Saxifragaceae demic species found mostly or en- Lepidium montanum with the weak Boykinia: transferred to Telesonix tirely within Utah. Most of these explanation that the dozen or so Micranthes: formerly included in endemics are restricted to unusual vars. recognized by Reed Rollins and Saxifraga geologic substrates, of which Utah is C.L. Hitchcock intergrade is not an particularly rich. Welsh and Atwood advancement of the science and Theophrastaceae tease apart these patterns in their risks trivialization of numerous dis- New family for UT flora— only latest contribution to Utah botany. tinct and localized endemics (such includes genus Samolus The book includes a lot of back- as vars. claronense and neeseae). (previously in Primulaceae) ground information on the unique Many new Draba species are recog- attributes and flora of different geo- nized, including some like D. santa- graphic areas of the state, as well as quinensis from Utah that are quite long, slender styles of desert areas a history of botanical exploration of localized, but other equally uncom- of southwestern Wyoming and Utah. Welsh is an engaging story- mon species seem to have been sum- adjacent Colorado and Utah is not teller and the book is entertaining, marily discarded. Despite the con- “identical” to D. pectinipila, a even if your passion is not plant ge- tentions of the authors, the type of dwarf alpine species with short, ography. Draba juniperina, a tall species with stubby styles endemic to alpine

11 Utah Native Plant Society

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