Farewell to the Aceraceae: Changes in the Angiosperm Family Tree Changes in the Angiosperm Family Tree

Home , Aceraceae, Acer grandidentatum, Androstephium, Carolus (plant), Celtis, Edward Palmer (botanist)

Sego Lily September 2010 33 (5)

September 2010 (volume 33 number 5)

In this issue: Farewell to the Aceraceae: Farewell to the Aceraceae: Changes in the Angiosperm Family Tree Changes in the Angiosperm Family Tree ...... 1, 4 Chapter News ...... 2 Bulletin Board ...... 3 Bladderworts: Terrors of the Wetlands ...... 6 On the False Report of an Edward Palmer Collection as a New Record for the Genus Oligomeris (Resedaceae) for Utah ...... 8 Native Plant Profiles: Four-wing Saltbush ...... 10

By Walter Fertig

The Maple family is dead. For sure there are still plenty of species of maples (Acer) across North America and Asia, but the maple family (Aceraceae) is gone — cut down by a new generation of taxonomists wielding DNA datasets and modern phylogenetic theory. The maples and their close cousins the horse-chestnuts (Hippocast- anaceae) are now part of an expanded Soapberry family (Sapind- aceae). The milkweed family (Asclepiad- Arborus angiospermus, an aceae) is also no more – absorbed by evolutionary or phylogenetic the Dogbanes (Apocynaceae). Gone family tree of the flowering too are the Goosefoots (Chenopod- plants or angiosperms, based iaceae), Duckweeds (Lemnaceae), on recent taxonomic revisions Pyrolas (Pyrolaceae), and Waterleafs of the Angiosperm Phylogeny (Hydrophyllaceae). Some familiar Group (APG). Cartoon by W. groups like the Lilies (Liliaceae) and Fertig. Figworts (Scrophulariaceae) have received [continued on page 4]

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

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ennials. The lecture and plant ex- Chapter News change are free to the public but donations will gladly be accepted for Escalante: September 24-25— the Richfield County Library. Escalante Canyons Art Festival, 10 The lecture starts at 9 AM with AM to 5 PM. Please contact Harriet the plant exchange from 10 AM until about helping with the Native Plant noon. All gardeners are encouraged table. to contribute extra plants from their Saturday, October 2—National gardens and take home new ones. Public Lands Day Re-vegetation The goal of the event is to promote Project at Calf Creek Campground water wise gardening, recycling and Escalante River Trailhead. 30 plants, and learning something new. willing laborers are needed to plant For more information, call Barbara native plants in designated spots. (435-896-8798), Ron Parsons (435- Please contact Jeanie Linn by 31 527-4751) or Lisa White August ([email protected], 826- ([email protected]).— Janett 5624) if you can help. Warner Tuesday, October 12: ―Dinosaur ennial plant exchange on Septem-

Salads: Flora of the Cretaceous age ber 18th at the Richfield City Park Manzanita (Kane County): Our Kaiparowits Formation, Grand Pavilion on 300 North Main. annual fall native plant sale will take Staircase-Escalante National Monu- UNPS member Barbara Jensen is place on Saturday, 11 September ment‖ by Dr. Alan Titus, monument the inspiration behind this event. 2010, from 9-11 AM at the Kanab paleontologist at 7 PM at the Esca- Barbara, a real Green recycling Farmer’s Market. Janett Warner of lante Interagency Visitor Center pioneer, didn’t want to throw away Wildland Nursery in Joseph, UT will auditorium.—Harriet Priska the extra plants in her garden. be on hand with a selection of native To start off the event, Ross shrubs, small trees, wildflowers, and Fremont (Richfield Area): The Murdock, greenhouse manager at grasses adapted for southern Utah Fremont Chapter and Sevier County Southern Utah University, will give gardening. - W. Fertig Master Gardeners are hosting a per- a presentation on growing per-

2 Sego Lily September 2010 33 (5) Salt Lake: Our potluck in City Bulletin Board Creek Canyon was held August 14th.

After paying homage to the rare Annual UNPS Member’s Meeting: The Society’s annual fall members Utah angelica (Angelica wheeleri) - meeting is scheduled for Saturday, October 23 on the campus of Utah Valley one topped out at 8 feet tall - we University in Orem, UT. Dr. Jason Alexander, curator of the UVU Herbar- hiked the trail from the end of the ium, will speak about current floristic projects for students and volunteers road. Lynn Bohs introduced us to being sponsored by the university. The meeting will begin at 4PM in Room some neat tricks for identifying PS110 of the Pope Science Building (located next to the Student Center). As plants that were not in flower. Then in years past, the meeting will include a New World potluck dinner featur- we enjoyed an excellent picnic sup- ing cuisine of North and South America. After dinner, Jason’s lecture will per. Next day we had 10 people hike be in one of the nearby lecture halls in the Pope Science Building. The UVU up to the newly discovered popula- herbarium is also located two doors down. tion of Woodnymph (Moneses uni- Utah Valley University is located just off I-15 at the University Parkway flora) at Brighton. The very late exit. Parking lot ―L‖ is located closest to the science building and has free Spring and slow Summer has meant parking on the weekend. If this lot should be full for some unforeseen rea- that flowers in general are staying son, additional parking is available nearby in Lot N near the entrance to the arounf longer; about 80 species new library or in lots T or U by the Events Center. There is a lot of con- were in bloom and the Wood- struction taking place on campus this fall, so some of the enclosed walkways nymphs were are their best. This between buildings will not be open. Consult the printable campus map at will surely become an annual pil- the UVU website (http://herbarium.uvu.edu/location.shtml) for more de- grimage, like our early Spring trip to tailed directions. see Anderson’s buttercup (Ranun- culus andersonii).—Bill Gray Utah Valley Herbarium Events: The Utah Valley University Herbar-

ium is hosting a series of events for member of the Utah Native Plant Soci- Southwestern/Bearclaw ety. Herbarium volunteer mounting days are continuing the first Saturday Poppy: Our September meeting of every month this fall. Three are currently scheduled on the 4th of Sep- will be held on Wednesday, Sept. tember, the 2nd of October, and the 6th of November, from noon until 4 8th at 7 PM at the Springdale Can- PM in the Herbarium (PS108). Due to the holidays, there will be no volun- yon Community Center. Donna teer days in December or January. Parking is free on Saturdays in Lot N Peppin, botanist intern for Zion Na- near the library. For further information on either of these events, please tional Park is working towards get- email or call Jason Alexander at [email protected] or 801-863-6806. ting the Zion working herbarium up and available for the public. Her UNPS Fall Student Scholarship talk is entitled ―Seeding Natives af- Winners: Andrew Rayburn, a ter Wildfire and Development of graduate student from Utah State Zion National Park’s Working Her- University, was awarded the Fall barium‖. Please note that the chap- 2009 research scholarship by the ter is moving its monthly meetings Utah Native Plant Society’s scholar- to the second Wednesday of the ship/grants committee. Andrew and month.—Barbara Farnsworth colleagues Jacob Davidson and

Hillary White are studying the en- Utah Valley: Anyone wishing to dangered Maguire’s primrose pick up a few native plants can come (Primula maguirei) and its relation- to the Wasatch Heritage Garden ship to soil chemistry and microhabi- (1040 N 900 E in Provo) at 10 AM tat variability within its limited range on Saturday, September 11. Bitsy in Logan Canyon. In particular, Schultz will be on hand to dig up these young researchers are looking several chokecherry and hackberry at possible facilitation of primrose seedlings that have grown out of seedling establishment by moss their intended spots at the garden. patches growing in cracks and ledges There are a few other plants that of its limestone cliff habitat. The re- could be transplanted and seeds to sults of this study will help land man- collect.— Celeste Kennard agers better understand the habitat

needs of Maguire’s primrose and contribute to the ecological literature on Right: Maguire’s primrose (Primula maguirei) is known only from the dolo- relationships between mosses and seedlings. UNPS is pleased to contribute mitic limestone cliffs of Logan Canyon $1000 in our scholarship/grant fund to help Andrew and colleagues recover in northeastern Utah and has been travel costs and fees for soil analysis associated with their project. Andrew listed as Endangered under the Endan- and colleagues will provide a summary of their findings in a future issue of gered Species Act since 1985. Photo by the Sego Lily. - W. Fertig Hillary White.

3 Utah Native Plant Society

Farewell to the Aceraceae: Changes in the Angiosperm Family Tree

[continued from page 1] extreme makeovers and while still alive, are barely recognizable. Meanwhile, several formerly obscure families, like the Lopseeds (Phrymaceae), Broomrapes (Orobanchaceae), and Plantains (Plantaginaceae) have at- tained prominence thanks to an in- flux of new species transferred from elsewhere. So what is going on? These changes are the result of studies by the Angiosperm Phylog- eny Group (APG), an international consortium of research institutes and professional taxonomists. The APG has been at work for nearly two decades applying modern research methods and theory to several centuries-old riddles. What is the most primitive group of angiosperms? Above: Bigtooth Maple (Acer grandidentatum) from Bells Canyon Trail, just east of Lower Bells Canyon Reservoir, Salt Lake County, UT. The Maple family How natural are existing orders and (Aceraceae) is now part of an expanded Soapberry family (Sapindaceae) according families? What does the family tree to the Angiosperm Phylogeny Group. Photo by Steve Hegji. (phylogeny) of flowering plants look like? Through sharing datasets and chaotic jumble of plant species. Magnolia group (Magnoliidae) is findings, the APG is attempting to Since Linnaeus’s time, plant thought to be the most primitive forge an elusive consensus among taxonomists have been striving to and closest to the putative ancestral taxonomists. The work of APG, create ever more natural combina- flowering plant. Typical Magnoliids, (originally published in 1999, up- tions of species by including infor- such as the magnolias (Magnoli- dated in 2003, and most recently mation from many sources, such aceae), buttercups (Ranunculaceae), revised in 2009) has corroborated as floral and fruit morphology, and water-lilies (Nymphaeaceae) many hypotheses of species relation- embryology, wood anatomy, leaf have numerous, separate sepals and ships among the angiosperms but architecture, cytology, genetics, petals, numerous stamens, many has also challenged long-held as- and the fossil record. Starting in unfused pistils, and pollen opening sumptions, much to the consterna- the 1860s with the acceptance of by a single germination pore. Other tion of some botanists. Darwin’s theory of evolution, the dicot lines include the mostly wind- Taxonomy has two main pur- primary emphasis of taxonomy has pollinated and petal-less Hama- poses: to provide standardized shifted from creating mere order meliidae (oaks, elms, birches, and names for distinct species and sub- to identifying the underlying sycamores), chemically-unique species/varieties and to organize genealogical relationships among Caryophyllidae (carnations, buck- these taxa into a logical sequence. species and families. wheats, and cacti), the large and The rules for naming species were Over the last 40 years the domi- somewhat amorphous Rosidae largely developed by Carolus Lin- nant angiosperm classification (roses, peas, maples, euphorbs, and naeus in the mid 1700s and since system has been that of the late parsleys) and Dilleniidae (mustards, formalized and periodically updated Arthur Cronquist of the New York heaths, violets, and willows), and in the International Code of Botani- Botanical Garden*. Cronquist split the Asteridae (asters, mints, cal Nomenclature. Hundreds of the flowering plants into six sub- phloxes, gentians) considered to be classification systems have been classes of dicots and five sub- the most advanced group because of proposed over the last three millen- classes of monocots, with each the pronounced reduction and fu- nia, beginning with the simple subunit representing a major evo- sion of floral parts. growth form approach (tree, shrub, lutionary line. Of these, the The monocots are believed to de- perennial herb …) of Theophrastus rive from the magnoliids through in ancient Greece. Linnaeus’s own *Russian botanist Armen Takhtajan and the primitive Alismatidae (mostly ―sexual system‖, based primarily on American Robert Thorne independently aquatic species with numerous sta- the number and degree of fusion of derived comparable, though somewhat mens and separate pistils such as stamens per flower, was an early more complex, systems at about the same the arrowheads and pondweeds). attempt to apply repeatable criteria time as Cronquist, but their works have not been as widely used in North America. Additional monocot lines include to the problem of organizing the 4 Sego Lily September 2010 33 (5)

the Arecidae (palms and arums), Family Changes Based on Recent Taxonomic Research Commelinidae (bromeliads, sedges, by the Angiosperm Phylogeny Group and grasses), Zingiberidae (bananas and gingers), and Liliidae (lilies, iris, Old Family New Family and orchids). Like the Asteridae, Aceraceae (maples) Sapindaceae (soapberries) the Liliidae are considered the most Asclepiadaceae (milkweeds) Apocynaceae (dogbanes) evolutionarily advanced group Buddlejaceae (butterfly-bushes) Scrophulariaceae (figworts) within their class. Callitrichaceae (water-starworts) Plantaginaceae (plantains) The systems advocated by Cron- Chenopodiaceae (goosefoots) Amaranthaceae (amaranths) quist, Taktahjan, and Thorne were & Sarcobataceae (greasewood) derived from their authors’ encyclo- Cuscutaceae (dodders) Convolvulaceae (morning-glories) pedic knowledge of flowering plant Fumariaceae (fumitories) Papaveraceae (poppies) diversity and the taxonomic litera- Hippuridaceae (mares’-tails) Plantaginaceae (plantains) ture. By contrast, the Angiosperm Hippocastanaceae (horse-chestnuts) Sapindaceae (soapberries) Phylogeny Group’s taxonomy is de- Hydrophyllaceae (waterleafs) Boraginaceae (borages) rived from pooling datasets and ex- Lemnaceae (duckweeds) Araceae (arums) periences of numerous individual Liliaceae (lilies, in part: still Alliaceae (onions) researchers, augmented by break- includes Erythronium, Fritill- Amaryllidaceae (daffodils) throughs in analyzing DNA that aria, Lilium, Lloydia) Asparagaceae (asparagus) were unavailable even two decades Colchicaceae (crocus) ago. In addition, the APG has ap- Melanthiaceae (bunchflowers) plied formal cladistic methodology Ruscaceae (butcher’s brooms) to the problem of family relation- Themidaceae (funnel-lilies) ships. Monotropaceae (pinesaps) Ericaceae (heaths) The basic premise of cladistics is Najadaceae (naiads) Hydrocharitaceae (frogbits) that species and families can be or- Pyrolaceae (pyrolas) Ericaceae (heaths) ganized based on deviations from an Scrophulariaceae (figworts, in Orobanchaceae (broomrapes: in- original set of shared characteristics. part, still includes Scrophularia cludes Castilleja, Cordylanthus, These changes can be depicted visu- Verbascum, Buddleja) Orthocarpus, & Pedicularis) ally as branches of a tree (each Phrymaceae (lopseeds: includes branch is a ―clade‖) and the distance Mimulus & Mimetanthe) between branches is analogous to Plantaginaceae (plantains: includes the degree of similarity between Collinsia, Penstemon, & Veronica) taxonomic groups. To be legitimate Tiliaceae (basswoods) Malvaceae (mallows) under the rules of cladistics, families Viscaceae (mistletoes) Santalaceae (sandalwoods) and higher taxonomixc groups must Zannichelliaceae (horned pondweed) Potamogetonaceae (pondweeds) include all species above a given fork in the tree (the decision of what fork Other changes: to choose is left to the taxonomist). Celtis goes from Ulmaceae (Celtidaceae) to Cannabaceae (hemps) Families that are nested within forks Nolina goes from Agavaceae to Ruscaceae (butcher’s brooms) comprising another, related family Sambucus & Viburnum go from Caprifoliaceae to Adoxaceae (moschatels) cannot stand alone, regardless of how distinct they might appear oth- erwise. Thus, the maple and horse- uted into more evolutionarily co- lily genera have been relocated to chestnut branches nest within that herent subgroups. Unfortunately, other monocot families and orders. of the soapberries and must be in- due to the naming rules set down The false asphodels (Tofieldia) cluded within an expanded family under the International Code, the turn out to be more closely related concept of Sapindaceae. Likewise, family names Orobanchaceae, to the arrowheads and are now the milkweed clade falls within the Phrymaceae, and Plantaginaceae placed in their own family dogbanes, duckweeds within the had to be retained, even though (Tofieldiaceae). Likewise, camas arums, and so forth (see chart at they are named for relatively unfa- (Camassia) is better placed with right). miliar genera. the yuccas and agaves (Agav- Other situations are more com- Another family that has been aceae). Other lily genera have plex, such as the old Scrophulari- split up considerably is the Lili- been split into two main clades aceae where genera once included in aceae. For years, specialists have based on seed and nectary fea- the Figwort family were scattered recognized that the group was un- tures. One, the asparagus line, among numerous branches and in- natural and served as a catch-all includes the onions (Allium), fun- tertwined with Orobanchaceae, for a diverse assemblage of mono- nel-lilies (Androstephium), and Phrymaceae, and Plantaginaceae. cots with six tepals and six sta- false Solomon’s seal (Maianthem- Either all of these families had to be mens. Based on recent genetic and um), which turn out to be more merged into one very amorphous morphological studies, several related to the irises, orchids, and family, or they had to be reconstit- agaves than the true lilies. While 5 Utah Native Plant Society the lily family remains, it is much and several small orders found- family synonyms and concepts. reduced and retains mostly the true mostly in the south Pacific and We live in an era of unstable tax- lilies (Lilium), tulips (Tulipa), Australia. From this basal group, onomy, and this is not likely to checker-lilies (Fritillaria), and trout the angiosperms split into the change any time soon. Efforts to lilies (Erythronium). There is still magnoliid line (analogous to Cron- create more natural taxonomic sys- disagreement as to whether the sego quist’s concept with a few of the tems, like that proposed by APG, are lilies and mariposas (Calochortus) most primitive forms and the but- worthwhile, even though they may belong here or in their own family, tercups removed), the monocots, be upsetting when they impact our Calochortaceae. Other former lily and the ―true dicots‖ or eudicots. favorite families or world view. Tax- family members have been segre- Among the eudicots, the butter- onomy is, after all, a legitimate sci- gated, including the catbriers cups diverged early, as did the ence and not merely pasting and re- (Smilacaceae), Trilliums (Trilli- Caryophyllidae. Two main arranging stamps in a binder. Some aceae), and death-camas (Melan- branches later arose: the Rosids of the changes proposed by APG will thiaceae). (which include most of Cronquist’s prove to be wrong in light of new Some of the changes proposed by Rosidae, Hamamelidae, and Dil- discoveries and changes in theory the APG remain controversial. The leniidae) and the Asterids (ex- (cladistics is not without its logical borages (Boraginaceae) traditionally panded from the original Asteridae shortcomings, particularly the prob- have been allied with the mints to include the umbels (Apiaceae), lems of hybridization and reticulate (Lamiaceae) on the basis of similar hollies (Aquifoliales), dogwoods evolution). The goal of the perfect, fruit structures: four 1-seeded nut- (Cornales), and heaths (Ericales). natural classification will remain lets. DNA evidence suggests these Of course no classification is elusive, just as it has since Lin- two groups are only distantly related ever complete or universally ac- naeus’s time nearly 250 years ago. within the Asterid clade and that the cepted. Numerous refinements borages should contain the water- were made in the third edition of References leafs (Hydrophyllaceae), despite the the APG system published in 2009 Angiosperm Phylogeny Group. 2003. An latter group (Phacelia, Hydrophyl- and more changes will likely arise update of the Angiosperm Phylogeny lum and relatives) having capsule and be posted on the APG website Group classification for the orders and families of flowering plants: APG II. fruits with numerous seeds. in the future (www.mobot.org/ Botanical Journal of the Linnaean Besides re-arranging plant fami- mobot/research/apweb/). Tax- Society 141:399-436. lies, the APG has altered Cronquist’s onomists are still free to use sys- Spears, P. 2006. A Tour of the Flower- long-standing family tree. The most tems of their choosing in technical ing Plants Based on the Classification primitive flowering plants are now manuals, floras, and species check- System of the Angiosperm Phylogeny thought to be a group of herbs and lists. Users of these products will Group. Missouri Botanical Garden shrubs that includes the water-lilies still need to be fluent in multiple Press, St. Louis, MO. 308 pp.

Bladderworts: Terrors of the Wetlands

By Stuart Markow Left: Common bladderwort, Utricularia Adapted from Castilleja, the news- vulgaris or U. macrorhiza has yellow, letter of the Wyoming Native Plant snapdragon-like flowers and leaves Society with bladders borne amid the rounded leaflet segments. Illustration from Brit-

ton and Brown (1913). The idea of plants killing and eating animals has intrigued man for centuries. This fascination with car- absorb critters was roundly rejected nivorous plants may result from: 1) by biologists for many years. The the thought that there just might be concept was energetically de- plants that actually eat people as nounced with the same scientific portrayed in such movie classics as reasoning that historically has been Little Shop of Horrors and The Lost used to deny reality on a number of World, and 2) an apparent situation fronts: it just didn’t seem right! of serious role reversal; people are Thus, those who did suggest that used to thinking of animals eating plants might be snacking on insects plants rather than vice-versa. In were ridiculed or ignored. It wasn’t fact, while the ability of certain until the late 1800s that, with a se- plants to capture insects and other ries of elegant experiments using small invertebrates has been ac- sundews and Venus flytraps, Charles cepted for a long time, the thought Darwin provided such conclusive that these plants might digest and evidence for insectivory that the 6 Sego Lily September 2010 33 (5) skeptics received little further at- a very finely balanced tension in- tention. Subsequent investigation volving hydrostatic pressure coun- by Darwin and others served to tered by the door’s weak grip on the elucidate the various mechanisms edges of the bladder walls. The of carnivory in plants. slightest touch of the trigger hairs is Today it is well documented enough to break this tension. that plants can and do prey on not The exact mechanism by which only insects, but a wide variety of the touch stimulus opens the door invertebrates and even vertebrate has not yet been determined. Some species as diverse as frogs, fish, researchers claim that the trigger birds, and mice. The number of hairs act as a lever, lifting the door’s plants able to perform this feat is lower edge and breaking the tenuous not well established, with reported connection between the door and figures ranging from 400 to 600 the bladder walls. Others argue that species. Part of this discrepancy is the mechanical stimulus is trans- due to the taxonomic concepts duced to an electrical signal which subscribed to by various authors, reduces the rigidity of the door, but perhaps more importantly causing it to buckle under the in- there is not complete agreement as wardly directed hydrostatic pres- to what constitutes a carnivorous sure. In either case, when the door plant. In almost all cases, the ana- Above: Flatleaf bladderwort, Utricu- loses its precarious grip it swings tomical equipment and processes laria intermedia is distinctive in hav- open and the prey is swept into the used are simply modifications of ing bladders borne on separate leaf bladder with the inrush of water. structures and processes found segments. It is rare in Utah, being Once the vacuum is released the known from only three locations in throughout the plant kingdom. Cache, Rich, and Wayne counties. door swings back to its normal, None of the individual features— Illustration from Britton and Brown closed position and the victim is traps, lures, directional guides, (1913). trapped. Glands within the bladder secretory glands, absorbing secrete digestive enzymes and acids, glands—are unique to carnivorous and the same glands absorb the nu- plants. door flies open releasing a partial trients. In North America, there are five vacuum which sucks the victim The activation of a trap results in main groups of carnivorous plants. inside. The door then shuts, im- some of the fastest movement ever The sundews (Drosera) have prisoning the prey within. recorded in plants. Older literature sticky leaves that catch small bugs Utricularia (Lentibulariaceae reports that the sequence of events like flypaper. Venus flytrap family) is a large genus of nearly from tripping to re-closing the blad- (Dionaea) has an ingenious hinged 300 species distributed world- der occurs in about 1/50th of a sec- leaf and trigger. Pitcher plants wide. Most are aquatic, some are ond. However, recent investigations (Sarracenia and Darlingtonia) moist-terrestrial, a few are even suggest that the trapping process have leaves shaped like a vase that epiphytic. All three Utah species occurs at a much faster rate, perhaps drown their prey in liquid pools. are strictly aquatic, lurking in shal- as quickly as 1/500th of a second. Butterworts (Pinguicula) have low wetlands and lunching on in- Unfortunately, the small size of the sticky, butter-yellow, basal leaves sects, water fleas, rotifers, cope- traps makes the adventure difficult that trap insects. Bladderworts pods, and other minute critters. to observe and, while this operation (Utricularia) catch tiny underwa- The traps are scattered about provides as dramatic a display as ter creatures through the power of the plants in great numbers, some- plants have to offer, few people have suction. Of these, only the blad- times on the stems, sometimes on actually seen bladderworts in action. derworts occur naturally in Utah. the leaves, sometimes on both. Because of their unique capabili- None appear to be man-eaters. They are generally oval-shaped, ties, carnivorous plants are often The bladderworts feature the with antennae-like appendages collected and cultivated as novelties. most sophisticated trapping de- that appear to funnel prey to the Without special, individualized care vices to be found among the car- trap entrance. A hinged door in by knowledgeable horticulturists, nivorous plants. These consist of front of the bladder is attached to they usually die within a few tiny, very numerous bladders with the top of the opening and swings months. With the exception of the one-way entrances adapted to inward. At the base are pointed Common bladderwort (Utricularia catching miniscule swimming bristles which act as trigger hairs. vulgaris), all of Utah’s carnivorous creatures in standing water or wet The trap is set by removal of most plants are fairly rare within the soil. The bladder-like trap is pro- of the water through glands lo- state, so removing them from their vided with a baited trap door com- cated both inside and outside of preferred sites cannot be justified. plete with a tripping device. Once the bladder. This removal gener- Besides, it may turn out that some of a small animal brushes against the ates a partial vacuum and, as a these are man eaters after all. It is door, the trap is sprung and the result, the door is held in place by best to leave them be.

7 Utah Native Plant Society

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

By Jason Andrew Alexander "southern Utah, northern Arizona, overall confusion surrounding Utah Valley University Herbarium & c." (see image of the label from Palmer's 1877 collection, which was Palmer 47, below). Between Sep- first proposed by McVaugh & Kear- In the July issue of the Sego Lily, tember and December 1877, they ney (1943) in a treatment of prob- a new taxon, Oligomeris linifolia assembled and numbered the col- lematic type specimens for the Ari- (Vahl) J.F. Macbride (Resedaceae) lections into at least 18 sets to be zona Flora. Problems with speci- was reported for Utah based on Ed- sold to other botanists and her- mens or Gray's publications can be ward Palmer 47, an 1877 collection baria. Palmer and Parry rapidly attributed to his handling of the deposited in the U.S. National Her- distributed these sets to raise specimens at GH. It appears that barium (US) at the Smithsonian In- funds for a combined trip to Mex- Gray split mixed or misidentified stitution. This report originated ico scheduled for departure in specimens and reassembled those from the newly published Volume 7 January of 1878 (McVaugh 1956: elements onto separate sheets. As a of the Flora of North America 72-74). Second, neither Palmer result, many GH specimens are (FNA). Attributing this species and nor Parry wrote tentative identifi- missing the original label assigned collection to Utah, however, is prob- cations on the specimen labels. by Palmer. Of all the other original lematic. As has been the case for Instead the specimens were sorted sets examined by me in the last dec- most of the past century, Edward taxonomically, assigned a hand- ade, only specimens at GH and US Palmer's collections have more fre- written serial number, resorted were found to be regularly missing quently been mis-cited by 20th cen- into sets, and distributed without Palmer's pre-printed label. Most tury taxonomists than any other determinations. By late 1877, Gray specimens at US were assigned a 19th century Utah collector. was sent a complete set of speci- "replacement" Department of Agri- For many early expeditions it can mens for determination. culture label by George Vasey and be a challenge relocating the popula- Most of the evidence, however, sometimes Palmer's original is miss- tions from which specimens were supports the hypothesis that Gray ing. Additionally, Gray transcribed obtained. However, due to an un- was the primary contributor to the Palmer's original list upon receipt usually persistent series of publica- and amended some of Palmer's tion and typographical errors which identifications with his own deter- originated in the 1870's with Asa Below: Palmer’s 1877 collection of Ol- minations into his manuscript of Gray at the herbarium of Harvard igomeris linifolia is from Nevada, not collections (Gray, no date), now de- University (GH), Edward Palmer's Utah. posited in the Harvard Botany Li- specimens have been the most prob- braries. It can be inferred that lematic. Palmer's collections were distributed by eastern taxonomists such as George Vasey (his 1869- 1870 collections), Asa Gray (his 1874 and 1877 collections) or Sereno Wat- son (his 1875 and 1879 collections). Palmer sent most of his original and duplicate sets to these botanists, who later sold & distributed them after identification (McVaugh 1956). The specimens of 1877 were one of the only collections that Palmer himself distributed, with some assis- tance from Charles Christopher Parry, a prolific collector of Rocky Mountain plants who spent much of his later years in Iowa. Some problems with the 1877 collection are related to the manner in which Palmer and Parry assembled the specimens for distribution. First, Palmer distributed his 1877 collections (and many of his later collections through the 1890's), with a handwritten serial number only, on a preprinted label entitled 8 Sego Lily September 2010 33 (5) sometime in December of 1877 Gray In regard to the citation of Oli- finished his determinations and be- gomeris linifolia as a taxon new to gan writing his type descriptions of Utah, it appears that this is simply new taxa, some of which were pub- another case in which the authors lished in Gray (1878). Afterwards, for the Resedaceae treatment for some form of a list was sent to each FNA failed to consult Asa Gray's of the botanists and herbaria to manuscript and (as is too commonly which a set was sent, containing the done) assumed this specimen was serial number, the determinations, collected in southern Utah. In the and Gray's abbreviations of Palmer's image on the previous page, the col- collection localities. In fact, the lection number 47, in the upper left identifications on labels of the speci- hand corner, is the only part of the mens of the 18 distributed sets all label that was originally written by are written in the unique handwrit- Palmer & Parry. The identification ing common to the collections of and the collection locality are writ- those botanists or herbaria to which ten in the hand of the worker who, the set was sent, and not in Gray's, in the late 1870's, was curating the Parry's or Palmer's hand. All have Columbia University Herbarium determinations identical to those (now part of the New York Botanical found in Gray's later manuscript Garden Herbarium, NY). There are and a small number of specimens in at least 7 duplicates of Palmer 47 in these sets even have Gray's locali- eastern herbaria, including the two ties. cited in the North American Flora, Though Gray can be attributed as and this one from NY is the only one the primary source of error regard- I have seen with Gray's manuscript ing Palmer's 1877 specimens, the Above: Flax-leaf whitepuff, a.k.a. Oli- collection locality. The correct cita- majority of western taxonomists gomeris linifolia is an annual or short- tion for this specimen is as follows: have all unintentionally contributed lived perennial forb with slender, suc- to this accumulation of error over culent leaves, tiny, white, 2-petaled Oligomeris linifolia (Vahl) J.F. the past 150 years. In addition, flowers, and flattened-globular fruits. Macbride; U.S.A. NEVADA. Clark many taxa cited for Utah (those that The species was originally described Co.: between St. Thomas and the have not been recollected since his from the Canary Islands and ranges Muddy River Valley [GH Manu- across the deserts of northern Africa 1877 expedition, for the most part) and the Middle East. Populations also script locality: "Muddy"], 17-20 in many floristic treatments are ac- occur from SE California and south- April 1877, Dr. Edward Palmer 47 tually records from Arizona or Ne- ern Nevada to western Texas and (GH, NY!, 4 sheets, US, WIS). vada. Since "southern Utah" is the northern Mexico. Thomas Nuttall col- first state mentioned on Palmer's lected O. linifolia in southern Califor- References pre-printed label, many botanists nia as early as 1836. Some bio- GRAY, A. 1878. Contributions to the have historically assumed that Utah geographers have questioned whether botany of North America. Proc. Am. is the likely collection locality for a the North American populations are Acad. Arts Sci. 13: 361-367. GRAY, A. no date. ―Palmer - S. Utah majority of his 1877 collection. This native. A recent genetic study by Santiago Martin-Bravo and col- & Arizona 1877.‖ Miscellaneous error has been perpetrated, un- leagues comparing New and Old Plant Lists. Volume 3. Unpublished knowingly, by botanists who are un- World populations found significant manuscript deposited at the Gray aware that specific collection locali- differentiation, suggesting that the Herbarium Library, Harvard Univer- ties for specimens prior to the species arrived in North America by sity. Cambridge, Mass. p. 26-31. 1880's were frequently only re- long-distance migration in the Qua- MCVAUGH, R. 1943. Edward Palmer's corded in unpublished manuscripts ternary, well before the modern era of collections in Arizona in 1869, 1876, or published in catalogues, and not human-facilitated dispersal. Illustra- and 1877. Part 1. General discussion; regularly written on the specimen tion by W. Fertig. itinerary and sources. Amer. Midl. Naturalist. 29: 768--775. label (and frequently differed from much as a year, and assigned to MCVAUGH, R. 1956. Edward Palmer: any generic title preprinted on the plant explorer of the American West. label itself). nonexistent localities." The botanists to whom Palmer sent his University of Oklahoma Press. Nor- McVaugh (1956: vi-vii) states that man, Oklahoma. 430 pp. specimens for determination and in the 19th century, "the prevailing MCVAUGH, R. AND T. H. KEARNEY. 1943. attitude toward documentation of distribution, "wrote his labels and Edward Palmer's collections in Arizona specimens... was a careless one by transcribed his data in addition to in 1869, 1876, and 1877. Part 2. A con- today's standards... some their own..., and perhaps they sideration of some Palmer collections [specimens were] sent out with gen- should not be censured too harshly cited in the ―Flowering Plants and eralized geographical information for hasty and superficial attention Ferns of Arizona.‖ Amer. Midl. Natu- ralist. 29: 776--778. covering a whole year’s work … to the tedious details" (McVaugh, some have been misdated by as 1956: vii-viii).

9 Utah Native Plant Society

Native Plant Profiles: Four-wing Saltbush

By Walter Fertig

Four-wing saltbush (Atriplex canescens) is a fairly non-descript shrubby member of the goosefoot family (Chenopodiaceae — now Amaranthaceae) that ranges across much of the southern Great Plains, Great Basin, and desert Southwest at elevations up to 8000 feet. It is characterized by grayish, linear to spoon-shaped leaf blades, yellowish twigs, and small yellowish or brown wind-pollinated flowers clustered in slender inflorescences and lacking showy petals. Mature fruits can be eye-catching because of their relatively large size (1/2 to 1 inches long and broad) and four membranous, slightly toothed wings (thus, the name ―four-wing‖). Four-wing saltbush is often abundant, especially in sandy or saline areas, and easy to ignore. Above: Pistillate (“female”) specimen faint. Yellow dyes were made by But when it comes to reproduc- of Four-wing saltbush laden with boiling the leaves with raw alum. tion, Four-wing saltbush is anything young, green, four-winged fruiting Saltbush seeds are edible (with a but ordinary. More than 90% of the bracts in late summer. Each set of naturally mild, salty taste) and were bracts encloses a one-seeded dry fruit. saltbush plants in any population parched by the Navajo to make are dioecious, meaning that individ- Four-wing plants are unisexual (either staminate or pistillate) but can flour. ual shrubs are either completely sta- Four-wing is regarded as one of minate (―male‖) or pistillate switch sexes from year to year de- pending on environmental conditions. the most important browse species (―female‖). Staminate plants only Photo by W. Fertig. in the western United States. In produce flowers that have anthers part this is due to the plant’s ability and shed pollen, and usually occur to thrive in semi-arid saline and in harsher microsites in the environ- Male plants that became fe- sandy basin habitats that cover ment. By contrast, pistillate plants males enjoyed a reproductive ad- much of the intermountain west, only produce seed and are not capa- vantage over other pistillate plants where Four-wing saltbush is often ble of self-fertilization. About 10% by being able to flower earlier in a one of the dominant shrub species. of plants in a population are good season. The result of all this More importantly, the foliage and monoecious and have separate male switching is that sex ratios in salt- seeds are palatable and highly nutri- and female flowers on the same in- bush populations can become tious. They are eaten with relish by dividual. skewed and sexes partially segre- cattle, sheep, and goats (though not What makes Four-wing saltbush gated along gradients of environ- horses) as well as deer, elk, ante- unusual is that the sex of any given mental quality, with females pre- lope, and bighorn sheep. Native plant is not genetically fixed (as in dominating in richer sites and able browsers consume the plant mostly most animals), but can change to become larger and produce in winter when saltbush is especially based on environmental cues. Re- more fruits. rich in carotene and protein. Ro- searchers have discovered that over Whether Native Americans paid dents, sage grouse, Gambel’s quail a seven year period nearly 40% of attention to the sexual intrigue of and other nesting birds take advan- the saltbush plants in a population Four-wing saltbush is unknown, tage of the shrub’s intricate branch- switched sexes, with as many as but tribes did make extensive use ing for shelter from predators and 20% doing so each year. Pistillate of saltbush. Before the advent of the elements. If saltbush rangeland plants were more likely to change steel axes made it easier to harvest is well-rested in the summer sides than staminate ones, especially trees, many Indians used brittle (allowing fruits to ripen and seed- in years following an unusually cold saltbush stems as fuel. Smoke lings to sprout) it can sustain rela- winter, drought, or after an atypi- from burning leaves was thought tively heavy grazing use in winter. cally heavy fruiting season. to revive those feeling weak or Too much browsing, however, can 10 Sego Lily September 2010 33 (5)

Celebrating Wildflowers: Plants of the Week

This essay is partly adapted from a biography of Four-wing saltbush prepared for the US Forest Service’s Celebrating Wildflower’s website (www.fs.fed.us/wildflowers/ plant-of-the-week/ index.shtml). Visit this website for a different native plant essay each week, and for other interesting and useful tidbits about pollinators, rare plants, wildflower events, and other happenings on US Forest Ser- vice lands.

lead to trampling of the brittle stems Above: Silvery foliage of Four-wing may take 3-4 years to become fully and eventually drain the seedbank. saltbush. The narrow leaves are an established, at which time they can Fortunately, Four-wing saltbush adaptation for water conservation in become self-perpetuating through readily germinates from seed if its semi-arid environment. Below: vegetative layering or seed. Four-winged pistillate bracts sur- given half a chance. Seeds ripen in I find Four-wing saltbush to be an round the one-seeded fruit. The late August and September and may bracts turn brownish-tan when ripe. enjoyable plant to have in my yard remain on the female plant into De- When planting, the wings should be in Kanab. It requires almost no cember. Several months of cold removed. Photos by Steve Hegji. maintenance, other than periodic stratification are required to induce pruning of dead branches at the germination in nature. Given some base of the plant (these old stems go protection from insects, rabbits, ro- 1/2 to 3/4 of an inch deep, de- in the woodstove). My covey of dents, and browsers, seedling four- pending on soil moisture and Gambel’s quail hang out all year in wings can grow rapidly, adding as coarseness. Seedlings require little the dense growth, and mule deer much as 18 inches of new growth. water and are tolerant of full sun browse on the foliage in winter. It is Because of its tolerance of cold and wind. They are best grown in also fun to keep tabs on which weather, drought, and poor soils dry sandy or slightly alkaline soil plants produce fruit each year, and Four-wing saltbush has become a in late fall, early winter, or very which ones will perform a sex- favorite species for use in reclama- early spring. Four-wing stands change next year! tion projects. Once established, the plant’s extensive root system is excellent for containing soil erosion. Four-wing is also becoming more popular as an accent plant in desert landscaping. In Utah, it is hardy in nearly all settings except high elevations and wetlands. Plants can be grown from stem cuttings or from seed. Stock is also increasingly available from commercial nurser- ies. When growing from seed, experts recommend an after-ripening period of up to 10 months, followed by 30- 50 days of cold stratification. Before planting, seeds should be soaked for two hours and the membranous wings removed (especially if being planted with a drill or mechanical seeder). Seeds should be planted

11 Utah Native Plant Society

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