July-August 2010 Sego Lily Newsletter, Utah Native Plant Society

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Sego Lily July 2010 33 (4)

Copyright 2010 Utah Native Plant Society. All Rights Reserved.

The Scarlet-Flowered Species of Echinocereus in Utah

have not been studied extensively orthoroughly enough.

Unfortunately Utah is a backwa-ter for cactus research, so membersof the cactus family are less well un-derstood here than in better-researched states such as Arizona.To my knowledge we have never hada professor at one of our Utah col-leges or universities interested inCactaceae, with a stable of eagergraduate students to do the work.In this some- [continued on page 4]

July 2010 (volume 33 number 4)

By Dorde W. Woodruff

The scarlet-floweredEchinocerei are predomi-

nantly hummingbird-pollinated andhave large, showy, bright red flowersin cespitose clumps that can reachan impressive size of a hundred ormore heads. Sizeable clumps withtheir bright flowers are a strikingfeature of late spring. They are a

fairly common sight in Utah‟s canyons, foothills, plateaus, and thehigher parts of desert valleys. Theclumps commonly grow in rocky places and usually on slopes.

Botanists have often sought toidentify cactus specimens fromUtah on the basis of keys derivedfrom other states that don‟t work well here or are not valid. Many cactus populations in our state

In this issue:

The Scarlet-Flowered Species of Echinocereus in Utah . . . . . . . 1, 4

Chapter News . . . . . . . . . . . . . . . 2 Annual Field Trip Highlights . . . 3Bulletin Board . . . . . . . . . . . . . . . 3

New San Juan/Four CornersNative 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: InBloom Year-Round . . . . . . . . 9

Botanist‟s Bookshelf: SummerReading Special . . . . . . . . . . . . 10

Cover: A small clump of Claret cup(now called Echinocereus mojaven-sis ) in the San Rafael Swell. Photoby Dorde W. Woodruff



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Utah Native Plant Society

Officers

President: Walter Fertig (Kane Co) Vice President: Kipp Lee (Salt Lake Co)Treasurer: Charlene Homan (Salt Lake

Co)Secretary: Mindy Wheeler (Summit

Co)Board Co-Chairs: Bill King (Salt Lake

Co) and Dave Wallace (Cache Co)

UNPS Board: Loreen Allphin (UtahCo), Robert Fitts (Utah Co), Susan Fitts(Utah Co), Ty Harrison (Salt Lake Co),Celeste Kennard (Utah Co), MargaretMalm (Washington Co), Larry Meyer(Salt Lake Co), Therese Meyer (Salt

Lake Co), Leila Shultz (Cache Co),Maggie Wolf (Salt Lake Co).

CommitteesCommunications: Larry MeyerConservation: Bill King and Tony

Frates

Education: Ty HarrisonHorticulture: Maggie Wolf Invasive Weeds: Susan FittsRare Plants: Walter FertigScholarship: Bill Gray

Chapters and Chapter PresidentsCache: Michael PiepCedar City: Marguerite SmithEscalante: Harriet Priska

Fremont: Lisa WhiteManzanita: Walter FertigMountain: Mindy WheelerSalt Lake: Marni AmbroseSouthwestern/Bearclaw poppy: Mar-

garet MalmUtah Valley: Celeste Kennard

Website: For late-breaking news, theUNPS store, the Sego Lily archives,Chapter events, links to other websites(including sources of native plants andthe digital Utah Rare Plant FieldGuide), and more, go to unps.org.Many thanks to Xmission for

sponsoring our website.For more information on UNPS:

Contact Bill King (582-0432) or SusanFitts (801-756-6177), or write toUNPS, PO Box 520041, Salt Lake City,UT, 84152-0041 or [email protected]

___________________________________________________________________________________________________

Sego Lily Editor: Walter Fertig([email protected]). The deadline forthe September 2010 Sego Lily is 15 August 2010.

Copyright 2010 Utah Native Plant So-ciety. All Rights Reserved

The Sego Lily is a publication of theUtah Native Plant Society, a 501(c)(3)

not-for-profit organization dedicatedto conserving and promoting steward-ship of our native plants. Use of con-tent material is encouraged but re-quires permission (except where ex-empted by statute) and must be cor-rectly credited and cited. Articles,photographs and illustrations submit-ted to us remain the property of thesubmitting individuals or organiza-tions. Submit permission requests [email protected]. We encourage read-ers to submit articles for potentialpublication. By submitting an article,an implicit license is granted to print

the article in the newsletter or otherUNPS publications for reprint withoutpermission (in print and electronicmedia). When submitting an article,please indicate whether it has beenpreviously published or submitted forconsideration to other publications.

the trail on Carmel Limestone.Several other plants of interest were found with the cryptantha,including the yellow-floweredCharleston Mountain violet (Viola purpurea var. charlestonensis)and Rose‟s spring-parsley (Cymopterus purpureus var. ro-sei ). In all, we found 132 vascularplant species in bloom.

Our next field trip will be onSaturday, 17 July to the Red Can-

yon Botanical Area on Dixie Na-tional Forest, off UT Hwy 12 west of Bryce Canyon National Park. We

will carpool from the Grand Stair-case-Escalante NM visitor centerparking lot in Kanab at 8 AM andplan to arrive at the botanical area by 9:30 AM for a leisurely half day of searching for Claron endemicsamong the Bristlecone pines. Formore info, contact me([email protected]).— W . Fertig.

Salt Lake: There are three remain-ing field trips scheduled for summer2010:

Saturday, July 17: Upper Lambs

Canyon with Bill Stockdale andMindy Wheeler

Saturday, August 7: Brighton tosee Wood nymph ( Monesesuniflora) with Bill Nelsen

Saturday, August 14: Hiking andpotluck in Upper City Creek Canyon with Marni Ambrose.

For more information on thesetrips, contact Bill Gray at [email protected] or 801-532-3486.— Bill Gray

Chapter News

Manzanita (Kane County): On

May 28, a dozen Kanab area plantenthusiasts embarked on an expedi-tion to the East Rim of Zion Canyonto view the spring wildflowers of Zion National Park. Part of our mis-sion was also to confirm the pres-ence of Dwarf cryptanth (Crypt-antha humilis, below), a species which had been previously reportedfor Zion NP, but not photographedor documented with a museum voucher. We successfully located asmall patch of the low-growing,

white-flowered, bristly plants along



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Bulletin Board

New San Juan/Four Corners Native Plant Society: Many of youmay already be aware of the „Southwest Colorado Wildflowers‟ website cre-ated by Al Schneider (www.swcoloradowildflowers.com/) - it is a real treas-ure trove of information and photos about the plants of the Four Cornersarea. Al has now started a new society dedicated to the native plants of thearea called the San Juan/Four Corners Native Plant Society. He invites all

UNPS members and especially those from the southeast part of Utah to par-ticipate in his field trips and other events. Follow the link above to get moreinformation or contact Al directly if you would like to be included in emailnotification. - Bill Gray

Global Garlic Mustard Field Survey: Do you want to be part of the world‟s largest scientific research project on invasive species? The „GlobalGarlic Mustard Field Survey‟ is an international collaboration to obtainmuch-needed data on the abundance and distribution of Garlic mustard( Alliaria petiolata) across its native and introduced ranges. In our firstfield season last year, we received measurements and seed samples from 65populations, with a majority from Europe. Our goal for this summer is 150or more, with a stronger emphasis on the southern and mid-west to westernUnited States.

This year we are hoping to increase participation among educators, as well as land managers and citizen scientists who may not have much formalscience training. The survey involves a simple protocol that can be followeddirectly or incorporated into field courses and nature surveys. A populationtakes two people about 2-4 hours to measure. We are also planning to develop internet-based teaching modules and tools to aid with monitoring andmanaging this invasive plant. The sampling protocol, along with contactinformation 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 CedarBreaks National Monument in scenic southeastern Iron County, Utah (23miles east of Cedar City). Volunteer wildflower specialists will be on handfor guided hikes at 10 AMand 1 PM daily. A JuniorRanger “wildflower scaven-ger hunt” will be held onFridays and Saturdays dur-ing the festival at 3 PM. Allactivities begin at the FeeBooth area (adjacent to themain parking area, about 1.5miles inside the monument‟s

south entrance). The visitorcenter will also be hostingan ongoing electronic dis-play of wildflower imagesand offer free wildflowerphotography tip sheets, and discounts on wildflower-themed books. Formore information on the festival and associated activities, go to the CedarBreaks National Monument website, www.nps.gov/cebr or call 435-586-0787. There is an entrance fee of $4 for Cedar Breaks National Monumentfor persons 16 and older and visitors in July should be prepared for poten-tial afternoon showers and temperatures dropping to the 60s.

Annual Field TripHighlights

On June 12, the Manzanita Chap-ter hosted the UNPS state board forits yearly foray to southern Utah. Aspart of the festivities, the chaptersponsored the first botanical foray of

the Best Friends Animal Sanctuary,located 7 miles north of Kanab. BestFriends is a no-kill animal shelterfocusing on dogs, cats, horses, rab- bits, and exotic birds. The groundsof the sanctuary include severalhundred acres of unspoiled redrock canyons, pinyon-juniper forest, sanddunes, and riparian woodlands. Ourgoal was to develop a plant specieslist for the sanctuary.

Our small but enthusiastic teamof botanists and naturalists spent 6hours scouring the area in search of new plant species. EntomologistKen Kingsley (below) provided ex-pert commentary on a number of unusual insects in the area, includ-ing the yucca moth and the parasi-toid wasp that preys on its develop-

ing larvae within the flower of theNarrow-leaved yucca (Yucca angus-tissima).

Prior to the foray, I had collectedor observed 179 vascular plant spe-

cies in the Best Friends area. By day‟s end our group had observed129 plant species, of which 36 werenew to the sanctuary. Our day‟s work increased the flora of the area by nearly 17%, bringing the totalnumber to 215 species.

After the board meeting, ourgroup reconvened at the home of Best Friends co-founder Jana dePeyer for a potluck dinner and gen-eral merriment.—Walter Fertig



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Above: Hedgehog in the city: A clumpof Echinocereus mojavensis percheshigh above Salt Lake City in quartzite. Photo by Tony Frates.

Most of Benson‟s work was fin-ished well in advance of publica-tion of his 1982 major work TheCacti of the United States and Canada. Not only must so compre-hensive a work necessarily be doneover a period of years, but also be-

cause it is a large and expensive volume, the Stanford University Press had to wait several years af-ter completion of the manuscriptuntil it had accumulated funds toprint it, so in spite of Benson‟s longand patient work on the cacti, it was outdated even when it first waspublished. Nevertheless, it was themost comprehensive work on thefamily for a long time.

In 1985 Nigel Paul Taylor‟smonograph “The Genus Echinocer-eus” was published. Taylor is a bota-nist specializing in the Cactaceae at

the famous Royal Botanic Gardensat Kew, England. This book coversthe whole genus (not just those of the U. S. and Canada), and includesthree species in Sect. Triglochidia-tus: E. scheeri , a complex Mexicanspecies, E. triglochidiatus, and E. polyacanthus (which is now consid-ered a synonym of E. coccineus).

The treatment of Echinocereus for the 2003 volume of Flora of North America (FNA) was written by Allan Zimmerman and BruceParfitt, both one-time students of

Donald Pinkava‟s at Arizona StateUniversity in Phoenix. They in-cluded three U.S. species in Sect.Triglochidiatus: E. coccineus, E.triglochidiatus and E. arizonicus. Although recognizing E. coccineus,they did not include Utah in its dis-tribution.

Like Benson, Pinkava is one of the Grand Old Men of cactus re-search; unlike Benson, he is still with us and still active. His special-ties are counting chromosomes and

studying polyploidy, and also thegenus Opuntia. Now retired from Arizona State University, he is stillconsulting with botanists of the De-sert Botanical Garden in Phoenixand others. The work of Pinkava andhis students in studying chromo-some numbers has been importantin sorting out Echinocereus.

Last fall, in one of my periodictours of herbaria, I visited theGarrett Herbarium at the University of Utah (U of U) and found that inpreparation for his treatment of the

genus for Intermountain Flora(IMF), Marc A. Baker (also once astudent of Pinkava‟s) had visited thethree major Utah herbaria: UtahState University and BYU, in addi-tion to the U of U, and annotated thespecimens of Echinocereus. Speci-mens formerly determined as E.triglochidiatus or one of its varieties were annotated by Baker mostly as E. mojavensis and a few as E.

gloomy picture the large collectionof cactus specimens at the Brigham Young University (BYU) is a greatasset.

Many botanists don‟t like to deal with cacti and they are underrepre-sented in herbaria. Because of theirsucculence and also their armor,making decent dried specimens outof them is difficult, and the driedspecimens are more changed inform from their live state than arethose of non-succulent plants. Thestudy of live plants in their environ-ment is more important in this fam-ily. Their morphology is different,and botanists who don‟t deal withthem much don‟t have an eye for

them. Consequently, the study of cacti lags behind that of most otherplant families, though more advanced in the southern states wherethere is more academic interest thanin Utah.

These red-flowered hedgehogcacti have been placed in a sub-genus, section Triglochidiatus. Go-ing through the literature on Echi-nocereus shows that this section hasnot been at all well understood. Various authorities have subdivided

the basal species E. triglochidiatus into a confusing array of species or varieties.

Quoting the late Lyman Benson,the largest single figure in cactusresearch, “Twenty -nine years of this[research] has not produced the ulti-mate answer” to resolve “…the insta- bility of the populations of the pro-posed species and their extensiveand bewildering intergradation witheach other.” In recent years mostexperts have used Benson‟s classifi-cation for this section, in which he

reduced the various scarlet-floweredspecies to varieties of E. triglochi-diatus. He did produce some orderfor these plants in his varieties, which others have built on. It wasnot a perfect analysis, but was the best available. I have followed Ben-son in determining all the scarlet-flowered hedgehogs in Utah as E.triglochidiatus but without the va-rieties, which didn‟t seem to work well in Utah.

The Scarlet-Flowered Species of Echinocereus in Utah(continued from page 1)



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coccineus. Before this E. mojavensis was usually recognized as a variety of E. triglochidiatus , and in previ-ous years E. coccineus was not gen-erally recognized as occurring inUtah.

Preparation for major works likethe various volumes of the continu-ing series of IMF and FNA offers a

great opportunity in making resultsof research accessible, as the experts who were chosen to write up thegenera review data and visit herbaria.

Scientific papers on the cactidon‟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 itsmore technical offspring Haseltonia but are scattered in many journals.The Internet makes it easier, but

even so, it takes a formidableamount of time spent searching tokeep up with current botanical lit-erature, even in this one family.

The forthcoming volume of IMFthat includes the Cactaceae will bring us a much better understand-ing of the species of sectionTriglochidiatus. So how did thisnew understanding occur? Baker,now an environmental consultant in Arizona, has been working on Echi-nocereus for years. Baker deter-mines the boundaries of taxa on the

basis of morphology, geographicdistribution, polyploidy, and floraldimorphism (in some species maleand female flowers are produced onseparate plants). For morphology,he measures sizes in a set of 22characters of flowers and stems, andanalyzes them using advanced sta-tistical techniques: analysis of vari-ance (ANOVA), discriminant analy-sis (DA), multivariate analysis of variance (MANOVA), principle com-ponent analysis (PCA), and un-

weighted pair group method witharithmetic mean (UPGMA). He alsohas extensive field experience andhas counted many a chromosomeset. As good botanists do, he studiestype specimens and the occurrenceof the plant at the type locality tounderstand what the author in-tended, and does not recognize taxauntil he has enough data to do so.

So what does Baker recognize? U.S. species recognized by him in sect.Triglochidiatus are E. arizonicus, E.

Above: Claret cup(Echinocereus coc-cineus) near the Petrified Forest Trail in Zion National Park. Photo by Tony Frates.

cineus in IMF, while placing E. can-yonensis in synonomy . It is possiblethat further study will change this.

The theory of its difference is thatthese plants grow in areas that are

hotter and drier, where humming- birds are not abundant, and there-fore are becoming adapted to beesrather than hummers. Flowersadapted to bees are smaller thanhummingbird flowers. Bees don‟tneed as much nectar as the larger birds with their high metabolisms,and often seek pollen rather thannectar. Though bees visit the scarlet-colored hummingbird flowers, they prefer yellow flowers. The predomi-nant pigments in the Cactaceae are

betalains, the chemistry of whichprecludes anthocyanins with their blue hues, which bees favor. Thescarlet hues of the hummingbirdhedgehogs come from flavonoids,and these flavonoids are differentpigments not found in most cacti.

The recognition that E. coccineus is found in Utah solved a longtimepuzzle for me. In May of 1973 I cameupon a curious population of hedge-hog cacti on the Kolob road. These

coccineus, E. mojavensis, E. san-taritensis, E. triglochidiatus, and E. yavapaiensis. And he recog-nized some Utah specimens as E coccineus. Our limited populationof E. coccineus, as far as is known,is only in the eastern part of Wash-ington County and mostly in andnear Zion. The Utah plants aredifferent than typical E. coccineus.

Ours are part of the group named by Michigan botanists Elzada Clo- ver and Lois Jotter (professor andstudent), the first women to runthe entire Colorado River, as E.canyonensis, with the type speci-men collected down in the GrandCanyon at Hermit Falls.

They are characterized by smaller flowers, by some (but notall) plants displaying one of a se-ries of flower colors trending to- wards yellow and away from thescarlet color of the hummingbird

pollination syndrome, and by be-ing visited by hordes of bees ratherthan hummers as the predominantpollinators. Our plants are north-ern outliers, with plants like ourson the Arizona Strip connecting tothe type locality in the Grand Can- yon.

Because there is some intergra-dation, and because this group isnot sufficiently well known, Bakerincludes these plants with E. coc-



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plants had a somewhat differentcolor of flower in orangish hues,moving away from the usual purescarlet, and were attracting swarmsof bees. I made specimens of it butdid not know what to call it, puttingit under the umbrella of E. triglochi-diatus but stating that it was differ-ent and being visited by bees.

At that time, the best and mostcomplete reference for keying outUtah cacti was Benson‟s 1969 3rd edition of The Cacti of Arizona; Ialso referred to his other 1969 book,The Native Cacti of California. Bothof these carried E. coccineus and E.canyonensis in synonomy (along with many other synonyms) but with no descriptions. Another refer-ence was Boissevain and Davidson‟s1940 Colorado Cacti, which did in-clude E. coccineus. But their concept

of the species was not the tetraploid,dimorphic entity as it is understoodtoday, on the basis of the type, andtheir circumscription of it did not fitthe environment of my specimen.This is not to dismiss their work;they did well for that time when somuch less was known about the Cac-taceae.

Baker‟s annotation of E. mojavensis for all the other Utah speci-mens of sect. Triglochidiatus wasmore puzzling. For one thing, speci-mens in our East Desert are somewhat different from those in our West Desert. For another, becauseof all the wildly different treatmentsof this section in cactus literaturethough the years, until now, E. mojavensis, which is based on a speci-men from the Mojave River in Cali-fornia, had not been clearly and con-sistently differentiated from E.triglochidiatus, based on a speci-men from Wolf Creek in New Mex-ico.

Addressing the first issue are two

statements, the first from FNA:“That taxon [mojavensis] includescurly-spined plants (mainly in Cali-fornia) and straight-spined plants(including most populations in Ari-zona, Utah, and western Colorado).” And from Baker, “…UPGMA sug-gests that E. triglochidiatus subsp.mojavensis [this was written beforehe had enough data to feel satisfiedin recognizing this taxon in specificrank] may be composed of morethan one taxonomically definable

open and chunky look, and has onaverage shorter stems, fewer ribs,areoles farther apart, and shorterand fewer spines.

E. coccineus in Utah may be dis-tinguished not only by spine struc-ture, but also by its location in Washington County (as far as isknown). E. mojavensis also occurs

there, but in more mesic locations. If flowering, E. coccineus will display smaller flowers with perhaps differ-ent colors, and is predominantly visited by bees rather than hum-mingbirds. Each plant will have only male or only female flowers. Stemand spine measurements overlap with E. mojavensis but the ones I‟veseen seem to be taller and moreslender with finer spines on better-separated areoles. This is tentative,however, and spine surface and

flower characters are more usefuluntil we know this species in Utah better. So far I‟ve accumulated about11 locations from various sources,most in or near the south end of Zion, but also extending from PahTempe Hot Springs near Hurricaneto the Little Creek Mountains.

According to specimens in themajor Utah herbaria, E. mojavensis is found in all Utah counties exceptthe far northern tier of Box Elder, Weber, Cache, Rich, Morgan, Davis,and Summit.

Baker has annotated the speci-mens in our three major herbaria, but most of the smaller Utah herbaria have probably not yet caughtup with this recent work.

References:

Baker, M.A. 2006 A new florally dimor-phic hexaploid, Echinocereus yavapaiensis sp. nov. (section Triglochidia-tus, Cactaceae) from central Arizona Plant Systematics and Evolution 258:63-83

Baker, M.A. 2006 Circumscription of Echinocereus arizonicussubsp. arizoni-cus: Phenetic analysis of morphologicalcharacters in section Triglochidiatus (Cactaceae), Part II Madroño 53(4): 388-399

Above: True Echinocereus triglochi-

diatus from the Sandia Mountains in New Mexico. Photo by Bob Sivinski.

group, a western group repre-sented by the type and an easterngroup, perhaps represented by the basionym Cereus mojavensis Engelm. & J. M. Bigelow var. zuniensis J. M. Bigelow & Engelm,for which the type locality is inCanyon Diablo, east of Flagstaff, Arizona.” To resolve this, ananalysis such as that employed by Baker needs to be done.

As to the second issue, is E. mojavensis the only sect. Triglochi-diatus taxon in Utah other than E.coccineus? Baker‟s herbarium vis-its showed that although E.triglochidiatus as now understoodoccurs near the Four Corners, ithas not appeared in herbaria asoccurring in Utah. To best differ-entiate between these three speciesin the field or in herbarium speci-mens, Baker uses spine characters.

Viewed under 30x magnification, E. coccineus has spines round incross section and mostly smooth-surfaced with little trend toward apapillate surface. E. mojavensis also has round spines but they arepapillate. And E. triglochidiatus has spines that are smooth butangled in cross section, fewer, andstouter. In the field E. triglochi-diatus is the most noticeably dif-ferent vegetatively, with a more



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A Comparison of Utah Echinocereus Species

By Tony Frates and Dorde Woodruff

Species Synonyms and notes Location in Utah(none limited to UT)

Description

Echinocereuscoccineus

Scarlet claret cup, Scarlethedgehog

The specific epithet coc-cineus=scarlet is somewhat a misnomer forours since they are much visited by bees and thus

drifting from the scarlethummingbird-attractingcolor

A confusing array of synonymshave been used; this taxon wasnot formerly well understood

Utah plants are part of the pro-posed E. canyonensis; morestudy is needed for acceptanceof this as a valid taxon

Only known from eastern Washington County

Stems in closely set clumps;flowers dimorphic, with aslightly constricted “waist”;7-19 smooth spines per ar-eole, uniform in colorthough gray in age, round incross-section; flowerssmaller, narrower, < 5 cmlong, may be more yellowishthan in E. mojavensis

Visited by bees more thanhummingbirds

Tetraploid, n=22

Echinocereusengelmannii

Engelmann's hedgehog

Vars. chrysocentrus, variegatus,& purpureus have been recog-nized in the past, but differencesare inconsistent or insufficiently documented and vars are nolonger accepted

A southern Utah, GreatBasin, and Colorado Pla-teau species in Beaver,Garfield, Iron, Kane,Juab, Millard, Tooele,San Juan, and Washing-ton cos.

Stems single to several inopen clumps but not moundforming; central spines withcontrasting light and dark colors, flat or angled; flow-ers perfect, pale to dark rose-pink, to 9 cm tall and wide

Bee pollinated; easily distin-

guished from other Utah Echinocereus

Tetraploid, n=22

Echinocereusmojavensis

Claret cup; Hedgehog;Hummingbird hedgehog

Most often known as E.triglochidiatus var. mojavensis, with a confusing number of other synonyms. Sometimesmisspelled as “mohavensis”. Thespecific epithet is unfortunately a misnomer since it is not re-stricted to the Mohave Desert, but named for the Mojave River.There may be differences be-tween western and easternplants.

The name var. inermis has beenapplied to the spineless form of far east-central Utah, but thistaxon is no longer accepted.

The most widespread Echinocereus in Utah,occurring in all countiesexcept the northern tierof Box Elder, Cache,Davis, Morgan, Rich, andSummit cos.

Plants in closely set clumps;flowers perfect, scarlet, usu-ally >5 cm long; 5-11 spinesper areole, uniform in colorthough gray in age, mostly round in cross-section,papillate (use 30x lens)

Hummingbird pollinated.

Diploid, n=11



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dodder develops two different exten-sions from its haustoria: a hand-likeform that surrounds the phloem anda straw-like form that pierces thexylem. In this way dodder can ob-tain everything it needs (water, min-eral nutrients and food) from itshost.

Most species of dodder can para-sitize many different host plants, butstudies have shown that doddergrows better on some hosts com-pared to others. Furthermore sev-eral studies have shown that dodderis able to preferentially infect thosehosts that provide the most benefit.Colleen Kelly, from Oxford Univer-sity, demonstrated this and foundthat the choice is made based on

chemicals in the bark that doddercan detect before forming haustoria.Kelly also found that having twodifferent hosts was better than one,although a second independentstudy failed to confirm this result.

The effects of dodder go beyond just individual host plants. By theend of a growing season a singledodder plant may form thousands of haustorial connections with many different host species and cover an

Above: 1796 print of Cuscuta europaeaby the painter Jacob Sturm fromDeutschlands Flora in Abbildungen.

choose a host is also reminiscent of

an insect herbivore or parasite.Researchers have demonstratedthat dodder stems will orient toward a tomato plant or even a vialof tomato extract as long as anodor can be detected. Other stud-ies have shown that dodder canlocate a host by the quality of lightreflected off the host‟s leaves and will even preferentially move toward hosts with higher chlorophyllcontent, in other words, those thatare greener, and presumably withmore sugars.

As soon as a dodder stem twinesaround its host it begins to formhaustoria, specialized short stemsthat tap into the host vascular tis-sue. Flowering plants have twokinds of vascular tissue. Xylemcarries water and mineral nutri-ents from the roots up the stem,and phloem moves energy-richcarbohydrates to the roots or otherareas of need. To match this,

Dodder Doesn’t Dodder Around

By Peter Lesica(Adapted from Kelseya, the

newsletter of the Montana NativePlant Society)

Dodders (Cuscuta spp.) aresurely among the world‟s mostunusual plants. They are parasiticon other flowering plants and lack leaves or any photosynthetic tis-sues. This is odd enough, butunlike most other parasitic plants,such as broomrapes (Orobanchespp.), paintbrushes (Castilleja spp.) or even mistletoes( Arceuthobium spp.), doddershave no roots. They can be annu-als arising from seed each year or

perennials arising from overwin-tering stem segments. Our specieshave yellow or orange, twiningstems. Dodders are closely relatedto morning glories, but the flowersare inconspicuous though some-times with intricately ornamentedcorollas. There are about 150 spe-cies of dodder worldwide, mostcommon in subtropical and tropi-cal America. Eleven species arereported for Utah, and based onthe small number of herbarium

collections and my own experi-ence, most (with the exception of C. pentagona) are not common inthe state. In Utah they have often been collected parasitizing nativeand introduced legumes as well asspotted knapweed and othermembers of the Aster Family.

In some ways, dodder actsmore like an animal than a plant!Other parasitic plants, such as broomrape, have seeds that germi-nate only when they are contacted by host root exudates, but notdodder. Dodder seeds germinateon the surface of the ground andthen forage for their host. Thestems grow outward while at thesame time waving around untilthey reach a host plant. The juve-nile stems can reach up to 6 cm(2.4 inches), but none are able toreach twice that far, and they mustfind a host within a few days or dietrying. The way dodder is able to



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area the size of a small house. Of course this can have significant ef-fects on the plant community. Sincethey are somewhat host-specific,dodders can alter community struc-ture by preferentially damagingsome species more than others. Forexample, University of Montana‟sRay Callaway and his collaborators

found that dodder reduced thedominance of glasswort ( Salicornia)in favor of sea-lavender ( Limonium)in California coastal marshes. Dod-ders also may damage commercialcrops such as tomato, pumpkin andalfalfa. Indeed, this is why we know so much about dodder ecology.Dodders‟ negative impacts also ex-tend beyond simple parasitism. Forexample, they can be conduits be-tween host plants for viruses, in-cluding disease-causing pathogens.

Some diseases can spread morequickly 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 seriousinvasive weed in China as well asPuerto Rico. The native doddercauses a decline in the invader re-sulting in greater nutrient availabil-ity to native members of the com-munity.

Although it might seem like hostplants are defenseless against the

wiley dodder, this may not always bethe case. Recently researchers havefound that some host plants transfermessenger RNA (mRNA) into theirdodder parasites. Some of thesemRNAs can incapacitate dodder‟sgenetic machinery, thereby reducingits ability to make proteins andgrow. This discovery has spawnedan interest in genetically engineer-ing crop plants that produce dodder-destroying mRNA. With luck agri-culturalists may be able to turn dod-

der‟s voracious appetite against it.

References:Koch, A. M., C. Binder, & I. R. Sand-

ers. 2004. Does the generalist parasiticplant 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 cuesguide host location and host selection by parasitic plants. Science 313: 1964-67.

Utah Botanica Odds and Ends from Utah Botany

Zion Vegetation Program: In Bloom Year-Round!

In 2007, Zion NP received funding to put in seed increase fields at its Na-tive Plant Nursery, with the goal of growing and harvesting large amounts of seed and using that seed for large-scale restoration projects such as fire re-habilitation, annual brome competition, and campground revegetation.Part of this funding included the purchase of a new seed storage cooler, which would be used to store seed from Zion and Cedar Breaks and PipeSpring National Monuments at a consistent temperature year-round. TheZion Veg Program has long needed a dependable unit for long-term seedstorage, and we got it in the form of a 10x12 walk-in refrigerator with light-ing, a floor drain, and adjustable temperatures that can be set as low as 35°F(below right).

To celebrate the new cooler and do away with unnecessary “white space,”the Veg Program hired fellow plant nerd Shannon Eberhard (below left), anaspiring plant illustrator and a former employee of the Fire Effects Monitor-ing Team, to paint the cooler with native plant illustrations (above). Shan-non did an amazing job, and that means that the Veg employees are lucky enough to look at beautiful, accurate native plant paintings on a daily basis.Right awn!! - Rebecca Lieberg, Zion Lead Reveg Bio Tech



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By Walter Fertig

Ahh summer … time to unroll thehammock, pour a cool drink, andrelax with a good botany book … ortwo. The following are some recenttitles you might consider for yoursummertime reading pleasure.

Wasatch Wildflowers: A Field Guide, by Steve Hegji, 2010,Cedar Fort, Inc, Springville, UT.207 pp. While this slim guide doesnot cover every plant species innorthern Utah‟s Wasatch Range, itdoes include 200 of the most com-mon or showy wildflowers. Eachcolor photo is accompanied by a brief, often entertaining caption.For example, American bistort

( Polygonum bistortoides) is de-scribed as being “… a bit like a giantQ-tip sticking up above all the otherplants.” A series of thumbnail sym- bols is used to organize brief discus-sions of key flower, leaf, and habitattraits and other information. Hegjiis a gifted photographer with a realeye for composition and interestingperspectives. His book is geared to anon-technical audience, but thephotos will be appreciated by allplant and nature lovers.

National Wildlife Federa-tion Field Guide to Wildflow-ers of North America, by DavidBrandenburg, 2010, Sterling Pub-lishing, New York. 673 pp. Com-pact but hefty, this field guide hasmore than 4000 color photos of 2200 native and introduced wild-flowers found across North America.The book is organized taxonomically rather than by flower color andshape as in many other popular fieldguides. Descriptions of species arenecessarily brief, and only a few of

the more common or showy speciesin each genus are featured, along with color range maps. An innova-tive feature is the use of thumbnailimages of different flower types ar-ranged by color and shape located atthe beginning of the book whichhelps orient the user to the correctfamily or genus without resorting todifficult keys. This book covers too wide of a geographic area and toofew species to be especially useful

Botanist’s Bookshelf: Summer Reading Special

Above: Oenothera howardii by Steve Hegji.

for identifying anything but themost showy species, but is an ex-cellent introduction to the overalldiversity of genera and species

found in North America. It would be an excellent companion to amore formal taxonomic textbook in teaching students about floristicdiversity.

The Sibley Guide to Trees, by David Allen Sibley, 2009, Al-fred A. Knopf, New York. 426 pp.Over the past decade Sibley hascreated a small publishing empire based on his series of bird guides, but now has branched into trees of North America. Done in much thesame style as his original bird field

guide, The Sibley Guide to Trees covers nearly 600 native and natu-ralized tree species with watercolorpaintings of leaves, flowers, fruits,twigs, bark, and other identifyingcharacteristics. The book has brief descriptions of each species and adetailed range map, as well as es-says on families and genera. Un-like many recent tree guides, this book is organized phylogenetically and not by leaf arrangement(opposite vs. alternate) or shape.

I noticed some minor errors, suchas the range map of Utah juniperomitting most of the Colorado Pla-teau, but no more than one mightexpect from an ornithologist writ-ing about botany. The book isprobably too large for field use, but would be a valuable reference forkeying out specimens at home orfor general information.

California Mosses, by Billand Nancy Malcolm, Jim Shevock,

and Dan Norris, 2009, Micro-OpticsPress, 430 pp. One of my annualNew Year‟s resolutions is to becomemore competent identifying mossesand bryophytes. California Mosses may just be the book I‟ve been wait-ing for to help me. The authors haveamassed an amazing array of colorphotos of 600 moss species of Cali-fornia that illustrate growth form,leaf shape, and important cellulardetails. Many of the photos arequite beautiful and their subjectsresemble pieces of stained glass art. Accompanying the brief descriptionsare black-and-white drawings of stylized leaves that emphasize diag-nostic differences between generaand species. A table at the end of

the book compares and contraststhese leaf shapes to help the profes-sional or novice bryologist placetheir unknown specimen into itsproper sequence. Though the book covers California specifically, my bryologist buddy John Spence as-sures me that most of the speciesfrom Utah and western North Amer-ica are included.

Flora of North America: vol-umes 7 and 8, by the Flora of North America Editorial Committee,

2009-2010. The Flora of North America (FNA) project was con-ceived nearly 20 years ago and 16 of the planned 28 volumes have now been published. The ambitious goalof the project is to produce a taxo-nomic treatment of the entire floraof the North American continent,including bryophytes, ferns, gymno-sperms, and flowering plants. Vol-umes include keys to all native andnaturalized species, range maps, brief taxonomic descriptions, andoften a line drawing (at least one per

genus). The two most recent vol-umes cover 30 families of floweringplants, including the Saxifragaceae,Primulaceae, Ericaceae, Salicaceae,and Brassicaceae.

Because it covers all of North America, the taxonomic keys in theFNA are often difficult as they mustfocus on obscure characters to splitout so many taxa, or species tend tocome out in the key at multiplepoints. Those interested in keying



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Major Taxonomic Changesin FNA volumes 7 & 8

Brassicaceae (Cruciferae) Arabis: split into three genera - Arabis (sensu stricto), Boechera,& Turritis

Cardaria: lumped with Lepidium Cusickiella: new genus (formerly included in Draba)

Lesquerella: lumped with Phy-saria

Schoenocrambe: lumped with Sisymbrium or moved to new Genus Hesperidanthus

Sinapis: formerly included in Brassica

Cleomaceae (formerly Capparaceae)

Peritoma: new genus for Cleomelutea and C. serrulata.

Crassulaceae Rhodiola: formerly included in

SedumTillaea: lumped with Crassula

EricaceaeFamily expanded to include theMonotropaceae and Pyrolaceae.

Ledum transferred to Rhododen-dron

MyrsinaceaeNew family for UT flora, includes

Anagallis and Lysimachia (fromPrimulaceae). Genus Glaux transferred to Lysimachia

ResedaceaeOligomeris linifolia—new for UT based on an 1877 Palmer collec-tion from “Southern Utah”

Saxifragaceae Boykinia: transferred to Telesonix Micranthes: formerly included in

Saxifraga

TheophrastaceaeNew family for UT flora— only includes genus Samolus(previously in Primulaceae)

limestone fellfields of BeartoothButte in Wyoming (but not in Mon-tana as reported). Likewise, Drabacalcifuga is a cryptic but distinctspecies from Montana and Wyomingthat can be readily distinguishedfrom D. oligosperma but has beensynonymized without comment.

The value of FNA would increase

greatly if more emphasis was placedon discussions of taxonomic prob-lems associated with species (likethe Drabas mentioned above). Inthe interest of brevity, such discus-sions are frequently excluded orkept too short to be meaningful.This is unfortunate, because withoutadequate explanation taxonomicdecisions can appear arbitrary or weakly supported. I would also liketo see the new volumes return to theoriginal practice of having range

maps depict the approximate areainhabited by a species, and not justhave one dot per state. This is re-dundant anyway, given that statesand provinces are listed in the textfor each species. The best maps inthe FNA series were those done forthe two grass volumes which hadcounty level distribution. There isno good excuse for excluding suchinformation as county-level distribution maps are now readily avail-able through the work of JohnKartesz and the BONAP program.

Plant Endemism and Geoen-demic Areas of Utah. By Stanley Welsh and N. Duane Atwood. 2009.Self-published, Orem, UT, 97 pp.Only a handful of states have morespecies of vascular plants than Utah.The size of the state flora is caused,in part, by the high number of en-demic species found mostly or en-tirely within Utah. Most of theseendemics are restricted to unusualgeologic substrates, of which Utah isparticularly rich. Welsh and Atwood

tease apart these patterns in theirlatest contribution to Utah botany.The book includes a lot of back-ground information on the uniqueattributes and flora of different geo-graphic areas of the state, as well asa history of botanical exploration of Utah. Welsh is an engaging story-teller and the book is entertaining,even if your passion is not plant ge-ography.

out their local species are betterserved by good state or regional flo-ras which only have to deal with asubset of all the taxa in the FNA.The real value of the FNA comes inits monographic treatment of fami-lies and genera. The taxonomic con-cepts and nomenclature used in theFNA should guide the treatment of

species in state and regional florasin the future.

Numerous taxonomic changeshave been made in volumes 7 and 8,especially in the Brassicaceae, Cap-paraceae (now Cleomaceae), andamong genera formerly included inthe Primulaceae, Pyrolaceae, andMonotropaceae. Such changes can be upsetting to those of us morecomfortable with the names welearned in our youth, but are not a bad thing if they represent advances

in taxonomic concepts (taxonomy isa science after all, and not juststamp collecting). The accompany-ing table summarizes the more sig-nificant changes that affect species,genus, and family concepts in Utah.

Many of the changes seem rea-sonable. A lot of morphological andgenetic data support merging themustard genera Lesquerella and Physaria. Likewise, no one will losesleep over combining Cardaria with Lepidium. Other changes may bemore controversial, such as splitting

Arabis into Boechera and Turritis(though there is good evidence forthis), or recognizing new taxa of Boechera that are of complex apo-mictic or hybrid origin.

Personally, I‟m more concerned with species treatments that don‟tseem to match reality in the field.Elimination of all varieties of Lepidium montanum with the weak explanation that the dozen or so vars. recognized by Reed Rollins andC.L. Hitchcock intergrade is not an

advancement of the science andrisks trivialization of numerous dis-tinct and localized endemics (suchas vars. claronense and neeseae).Many new Draba species are recog-nized, including some like D. santa-quinensis from Utah that are quitelocalized, but other equally uncom-mon species seem to have been sum-marily discarded. Despite the con-tentions of the authors, the type of Draba juniperina, a tall species with

long, slender styles of desert areasof southwestern Wyoming andadjacent Colorado and Utah is not“identical” to D. pectinipila, adwarf alpine species with short,stubby styles endemic to alpine



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July-August 2010 Sego Lily Newsletter, Utah Native Plant Society