Sep-Oct 2010 Sego Lily Newsletter, Utah Native Plant Society

8/8/2019 Sep-Oct 2010 Sego Lily Newsletter, Utah Native Plant Society

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Sego Lily September 2010 33 (5)

Copyright 2010 Utah Native Plant Society. All Rights Reserved.

September 2010 (volume 33 number 5)

In this issue:

Farewell to the Aceraceae:Changes in the AngiospermFamily Tree . . . . . . . . . . . . . . . 1, 4

Chapter News . . . . . . . . . . . . . . . 2Bulletin Board . . . . . . . . . . . . . . . 3Bladderworts: Terrors of theWetlands . . . . . . . . . . . . . . . . . . . 6

On the False Report of an EdwardPalmer Collection as a NewRecord for the Genus Oligomeris(Resedaceae) for Utah . . . . . . . . 8

Native Plant Profiles: Four-wingSaltbush . . . . . . . . . . . . . . . . . . . . 10

Farewell to the Aceraceae:Changes in the Angiosperm Family Tree

By Walter Fertig

The Maple family is dead.For sure there are still plenty of spe-cies of maples (Acer) across NorthAmerica and Asia, but the maplefamily (Aceraceae) is gone cutdown by a new generation of tax-onomists wielding DNA datasetsand modern phylogenetic theory.The maples and their close cousinsthe horse-chestnuts (Hippocast-anaceae) are now part of an ex-

panded Soapberry family (Sapind-aceae).The milkweed family (Asclepiad-

aceae) is also no more absorbed bythe Dogbanes (Apocynaceae). Gonetoo are the Goosefoots (Chenopod-iaceae), Duckweeds (Lemnaceae),Pyrolas (Pyrolaceae), and Waterleafs(Hydrophyllaceae). Some familiargroups like the Lilies (Liliaceae) andFigworts (Scrophulariaceae) havereceived [continued on page 4]

Arborus angiospermus, anevolutionary or phylogeneticfamily tree of the floweringplants or angiosperms, basedon recent taxonomic revisionsof the Angiosperm PhylogenyGroup (APG). Cartoon by W.Fertig.


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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 & Tony FratesEducation: Ty Harrison

Horticulture: Maggie WolfImportant Plant Areas: MindyWheelerInvasive Weeds: Susan FittsRare Plants: Walter FertigScholarship/Grants: Therese Meyer

Chapters and Chapter PresidentsCache: Michael PiepCedar City: Marguerite Smith

Escalante: Harriet PriskaFremont: Lisa WhiteManzanita: Walter FertigMountain: Mindy WheelerSalt Lake: Marni AmbroseSouthwestern/Bearclaw poppy: Mar-

garet MalmUtah Valley: Celeste Kennard

Website: For late-breaking news, theUNPS store, theSego Lily archives,Chapter events, sources of nativeplants, the digital Utah Rare PlantField Guide, 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 November 2010Sego Lily is 15October 2010.

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

TheSego 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.

ennial plant exchange on Septem-ber 18th at the Richfield City ParkPavilion on 300 North Main.UNPS member Barbara Jensen isthe inspiration behind this event.Barbara, a real Green recyclingpioneer, didnt want to throw awaythe extra plants in her garden.

To start off the event, RossMurdock, greenhouse manager atSouthern Utah University, will givea presentation on growing per-

ennials. The lecture and plant ex-change are free to the public butdonations will gladly be accepted for

the Richfield County Library.The lecture starts at 9 AM withthe plant exchange from 10 AM untilnoon. All gardeners are encouragedto contribute extra plants from theirgardens and take home new ones.The goal of the event is to promotewater wise gardening, recyclingplants, and learning something new.For more information, call Barbara(435-896-8798), Ron Parsons (435-527-4751) or Lisa White([email protected]).JanettWarner

Manzanita (Kane County): Ourannual fall native plant sale will takeplace on Saturday, 11 September2010, from 9-11 AM at the KanabFarmers Market. Janett Warner ofWildland Nursery in Joseph, UT willbe on hand with a selection of nativeshrubs, small trees, wildflowers, andgrasses adapted for southern Utahgardening. - W. Fertig

Chapter News

Escalante: September 24-25

Escalante Canyons Art Festival, 10AM to 5 PM. Please contact Harrietabout helping with the Native Planttable.

Saturday, October 2NationalPublic Lands Day Re-vegetationProject at Calf Creek Campgroundand Escalante River Trailhead. 30willing laborers are needed to plantnative plants in designated spots.Please contact Jeanie Linn by 31August ([email protected], 826-5624) if you can help.

Tuesday, October 12: DinosaurSalads: Flora of the Cretaceous ageKaiparowits Formation, GrandStaircase-Escalante National Monu-ment by Dr. Alan Titus, monumentpaleontologist at 7 PM at the Esca-lante Interagency Visitor Centerauditorium.Harriet Priska

Fremont (Richfield Area): TheFremont Chapter and Sevier CountyMaster Gardeners are hosting a per-


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Bulletin Board

Annual UNPS Members Meeting: The Societys annual fall membersmeeting is scheduled for Saturday, October 23 on the campus of Utah ValleyUniversity in Orem, UT. Dr. Jason Alexander, curator of the UVU Herbar-ium, will speak about current floristic projects for students and volunteersbeing sponsored by the university. The meeting will begin at 4PM in RoomPS110 of the Pope Science Building (located next to the Student Center). As

in years past, the meeting will include a New World potluck dinner featur-ing cuisine of North and South America. After dinner, Jasons lecture willbe in one of the nearby lecture halls in the Pope Science Building. The UVUherbarium is also located two doors down.

Utah Valley University is located just off I-15 at the University Parkwayexit. Parking lot L is located closest to the science building and has freeparking on the weekend. If this lot should be full for some unforeseen rea-son, additional parking is available nearby in Lot N near the entrance to thenew library or in lots T or U by the Events Center. There is a lot of con-struction taking place on campus this fall, so some of the enclosed walkwaysbetween buildings will not be open. Consult the printable campus map atthe UVU website (http://herbarium.uvu.edu/location.shtml) for more de-tailed directions.

Utah Valley Herbarium Events: The Utah Valley University Herbar-ium is hosting a series of events for member of the Utah Native Plant Soci-ety. Herbarium volunteer mounting days are continuing the first Saturdayof every month this fall. Three are currently scheduled on the 4th of Sep-tember, the 2nd of October, and the 6th of November, from noon until 4PM in the Herbarium (PS108). Due to the holidays, there will be no volun-teer days in December or January. Parking is free on Saturdays in Lot Nnear the library. For further information on either of these events, pleaseemail or call Jason Alexander at [email protected] or 801-863-6806.

UNPS Fall Student ScholarshipWinners: Andrew Rayburn, agraduate student from Utah State

University, was awarded the Fall2009 research scholarship by theUtah Native Plant Societys scholar-ship/grants committee. Andrew andcolleagues Jacob Davidson andHillary White are studying the en-dangered Maguires primrose(Primula maguirei) and its relation-ship to soil chemistry and microhabi-tat variability within its limited rangein Logan Canyon. In particular,these young researchers are lookingat possible facilitation of primroseseedling establishment by moss

patches growing in cracks and ledgesof its limestone cliff habitat. The re-sults of this study will help land man-agers better understand the habitatneeds of Maguires primrose and contribute to the ecological literature onrelationships between mosses and seedlings. UNPS is pleased to contribute$1000 in our scholarship/grant fund to help Andrew and colleagues recovertravel costs and fees for soil analysis associated with their project. Andrewand colleagues will provide a summary of their findings in a future issue oftheSego Lily. - W. Fertig

Salt Lake: Our potluck in CityCreek Canyon was held August 14th.After paying homage to the rareUtah angelica (Angelica wheeleri) -one topped out at 8 feet tall - wehiked the trail from the end of theroad. Lynn Bohs introduced us tosome neat tricks for identifyingplants that were not in flower. Then

we enjoyed an excellent picnic sup-per. Next day we had 10 people hikeup to the newly discovered popula-tion of Woodnymph (Moneses uni-flora) at Brighton. The very lateSpring and slow Summer has meantthat flowers in general are stayingarounf longer; about 80 specieswere in bloom and the Wood-nymphs were are their best. Thiswill surely become an annual pil-grimage, like our early Spring trip tosee Andersons buttercup (Ranun-

culus andersonii).Bill Gray

Southwestern/BearclawPoppy: Our September meetingwill be held on Wednesday, Sept.8th at 7 PM at the Springdale Can-yon Community Center. DonnaPeppin, botanist intern for Zion Na-tional Park is working towards get-ting the Zion working herbarium upand available for the public. Hertalk is entitled Seeding Natives af-ter Wildfire and Development ofZion National Parks Working Her-

barium. Please note that the chap-ter is moving its monthly meetingsto the second Wednesday of themonth.Barbara Farnsworth

Utah Valley:Anyone wishing topick up a few native plants can cometo the Wasatch Heritage Garden(1040 N 900 E in Provo) at 10 AMon Saturday, September 11. BitsySchultz will be on hand to dig upseveral chokecherry and hackberryseedlings that have grown out of

their intended spots at the garden.There are a few other plants thatcould be transplanted and seeds tocollect.Celeste Kennard

Right: Maguires primrose (Primulamaguirei) is known only from the dolo-mitic limestone cliffs of Logan Canyonin northeastern Utah and has beenlisted as Endangered under the Endan-gered Species Act since 1985. Photo byHillary White.


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chaotic jumble of plant species.Since Linnaeuss time, plant

taxonomists have been striving tocreate ever more natural combina-tions of species by including infor-mation from many sources, suchas floral and fruit morphology,embryology, wood anatomy, leafarchitecture, cytology, genetics,and the fossil record. Starting inthe 1860s with the acceptance ofDarwins theory of evolution, theprimary emphasis of taxonomy hasshifted from creating mere orderto identifying the underlyinggenealogical relationships amongspecies and families.

Over the last 40 years the domi-nant angiosperm classificationsystem has been that of the lateArthur Cronquist of the New York

Botanical Garden*. Cronquist splitthe flowering plants into six sub-classes of dicots and five sub-classes of monocots, with eachsubunit representing a major evo-lutionary line. Of these, the

*Russian botanist Armen Takhtajan andAmerican Robert Thorne independentlyderived comparable, though somewhatmore complex, systems at about the sametime as Cronquist, but their works have notbeen as widely used in North America.

[continued from page 1] extrememakeovers and while still alive, arebarely recognizable. Meanwhile,several formerly obscure families,like the Lopseeds (Phrymaceae),

Broomrapes (Orobanchaceae), andPlantains (Plantaginaceae) have at-tained prominence thanks to an in-flux of new species transferred fromelsewhere.

So what is going on?These changes are the result of

studies by the Angiosperm Phylog-eny Group (APG), an internationalconsortium of research institutesand professional taxonomists. TheAPG has been at work for nearly twodecades applying modern researchmethods and theory to several cen-turies-old riddles. What is the mostprimitive group of angiosperms?How natural are existing orders andfamilies? What does the family tree(phylogeny) of flowering plants looklike? Through sharing datasets andfindings, the APG is attempting toforge an elusive consensus amongtaxonomists. The work of APG,(originally published in 1999, up-dated in 2003, and most recentlyrevised in 2009) has corroboratedmany hypotheses of species relation-

ships among the angiosperms buthas also challenged long-held as-sumptions, much to the consterna-tion of some botanists.

Taxonomy has two main pur-poses: to provide standardizednames for distinct species and sub-species/varieties and to organizethese taxa into a logical sequence.The rules for naming species werelargely developed by Carolus Lin-naeus in the mid 1700s and sinceformalized and periodically updated

in the International Code of Botani-cal Nomenclature. Hundreds ofclassification systems have beenproposed over the last three millen-nia, beginning with the simplegrowth form approach (tree, shrub,perennial herb ) of Theophrastusin ancient Greece. Linnaeuss ownsexual system, based primarily onthe number and degree of fusion ofstamens per flower, was an earlyattempt to apply repeatable criteriato the problem of organizing the

Farewell to the Aceraceae: Changes in the Angiosperm Family Tree

Above: Bigtooth Maple (Acer grandidentatum) from Bells Canyon Trail, just east ofLower Bells Canyon Reservoir, Salt Lake County, UT. The Maple family(Aceraceae) is now part of an expanded Soapberry family (Sapindaceae) accordingto the Angiosperm Phylogeny Group. Photo by Steve Hegji.

Magnolia group (Magnoliidae) isthought to be the most primitiveand closest to the putative ancestralflowering plant. Typical Magnoliids,such as the magnolias (Magnoli-aceae), buttercups (Ranunculaceae),and water-lilies (Nymphaeaceae)have numerous, separate sepals andpetals, numerous stamens, manyunfused pistils, and pollen openingby a single germination pore. Otherdicot lines include the mostly wind-pollinated and petal-less Hama-meliidae (oaks, elms, birches, andsycamores), chemically-uniqueCaryophyllidae (carnations, buck-wheats, and cacti), the large andsomewhat amorphous Rosidae(roses, peas, maples, euphorbs, andparsleys) and Dilleniidae (mustards,heaths, violets, and willows), and

the Asteridae (asters, mints,phloxes, gentians) considered to bethe most advanced group because ofthe pronounced reduction and fu-sion of floral parts.

The monocots are believed to de-rive from the magnoliids throughthe primitive Alismatidae (mostlyaquatic species with numerous sta-mens and separate pistils such asthe arrowheads and pondweeds).Additional monocot lines include


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Family Changes Based on Recent Taxonomic Researchby the Angiosperm Phylogeny Group

Old Family New Family Aceraceae (maples) Sapindaceae (soapberries) Asclepiadaceae (milkweeds) Apocynaceae (dogbanes)Buddlejaceae (butterfly-bushes) Scrophulariaceae (figworts)Callitrichaceae (water-starworts) Plantaginaceae (plantains)

Chenopodiaceae (goosefoots) Amaranthaceae (amaranths)& Sarcobataceae (greasewood)Cuscutaceae (dodders) Convolvulaceae (morning-glories)Fumariaceae (fumitories) Papaveraceae (poppies)Hippuridaceae (mares-tails) Plantaginaceae (plantains)Hippocastanaceae (horse-chestnuts) Sapindaceae (soapberries)Hydrophyllaceae (waterleafs) Boraginaceae (borages)Lemnaceae (duckweeds) Araceae (arums)Liliaceae (lilies, in part: still Alliaceae (onions)

includesErythronium, Fritill- Amaryllidaceae (daffodils)aria, Lilium, Lloydia) Asparagaceae (asparagus)

Colchicaceae (crocus)Melanthiaceae (bunchflowers)Ruscaceae (butchers brooms)

Themidaceae (funnel-lilies)Monotropaceae (pinesaps) Ericaceae (heaths)Najadaceae (naiads) Hydrocharitaceae (frogbits)Pyrolaceae (pyrolas) Ericaceae (heaths)Scrophulariaceae (figworts, in Orobanchaceae (broomrapes: in-

part, still includesScrophularia cludes Castilleja, Cordylanthus,Verbascum, Buddleja) Orthocarpus, & Pedicularis)

Phrymaceae (lopseeds: includesMimulus & Mimetanthe)

Plantaginaceae (plantains: includesCollinsia, Penstemon, & Veronica)

Tiliaceae (basswoods) Malvaceae (mallows) Viscaceae (mistletoes) Santalaceae (sandalwoods)

Zannichelliaceae (horned pondweed) Potamogetonaceae (pondweeds)

Other changes:Celtis goes from Ulmaceae (Celtidaceae) to Cannabaceae (hemps)Nolina goes from Agavaceae to Ruscaceae (butchers brooms)Sambucus & Viburnum go from Caprifoliaceae to Adoxaceae (moschatels)

the Arecidae (palms and arums),Commelinidae (bromeliads, sedges,and grasses), Zingiberidae (bananasand gingers), and Liliidae (lilies, iris,and orchids). Like the Asteridae,the Liliidae are considered the mostevolutionarily advanced groupwithin their class.

The systems advocated by Cron-

quist, Taktahjan, and Thorne werederived from their authors encyclo-pedic knowledge of flowering plantdiversity and the taxonomic litera-ture. By contrast, the AngiospermPhylogeny Groups taxonomy is de-rived from pooling datasets and ex-periences of numerous individualresearchers, augmented by break-throughs in analyzing DNA thatwere unavailable even two decadesago. In addition, the APG has ap-plied formal cladistic methodology

to the problem of family relation-ships.The basic premise of cladistics is

that species and families can be or-ganized based on deviations from anoriginal set of shared characteristics.These changes can be depicted visu-ally as branches of a tree (eachbranch is a clade) and the distancebetween branches is analogous tothe degree of similarity betweentaxonomic groups. To be legitimateunder the rules of cladistics, familiesand higher taxonomixc groups must

include all species above a given forkin the tree (the decision of what forkto choose is left to the taxonomist).Families that are nested within forkscomprising another, related familycannot stand alone, regardless ofhow distinct they might appear oth-erwise. Thus, the maple and horse-chestnut branches nest within thatof the soapberries and must be in-cluded within an expanded familyconcept of Sapindaceae. Likewise,the milkweed clade falls within the

dogbanes, duckweeds within thearums, and so forth (see chart atright).

Other situations are more com-plex, such as the old Scrophulari-aceae where genera once included inthe Figwort family were scatteredamong numerous branches and in-tertwined with Orobanchaceae,Phrymaceae, and Plantaginaceae.Either all of these families had to bemerged into one very amorphousfamily, or they had to be reconstit-

uted into more evolutionarily co-herent subgroups. Unfortunately,due to the naming rules set downunder the International Code, thefamily names Orobanchaceae,Phrymaceae, and Plantaginaceae

had to be retained, even thoughthey are named for relatively unfa-miliar genera.

Another family that has beensplit up considerably is the Lili-aceae. For years, specialists haverecognized that the group was un-natural and served as a catch-allfor a diverse assemblage of mono-cots with six tepals and six sta-mens. Based on recent genetic andmorphological studies, several

lily genera have been relocated toother monocot families and orders.The false asphodels (Tofieldia)turn out to be more closely relatedto the arrowheads and are nowplaced in their own family

(Tofieldiaceae). Likewise, camas(Camassia) is better placed withthe yuccas and agaves (Agav-aceae). Other lily genera havebeen split into two main cladesbased on seed and nectary fea-tures. One, the asparagus line,includes the onions (Allium), fun-nel-lilies (Androstephium), andfalse Solomons seal (Maianthem-um), which turn out to be morerelated to the irises, orchids, andagaves than the true lilies. While


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the lily family remains, it is muchreduced and retains mostly the truelilies (Lilium), tulips (Tulipa),checker-lilies (Fritillaria), and troutlilies (Erythronium). There is stilldisagreement as to whether the segolilies and mariposas (Calochortus)belong here or in their own family,Calochortaceae. Other former lily

family members have been segre-gated, including the catbriers(Smilacaceae), Trilliums (Trilli-aceae), and death-camas (Melan-thiaceae).

Some of the changes proposed bythe APG remain controversial. Theborages (Boraginaceae) traditionallyhave been allied with the mints(Lamiaceae) on the basis of similarfruit structures: four 1-seeded nut-lets. DNA evidence suggests thesetwo groups are only distantly related

within the Asterid clade and that theborages should contain the water-leafs (Hydrophyllaceae), despite thelatter group (Phacelia, Hydrophyl-lum and relatives) having capsulefruits with numerous seeds.

Besides re-arranging plant fami-lies, the APG has altered Cronquistslong-standing family tree. The mostprimitive flowering plants are nowthought to be a group of herbs andshrubs that includes the water-lilies

and several small orders found-mostly in the south Pacific andAustralia. From this basal group,the angiosperms split into themagnoliid line (analogous to Cron-quists concept with a few of themost primitive forms and the but-tercups removed), the monocots,and the true dicots or eudicots.

Among the eudicots, the butter-cups diverged early, as did theCaryophyllidae. Two mainbranches later arose: the Rosids(which include most of CronquistsRosidae, Hamamelidae, and Dil-leniidae) and the Asterids (ex-panded from the original Asteridaeto include the umbels (Apiaceae),hollies (Aquifoliales), dogwoods(Cornales), and heaths (Ericales).

Of course no classification isever complete or universally ac-

cepted. Numerous refinementswere made in the third edition ofthe APG system published in 2009and more changes will likely ariseand be posted on the APG websitein the future (www.mobot.org/mobot/research/apweb/). Tax-onomists are still free to use sys-tems of their choosing in technicalmanuals, floras, and species check-lists. Users of these products willstill need to be fluent in multiple

family synonyms and concepts.We live in an era of unstable tax-

onomy, and this is not likely tochange any time soon. Efforts tocreate more natural taxonomic sys-tems, like that proposed by APG, areworthwhile, even though they maybe upsetting when they impact ourfavorite families or world view. Tax-

onomy is, after all, a legitimate sci-ence and not merely pasting and re-arranging stamps in a binder. Someof the changes proposed by APG willprove to be wrong in light of newdiscoveries and changes in theory(cladistics is not without its logicalshortcomings, particularly the prob-lems of hybridization and reticulateevolution). The goal of the perfect,natural classification will remainelusive, just as it has since Lin-naeuss time nearly 250 years ago.

ReferencesAngiosperm Phylogeny Group. 2003. An

update of the Angiosperm PhylogenyGroup classification for the orders andfamilies of flowering plants: APG II.Botanical Journal of the LinnaeanSociety 141:399-436.

Spears, P. 2006.A Tour of the Flower-ing Plants Based on the ClassificationSystem of the Angiosperm PhylogenyGroup. Missouri Botanical GardenPress, St. Louis, MO. 308 pp.

Bladderworts: Terrors of the Wetlands

By Stuart MarkowAdapted from Castilleja, the news-letter of the Wyoming Native Plant

Society

The idea of plants killing and eat-ing animals has intrigued man forcenturies. This fascination with car-nivorous plants may result from: 1)

the thought that there just might beplants that actually eat people asportrayed in such movie classics asLittle Shop of Horrors and The LostWorld, and 2) an apparent situationof serious role reversal; people areused to thinking of animals eatingplants rather than vice-versa. Infact, while the ability of certainplants to capture insects and othersmall invertebrates has been ac-cepted for a long time, the thoughtthat these plants might digest and

Left: Common bladderwort, Utriculariavulgaris orU. macrorhiza has yellow,snapdragon-like flowers and leaveswith bladders borne amid the roundedleaflet segments. Illustration from Brit-ton and Brown (1913).

absorb critters was roundly rejectedby biologists for many years. The

concept was energetically de-nounced with the same scientificreasoning that historically has beenused to deny reality on a number offronts: it just didnt seem right!

Thus, those who did suggest thatplants might be snacking on insectswere ridiculed or ignored. It wasntuntil the late 1800s that, with a se-ries of elegant experiments usingsundews and Venus flytraps, CharlesDarwin provided such conclusiveevidence for insectivory that the


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door flies open releasing a partialvacuum which sucks the victiminside. The door then shuts, im-prisoning the prey within.

Utricularia (Lentibulariaceaefamily) is a large genus of nearly300 species distributed world-

wide. Most are aquatic, some aremoist-terrestrial, a few are evenepiphytic. All three Utah speciesare strictly aquatic, lurking in shal-low wetlands and lunching on in-sects, water fleas, rotifers, cope-pods, and other minute critters.

The traps are scattered aboutthe plants in great numbers, some-times on the stems, sometimes onthe leaves, sometimes on both.They are generally oval-shaped,with antennae-like appendages

that appear to funnel prey to thetrap entrance. A hinged door infront of the bladder is attached tothe top of the opening and swingsinward. At the base are pointedbristles which act as trigger hairs.The trap is set by removal of mostof the water through glands lo-cated both inside and outside ofthe bladder. This removal gener-ates a partial vacuum and, as aresult, the door is held in place by

a very finely balanced tension in-volving hydrostatic pressure coun-tered by the doors weak grip on theedges of the bladder walls. Theslightest touch of the trigger hairs isenough to break this tension.

The exact mechanism by whichthe touch stimulus opens the doorhas not yet been determined. Some

researchers claim that the triggerhairs act as a lever, lifting the doorslower edge and breaking the tenuousconnection between the door andthe bladder walls. Others argue thatthe mechanical stimulus is trans-duced to an electrical signal whichreduces the rigidity of the door,causing it to buckle under the in-wardly directed hydrostatic pres-sure. In either case, when the doorloses its precarious grip it swingsopen and the prey is swept into the

bladder with the inrush of water.Once the vacuum is released thedoor swings back to its normal,closed position and the victim istrapped. Glands within the bladdersecrete digestive enzymes and acids,and the same glands absorb the nu-trients.

The activation of a trap results insome of the fastest movement everrecorded in plants. Older literaturereports that the sequence of eventsfrom tripping to re-closing the blad-der occurs in about 1/50th of a sec-

ond. However, recent investigationssuggest that the trapping processoccurs at a much faster rate, perhapsas quickly as 1/500th of a second.Unfortunately, the small size of thetraps makes the adventure difficultto observe and, while this operationprovides as dramatic a display asplants have to offer, few people haveactually seen bladderworts in action.

Because of their unique capabili-ties, carnivorous plants are oftencollected and cultivated as novelties.

Without special, individualized careby knowledgeable horticulturists,they usually die within a fewmonths. With the exception of theCommon bladderwort (Utriculariavulgaris), all of Utahs carnivorousplants are fairly rare within thestate, so removing them from theirpreferred sites cannot be justified.Besides, it may turn out that some ofthese are man eaters after all. It isbest to leave them be.

Above: Flatleaf bladderwort, Utricu-laria intermedia is distinctive in hav-ing bladders borne on separate leafsegments. It is rare in Utah, beingknown from only three locations inCache, Rich, and Wayne counties.Illustration from Britton and Brown(1913).

skeptics received little further at-tention. Subsequent investigationby Darwin and others served toelucidate the various mechanismsof carnivory in plants.

Today it is well documentedthat plants can and do prey on notonly insects, but a wide variety ofinvertebrates and even vertebrate

species as diverse as frogs, fish,birds, and mice. The number ofplants able to perform this feat isnot well established, with reportedfigures ranging from 400 to 600species. Part of this discrepancy isdue to the taxonomic conceptssubscribed to by various authors,but perhaps more importantlythere is not complete agreement asto what constitutes a carnivorousplant. In almost all cases, the ana-tomical equipment and processes

used are simply modifications ofstructures and processes foundthroughout the plant kingdom.None of the individual featurestraps, lures, directional guides,secretory glands, absorbingglandsare unique to carnivorousplants.

In North America, there are fivemain groups of carnivorous plants.The sundews (Drosera) havesticky leaves that catch small bugslike flypaper. Venus flytrap(Dionaea) has an ingenious hinged

leaf and trigger. Pitcher plants(Sarracenia andDarlingtonia)have leaves shaped like a vase thatdrown their prey in liquid pools.Butterworts (Pinguicula) havesticky, butter-yellow, basal leavesthat trap insects. Bladderworts(Utricularia) catch tiny underwa-ter creatures through the power ofsuction. Of these, only the blad-derworts occur naturally in Utah.None appear to be man-eaters.

The bladderworts feature the

most sophisticated trapping de-vices to be found among the car-nivorous plants. These consist oftiny, very numerous bladders withone-way entrances adapted tocatching miniscule swimmingcreatures in standing water or wetsoil. The bladder-like trap is pro-vided with a baited trap door com-plete with a tripping device. Oncea small animal brushes against thedoor, the trap is sprung and the


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By Jason Andrew AlexanderUtah Valley University Herbarium

In the July issue of theSego Lily,

a new taxon, Oligomeris linifolia(Vahl) J.F. Macbride (Resedaceae)was reported for Utah based onEd-ward Palmer 47, an 1877 collectiondeposited in the U.S. National Her-barium (US) at the Smithsonian In-stitution. This report originatedfrom the newly published Volume 7of theFlora of North America(FNA). Attributing this species andcollection to Utah, however, is prob-lematic. As has been the case formost of the past century, Edward

Palmer's collections have more fre-quently been mis-cited by 20th cen-tury taxonomists than any other19th century Utah collector.

For many early expeditions it canbe a challenge relocating the popula-tions from which specimens wereobtained. However, due to an un-usually persistent series of publica-tion and typographical errors whichoriginated in the 1870's with AsaGray at the herbarium of HarvardUniversity (GH), Edward Palmer'sspecimens have been the most prob-

lematic. Palmer's collections weredistributed by eastern taxonomistssuch as George Vasey (his 1869-1870 collections), Asa Gray (his 1874and 1877 collections) or Sereno Wat-son (his 1875 and 1879 collections).Palmer sent most of his original andduplicate sets to these botanists,who later sold & distributed themafter identification (McVaugh 1956).The specimens of 1877 were one ofthe only collections that Palmerhimself distributed, with some assis-

tance from Charles ChristopherParry, a prolific collector of RockyMountain plants who spent much ofhis later years in Iowa.

Some problems with the 1877collection are related to the mannerin which Palmer and Parry assem-bled the specimens for distribution.First, Palmer distributed his 1877collections (and many of his latercollections through the 1890's), witha handwritten serial number only,on a preprinted label entitled

overall confusion surroundingPalmer's 1877 collection, which wasfirst proposed by McVaugh & Kear-ney (1943) in a treatment of prob-

lematic type specimens for theAri-zona Flora. Problems with speci-mens or Gray's publications can beattributed to his handling of thespecimens at GH. It appears thatGray split mixed or misidentifiedspecimens and reassembled thoseelements onto separate sheets. As aresult, many GH specimens aremissing the original label assignedby Palmer. Of all the other originalsets examined by me in the last dec-ade, only specimens at GH and US

were found to be regularly missingPalmer's pre-printed label. Mostspecimens at US were assigned a"replacement" Department of Agri-culture label by George Vasey andsometimes Palmer's original is miss-ing. Additionally, Gray transcribedPalmer's original list upon receiptand amended some of Palmer'sidentifications with his own deter-minations into his manuscript ofcollections (Gray, no date), now de-posited in the Harvard Botany Li-braries. It can be inferred that

"southern Utah, northern Arizona,& c." (see image of the label fromPalmer 47, below). Between Sep-tember and December 1877, they

assembled and numbered the col-lections into at least 18 sets to besold to other botanists and her-baria. Palmer and Parry rapidlydistributed these sets to raisefunds for a combined trip to Mex-ico scheduled for departure inJanuary of 1878 (McVaugh 1956:72-74). Second, neither Palmernor Parry wrote tentative identifi-cations on the specimen labels.Instead the specimens were sortedtaxonomically, assigned a hand-

written serial number, resortedinto sets, and distributed withoutdeterminations. By late 1877, Graywas sent a complete set of speci-mens for determination.

Most of the evidence, however,supports the hypothesis that Graywas the primary contributor to the

Below:Palmers 1877 collection ofOl-igomeris linifolia is from Nevada, notUtah.

On the False Report of an Edward Palmer Collection as a New Recordfor the Genus Oligomeris (Resedaceae) for Utah


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sometime in December of 1877 Grayfinished his determinations and be-gan writing his type descriptions ofnew taxa, some of which were pub-lished in Gray (1878). Afterwards,some form of a list was sent to eachof the botanists and herbaria towhich a set was sent, containing theserial number, the determinations,

and Gray's abbreviations of Palmer'scollection localities. In fact, theidentifications on labels of the speci-mens of the 18 distributed sets allare written in the unique handwrit-ing common to the collections ofthose botanists or herbaria to whichthe set was sent, and not in Gray's,Parry's or Palmer's hand. All havedeterminations identical to thosefound in Gray's later manuscriptand a small number of specimens inthese sets even have Gray's locali-

ties.Though Gray can be attributed asthe primary source of error regard-ing Palmer's 1877 specimens, themajority of western taxonomistshave all unintentionally contributedto this accumulation of error overthe past 150 years. In addition,many taxa cited for Utah (those thathave not been recollected since his1877 expedition, for the most part)in many floristic treatments are ac-tually records from Arizona or Ne-vada. Since "southern Utah" is the

first state mentioned on Palmer'spre-printed label, many botanistshave historically assumed that Utahis the likely collection locality for amajority of his 1877 collection. Thiserror has been perpetrated, un-knowingly, by botanists who are un-aware that specific collection locali-ties for specimens prior to the1880's were frequently only re-corded in unpublished manuscriptsor published in catalogues, and notregularly written on the specimen

label (and frequently differed fromany generic title preprinted on thelabel itself).

McVaugh (1956: vi-vii) states thatin the 19th century, "the prevailingattitude toward documentation ofspecimens... was a careless one bytoday's standards... some[specimens were] sent out with gen-eralized geographical informationcovering a whole years work some have been misdated by as

Above: Flax-leaf whitepuff, a.k.a. Oli-gomeris linifolia is an annual or short-lived perennial forb with slender, suc-culent leaves, tiny, white, 2-petaledflowers, and flattened-globular fruits.The species was originally describedfrom the Canary Islands and rangesacross the deserts of northern Africaand the Middle East. Populations alsooccur from SE California and south-ern Nevada to western Texas andnorthern Mexico. Thomas Nuttall col-

lectedO. linifolia in southern Califor-nia as early as 1836. Some bio-geographers have questioned whetherthe North American populations arenative. A recent genetic study bySantiago Martin-Bravo and col-leagues comparing New and OldWorld populations found significantdifferentiation, suggesting that thespecies arrived in North America bylong-distance migration in the Qua-ternary, well before the modern era ofhuman-facilitated dispersal. Illustra-tion by W. Fertig.

much as a year, and assigned tononexistent localities." The bota-nists to whom Palmer sent hisspecimens for determination anddistribution, "wrote his labels andtranscribed his data in addition totheir own..., and perhaps theyshould not be censured too harshlyfor hasty and superficial attentionto the tedious details" (McVaugh,1956: vii-viii).

In regard to the citation ofOli-gomeris linifolia as a taxon new toUtah, it appears that this is simplyanother case in which the authorsfor the Resedaceae treatment forFNA failed to consult Asa Gray'smanuscript and (as is too commonlydone) assumed this specimen wascollected in southern Utah. In the

image on the previous page, the col-lection number 47, in the upper lefthand corner, is the only part of thelabel that was originally written byPalmer & Parry. The identificationand the collection locality are writ-ten in the hand of the worker who,in the late 1870's, was curating theColumbia University Herbarium(now part of the New York BotanicalGarden Herbarium, NY). There areat least 7 duplicates ofPalmer 47ineastern herbaria, including the two

cited in the North American Flora,and this one from NY is the only oneI have seen with Gray's manuscriptcollection locality. The correct cita-tion for this specimen is as follows:

Oligomeris linifolia (Vahl) J.F.Macbride; U.S.A. NEVADA. ClarkCo.: between St. Thomas and theMuddy River Valley [GH Manu-script locality: "Muddy"], 17-20April 1877,Dr. Edward Palmer 47(GH, NY!, 4 sheets, US, WIS).

ReferencesGRAY, A. 1878. Contributions to thebotany of North America.Proc. Am.Acad. Arts Sci. 13: 361-367.

GRAY, A. no date. Palmer - S. Utah& Arizona 1877. MiscellaneousPlant Lists. Volume 3. Unpublishedmanuscript deposited at the GrayHerbarium Library, Harvard Univer-sity. Cambridge, Mass. p. 26-31.

MCVAUGH, R. 1943. Edward Palmer'scollections in Arizona in 1869, 1876,and 1877. Part 1. General discussion;itinerary and sources.Amer. Midl.Naturalist. 29: 768--775.

MCVAUGH, R. 1956.Edward Palmer:plant explorer of the American West.University of Oklahoma Press. Nor-man, Oklahoma. 430 pp.

MCVAUGH, R.AND T. H. KEARNEY. 1943.Edward Palmer's collections in Arizonain 1869, 1876, and 1877. Part 2. A con-sideration of some Palmer collectionscited in the Flowering Plants andFerns of Arizona.Amer. Midl. Natu-ralist. 29: 776--778.


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faint. Yellow dyes were made byboiling the leaves with raw alum.Saltbush seeds are edible (with anaturally mild, salty taste) and wereparched by the Navajo to makeflour.

Four-wing is regarded as one ofthe most important browse species

in the western United States. Inpart this is due to the plants abilityto thrive in semi-arid saline andsandy basin habitats that covermuch of the intermountain west,where Four-wing saltbush is oftenone of the dominant shrub species.More importantly, the foliage andseeds are palatable and highly nutri-tious. They are eaten with relish bycattle, sheep, and goats (though nothorses) as well as deer, elk, ante-lope, and bighorn sheep. Nativebrowsers consume the plant mostly

in winter when saltbush is especiallyrich in carotene and protein. Ro-dents, sage grouse, Gambels quailand other nesting birds take advan-tage of the shrubs intricate branch-ing for shelter from predators andthe elements. If saltbush rangelandis well-rested in the summer(allowing fruits to ripen and seed-lings to sprout) it can sustain rela-tively heavy grazing use in winter.Too much browsing, however, can

By Walter Fertig

Four-wing saltbush (Atriplexcanescens) is a fairly non-descriptshrubby member of the goosefoot

family (Chenopodiaceae nowAmaranthaceae) that ranges acrossmuch of the southern Great Plains,Great Basin, and desert Southwestat elevations up to 8000 feet. It ischaracterized by grayish, linear tospoon-shaped leaf blades, yellowishtwigs, and small yellowish or brownwind-pollinated flowers clustered inslender inflorescences and lackingshowy petals. Mature fruits can beeye-catching because of their rela-tively large size (1/2 to 1 inches long

and broad) and four membranous,slightly toothed wings (thus, thename four- wing). Four-wing salt-bush is often abundant, especially insandy or saline areas, and easy toignore.

But when it comes to reproduc-tion, Four-wing saltbush is anythingbut ordinary. More than 90% of thesaltbush plants in any populationare dioecious, meaning that individ-ual shrubs are either completely sta-minate (male) or pistillate(female). Staminate plants onlyproduce flowers that have anthersand shed pollen, and usually occurin harsher microsites in the environ-ment. By contrast, pistillate plantsonly produce seed and are not capa-ble of self-fertilization. About 10%of plants in a population aremonoecious and have separate maleand female flowers on the same in-dividual.

What makes Four-wing saltbushunusual is that the sex of any givenplant is not genetically fixed (as in

most animals), but can changebased on environmental cues. Re-searchers have discovered that overa seven year period nearly 40% ofthe saltbush plants in a populationswitched sexes, with as many as20% doing so each year. Pistillateplants were more likely to changesides than staminate ones, especiallyin years following an unusually coldwinter, drought, or after an atypi-cally heavy fruiting season.

Native Plant Profiles: Four-wing Saltbush

Above: Pistillate (female) specimenof Four-wing saltbush laden withyoung, green, four-winged fruitingbracts in late summer. Each set ofbracts encloses a one-seeded dry fruit.Four-wing plants are unisexual(either staminate or pistillate) but canswitch sexes from year to year de-pending on environmental conditions.

Photo by W. Fertig.

Male plants that became fe-males enjoyed a reproductive ad-vantage over other pistillate plantsby being able to flower earlier in agood season. The result of all thisswitching is that sex ratios in salt-bush populations can becomeskewed and sexes partially segre-gated along gradients of environ-mental quality, with females pre-dominating in richer sites and able

to become larger and producemore fruits.

Whether Native Americans paidattention to the sexual intrigue ofFour-wing saltbush is unknown,but tribes did make extensive useof saltbush. Before the advent ofsteel axes made it easier to harvesttrees, many Indians used brittlesaltbush stems as fuel. Smokefrom burning leaves was thoughtto revive those feeling weak or


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lead to trampling of the brittle stemsand eventually drain the seedbank.

Fortunately, Four-wing saltbushreadily germinates from seed ifgiven half a chance. Seeds ripen inlate August and September and mayremain on the female plant into De-cember. Several months of coldstratification are required to inducegermination in nature. Given someprotection from insects, rabbits, ro-dents, and browsers, seedling four-wings can grow rapidly, adding asmuch as 18 inches of new growth.

Because of its tolerance of coldweather, drought, and poor soilsFour-wing saltbush has become afavorite species for use in reclama-tion projects. Once established, theplants extensive root system is ex-cellent for containing soil erosion.Four-wing is also becoming morepopular as an accent plant in desertlandscaping. In Utah, it is hardy innearly all settings except high eleva-tions and wetlands. Plants can begrown from stem cuttings or from

seed. Stock is also increasinglyavailable from commercial nurser-ies.

When growing from seed, expertsrecommend an after-ripening periodof up to 10 months, followed by 30-50 days of cold stratification. Beforeplanting, seeds should be soaked fortwo hours and the membranouswings removed (especially if beingplanted with a drill or mechanicalseeder). Seeds should be planted

Above: Silvery foliage of Four-wingsaltbush. The narrow leaves are anadaptation for water conservation inits semi-arid environment. Below:Four-winged pistillate bracts sur-round the one-seeded fruit. Thebracts turn brownish-tan when ripe.When planting, the wings should beremoved. Photos by Steve Hegji.

1/2 to 3/4 of an inch deep, de-pending on soil moisture andcoarseness. Seedlings require littlewater and are tolerant of full sun

and wind. They are best grown indry sandy or slightly alkaline soilin late fall, early winter, or veryearly spring. Four-wing stands

Celebrating Wildflowers:Plants of the Week

This essay is partly adaptedfrom a biography of Four-wingsaltbush prepared for the USForest Services CelebratingWildflowers website

(www.fs.fed.us/wildflowers/plant-of-the-week/index.shtml). Visit this web-site for a different native plantessay each week, and for otherinteresting and useful tidbitsabout pollinators, rare plants,wildflower events, and otherhappenings on US Forest Ser-vice lands.

may take 3-4 years to become fullyestablished, at which time they canbecome self-perpetuating throughvegetative layering or seed.

I find Four-wing saltbush to be anenjoyable plant to have in my yardin Kanab. It requires almost nomaintenance, other than periodicpruning of dead branches at thebase of the plant (these old stems goin the woodstove). My covey ofGambels quail hang out all year inthe dense growth, and mule deerbrowse on the foliage in winter. It isalso fun to keep tabs on whichplants produce fruit each year, andwhich ones will perform a sex-change next year!


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Sep-Oct 2010 Sego Lily Newsletter, Utah Native Plant Society