Determining the Nativity of Plant Species

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Sego Lily September 2011 34 (5)

September 2011 (volume 34 number 5)

In this issue:

Determining the Nativity of Plant Species ...... 1, 4 Chapter News ...... 2 Bulletin Board ...... 3 Utah Botanica ...... 11 A New Sandwort for Utah Ode to the Staminode II

Cover: Common purslane (Portulaca oleracea) occurs world- wide in weedy habitats, but does that make it non-native? For the answer, see page 5. Photo by Al Schneider (www.swcolorado wildflowers.com).

Determining the Nativity of Plant Species

By Walter Fertig

Competition from invasive, non-native plant and animal species is considered the second greatest threat to biological diversity in the United States after habitat loss. An estimated 5000 plant species have been deliberately or accidentally introduced into the United States over the last five centuries. Of these, about 500 are considered serious agricultural pests. Pimental et al. (2000) estimate that reduced crop yields and expenses for the control of invasive weedy plants cost the US agricultural economy $26.4 billion per year. Invasive non-native species also impact native plant species and communities through direct competition, altered fire regimes, and reduced habitat quality and complexity. Control of non-natives has become an increasingly important task of land managers, siphoning away money and resources that might be better put towards other conservation issues. Given the importance of non-native plant species, surprisingly little attention has been paid to how nativity is actually determined (Willis & Birks 2006). Typically, plant taxa are considered native if they are found within their presumed area of evolutionary origin, or arrived without human intervention (Pysek et al. 2004). Non-natives, by con- trast, are known or suspected to have been brought to a new area by humans on purpose (to be grown for food, fiber, or ornamentation) or to have escaped by accident. Most introductions fail to become [continued on page 4]

Horticulture: Maggie Wolf Sego Lily Editor: Walter Fertig Important Plant Areas: Mindy ([email protected]). The deadline for Wheeler the November 2011 Sego Lily is 15 Invasive Weeds: Susan Fitts October 2011. Rare Plants: Walter Fertig Scholarship/Grants: Therese Meyer Copyright 2011 Utah Native Plant So- ciety. All Rights Reserved Chapters and Chapter Presidents Officers Cache: Michael Piep The Sego Lily is a publication of the Co-Presidents: Kipp Lee (Salt Lake Co.) Cedar City: Marguerite Smith Utah Native Plant Society, a 501(c)(3) & Walter Fertig (Kane Co) Escalante: Toni Wassenberg not-for-profit organization dedicated Vice President: vacant 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: Bill Gray quires permission (except where ex- Co) Southwestern/Bearclaw poppy: Mar- empted by statute) and must be cor- Board Chair: Larry Meyer (Salt Lake garet Malm rectly credited and cited. Articles, Co) Utah Valley: Celeste Kennard photographs and illustrations submitted to us remain the property of the UNPS Board: Jason Alexander (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), Therese Meyer Field Guide, and more, go to unps.org. publication. By submitting an article, (Salt Lake Co), Leila Shultz (Cache Co), Many thanks to Xmission for an implicit license is granted to print Dave Wallace (Cache Co.), Maggie Wolf sponsoring our website. the article in the newsletter or other (Salt Lake Co). For more information on UNPS: UNPS publications for reprint without Contact Bill King (801-582-0432) or permission (in print and electronic Committees Susan 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.

______Chapter News Manzanita (Kane County): For Fremont (Richfield area): The our August adventure, the chapter Fremont Chapter had a busy sum- visited the organic farm of John and mer! For June’s presentation, local Rhonda Flatberg along the Virgin photographer Kreig Rasmussen River, just north of Glendale. The shared tips on how to get great pic- Flatbergs have spent over two dec- tures. It was a great session, and ades slowly restoring a degraded wet everyone came away with a few meadow through rest and rotation more tricks for getting amazing pho- of a small herd of cattle. They have tos. In July, chapter members had a also constructed several solar- campout at the cabin up Seven Mile heated structures and greenhouses to enjoy native plants in their world. and are living completely off the Our members are also constantly grid. Attendees got to admire the working to maintain our native construction and also tromp about plant gardens at Fremont Indian the wet meadow and riparian vege- State Park and the Sevier County tation to admire the late summer Administration Building. On a sad flora. The Fremont Chapter is still look- note, one of our Our next event will be an evening ing to get back into the calendar- long-time chap- hike in Kanab Creek Canyon on the making business as a fundraiser—we ter members, premises of the Best Friends Animal still have some spots for various Elissa Jean Sanctuary on Tuesday, September chapters’ photos, but we’d like to get Wood Stevens, 13 from 6:30-dusk. Participants will the calendar into final production passed away on be invited to help locate new species AS SOON AS POSSIBLE in order to July 31, 2011, for the sanctuary plant list, currently allow holiday sales. For any ques- from complica- being developed by your chapter’s tions, or to submit your high- tions due to umbel leader. Please meet at the resolution photo and information, pancreatic can- Best Friends Welcome Center park- please contact Janett Warner at cer.—Lisa ing lot at 6:15 to then get going at [email protected] . White 6:30.—W. Fertig 2 Sego Lily September 2011 34 (5) Bulletin Board

UNPS Annual Meeting, 5 November 2011: The Salt Lake Chapter will be hosting the annual Utah Na- tive Plant Society members meeting and potluck on Saturday, November 5 at Red Butte Garden from 1-5 PM. Past Society President Bill Gray will address the group about his project to catalogue and relocate photo points in Utah and the southwest taken by pioneer botanist and noted curmudgeon Marcus E. Jones. Dr. Gray’s work has been funded, in part, through a small grant from UNPS. The meeting will also include the election of new society officers and board members and is an excellent opportunity to socialize with kindred native plant lovers.

Herbarium Meeting: The Intermountain Herbarium of Utah State University in Logan is hosting the annual regional meeting of western herbaria this fall from September 30-October 2. The meeting will include updates on the activities of various herbaria in the region and ongoing efforts to develop a regional consortium of collections and present herbarium data digitally on the internet. Dr. John Spence, expert bryologist from Glen Canyon National Rec- reation Area, will lead a field trip and workshop focusing on the identification of bryophytes. For additional information, contact Dr. Michael Piep, assistant curator of the Intermountain Herbarium ([email protected]).

Pollinator Conservation Talk, 12 September 2011: Bees and other pollinators are a fundamental part of our environment. There are more than 4000 native bee species in North America, and their work is something that touches us all through the food we eat, the clothes we wear, and the landscapes we enjoy, and yet their fu- ture is clouded by habitat loss, pesticide use, and introduced diseases. Join Mace Vaughan, Pollinator Program Director of the Xerces Society and Joint Pollinator Conservation Special- ist for the USDA’s Natural Resources Conservation Service, and coauthor of the newly released book, Attracting Na- tive Pollinators, to learn more about the fascinating lives of these insects and glean information about how you can care for these vital animals where you live. Whether you are an urban gardener, a suburban park manager, a working farmer, or just care about where your food comes from, this evening has something for you. The meeting will be at the Gore Auditorium of Westminster College in Salt Lake City on Monday, September 12 from 7:30 to 9 PM. Admis- sion is free and parking available under the athletic field off 1700 South.

Pollinator Workshops: The Xerces Society is also sponsoring two one-day courses in Pollinator Conserva- tion Planning in St. George (Weds., September 14) and Richfield (Thurs., September 15) by Mace Vaughan. These classes will equip conservationists, land managers, farm educators, and agricultural professionals with the latest science-based approaches to increasing crop security and reversing the trend of pollinator decline, especially in heavily managed agricultural landscapes. To register, click on the Xerces Society webpage (http://www.xerces.org/events/).

Salt Lake Chapter: The next meeting of the SL Chapter will focus on UFOs—unidentified flowering objects! Bill Gray will be the master of ceremonies, but needs your help with submissions of UFO images at least one week prior to the meeting. UFO night will be Wednesday, Sep- tember 7 at 7 PM at the Salt Lake City REI (3285 E 3300 South). Ad- mission is free and open to all.

Right: An Unidentified Flowering Ob- ject observed by Bill King in southern Nevada in late winter. Can you guess what this plant is or its planet of origin? Answer in the next issue.

3 Utah Native Plant Society

Determining the Nativity of Plant Species (Continued from page 1) established or naturalized without further human assistance and occur only in cultivated settings. Deci- sions about nativity have traditionally been made by the authors of regional floras, often without pro- viding a rationale. Determining the nativity of two small subsets of plant taxa is especially problematic. The first group includes species that are widely disjunct from the core area of their range. Disjunctions can arise naturally by long-distance dispersal or vicariance (contraction and isolation of once continuous ranges). In some instances, native Americans may have augmented the spread of species before European contact, as in the cases of maize (Zea mays), kid- ney beans, (Phaseolus vulgaris), and various squashes (Cucurbita spp.). The second group is com- posed of weedy species that commonly occur in disturbed habitats historical data (herbarium records, Above: Yellow bee plant (Cleome lutea and have been known since very floristic checklists, historic litera- or Peritoma lutea) is a native North early in the European colonization ture), geographic distribution, fre- American wildflower that commonly of the world. This group includes quency of known naturalization, occurs in weedy habitats and might, therefore, be mistaken for being non- some cosmopolitan weeds, such as genetic diversity, reproductive pat- purslane (Portulaca oleracea), com- native. Fossil remains of bee plant pol- tern (asexual or human-mediated len and seeds from pre-Columbian ar- mon reedgrass (Phragmites aus- vs. sexual or unmediated), and chaeological sites provides solid evi- tralis), and Kentucky bluegrass (Poa possible means of introduction. dence that Cleome species are truly na- pratensis), as well as such North Few of these are definitive by tive to North America. Photo by Al American ―weeds‖ as bug-seed themselves, but taken together Schneider (www.swcoloradowild (Corispermum spp.), ragweed they can provide evidence that a flowers.com). (Ambrosia spp.), and biennial given species is native or intro- wormwood (Artemisia biennis). duced. I will apply each of Webb’s Incorrectly determining the na- criteria to specific case studies question. Pollen assemblages are tive status of a species can have seri- from the putative non-native flora naturally biased towards species ous ramifications. Native disjunct of North America. My focus will that are wind pollinated and grow in species are often rare and of conser- primarily be on species affected by the vicinity of deposition basins, vation interest as these isolated European contact, rather than on such as bogs or sediment-filled populations are potential sources of ―archaeophytes‖ that were trans- lakes. Even if conditions for deposi- speciation and vulnerable to chance ported widely by humans in Pre- tion are favorable, the pollen of extinction events. If misidentified Columbian times. some species does not preserve well as non-native, these populations (such as Juncus) and identification might be subjected to weed control Fossil Evidence to species or even genera can be dif- efforts or ignored when they could Reliably dated and identified ficult in some taxonomic groups, benefit from benevolent steward- fossils provide the most concrete such as Poaceae and Chenopodi- ship. evidence that a plant species was aceae. Care must also be taken to To reduce this uncertainty, Webb present in an area prior to Euro- ensure that modern pollen has not (1985) recommended eight criteria pean contact, and thus native. Un- contaminated the sample. Macro- for assessing the nativity of a spe- fortunately, fossil and pollen evi- fossils (including preserved leaves, cies. These criteria include: fossil dence are lacking for the majority twigs, or floral parts) typically have evidence (pollen and macrofossils), of disjunct or weedy species in a much better taxonomic resolution, but are also rarely preserved, with 4 Sego Lily September 2011 34 (5) the exception of bog or lake habitats Common purslane (Portulaca have also been discovered from tidal and packrat middens. Of course the oleracea) is one of the world’s wetlands of Connecticut and the absence of a species from the fossil most widespread and abundant North Atlantic Coast that are 2000- record cannot be taken as proof that weedy species. Traditionally, it 4000 years old. Although present in it was not present. has been considered native to In- pre-Columbian times, fossil evi- Despite these limitations, fossils dia or Saharan Africa but intro- dence suggests that P. australis was have provided evidence that several duced elsewhere. The species has a minor component of coastal wet- commonly presumed non-native been known in North America lands until relatively modern times. species were, in fact, present in parts since Columbus documented it in Many other weedy native species of North America prior to Colum- Cuba and early 19th century ex- appear in pre-European fossil sites bus. Whether these native weeds plorers found it in Missouri and in North America and provide evi- subsequently spread to new parts of Colorado (Williams 2003). Re- dence for Native American agricul- the continent following the expan- cently, pollen and seed identifiable tural and land-disturbance activities sion of European culture is a sepa- as P. oleracea have been found at (Jackson 1997). Such species in- rate question that is not as easily archaeological sites in Kentucky, clude ragweeds (Ambrosia spp.), resolved. Louisiana, and Ontario that date beeplants (Cleome or Peritoma For nearly a century, monogra- from 600-3000 years ago. At spp.), spurges (Euphorbia and phers considered the North Ameri- Crawford Lake in Ontario, Chamaesyce spp.), globe-mallows can species in the genus Corisperm- purslane pollen and seed can be (Sphaeralcea spp.), and tidestromia um to be conspecific with Eurasian dated in annual lake sediments to (Tidestromia spp.). These species taxa based on their morphologic 1430-1489 AD and are found in are generally not considered adven- similarity and propensity for weedy, association with maize, sunflow- tive in North America because they disturbed habitats. Species of Co- ers, and cultivated bean fossils, are not found outside of the conti- rispermum are commonly called suggesting that purslane may have nent, or have only invaded other bugseeds due to their relatively been a food source for native peo- areas in recent years. large, flattened, and often membra- ples. Both the seeds and leaves of Fossil data have also been used to nous-margined fruits that superfi- purslane are edible and have the date the first appearance and subse- cially resemble a squashed bug. highest amounts of omega-3 fatty quent spread of non-native species Bugseed fruits are distinctive and acids and antioxidants of any that invaded North America follow- easy to identify. Betancourt et al. plants that have been tested. ing European contact. Pollen sam- (1981) discovered fossilized seeds of Carpetweed (Mollugo verticil- ples from Crawford Lake show that Corispermum from soil samples in lata) is often considered native to Rumex and Plantago pollen Alaska, Yukon, and the American the New World tropics, though (ostensibly representing R. ace- southwest and dated them using herbarium collections place it in tosella. P. lanceolata, and P. major) tandem accelerator mass spectrome- Ohio as early as 1828. Chapman first appear in the early 1820s, coin- try at ages between 4000 (New Mex- et al. (1974) report seeds of carpet- ciding with the establishment of ico) and 38,000 years (Alaska), pro- weed from an archaeological site at European agriculture in southern viding firm evidence that at least Icehouse Bottom, Tennessee dat- Canada (McAndrews 1988). Pollen some bugseed species were present ing to approximately 1170 years samples from other locations in in North America well before Euro- ago. Additional pre-Columbian North America show that Russian- pean settlement. Mosyakin (1995) carpetweed fossils are known from thistle (Salsola spp.), hemp re-assessed the taxonomy of the Louisiana and Alabama (Byrne (Cannabis sativa), hops (Humulus group and found subtle, but consis- and McAndrews 1975). Although lupulus), alfilaria (Erodium cicutar- tent morphological characters to related species of carpetweeds are ium), purple loosestrife (Lythrum recognize at least nine native North edible, M. verticillata is not widely salicaria), and various cereal American taxa. One of the new spe- used and probably grew as a grasses appeared or became abun- cies, C. navicula, is a narrow en- ―weed‖ in disturbed soil of Native dant only following white settle- demic of sand dunes in North Park, American garden plots. ment. Colorado and might reasonably be The common reed (Phragmites considered a species of conservation australis) is an aggressive invader Historical Data concern. Two other Eurasian spe- of wetland sites. Genetic evidence Webb (1985) notes that historical cies (C. hyssopifolium and C. suggests that there are both native records can show when a species nitidum) are still thought to be in- and non-native genotypes of the was first recorded in an area, but troduced in North America, but nei- species in North America cannot prove nativity. Evidence ther is apparently well established. (Saltonstall 2002). Hansen (1978) from ship manifests, seed cata- It should be noted that Thomas Nut- discovered macrofossil remains of logues, and published reports can tall, author of the premier early 19th P. australis from giant ground corroborate the introduction of most century flora of North America, con- sloth coprolites (fossil poop) in the crop and ornamental species to sidered at least one bugseed, C. Grand Canyon of Arizona that date North America (Mack 1990). Prob- americanum, to be native. to 40,000 years ago. Macrofossil lems remain proving the nativity of remains of Phragmites rhizomes disjunct or wide-ranging weedy spe-

5 Utah Native Plant Society cies, though early herbarium speci- Report of 1879 that they found side of the Teton Range in Wyoming mens, naturalist notebooks, or pub- dunes where ―… fine sand is blown in the early 1980s. Hartman et al. lished reports can at least provide a up upon the hillsides for a distance (1985) reported that the stand of first date of appearance. Combined of 500 to 600 feet‖. The only mature trees, located outside a with other evidence, historical data dunes fitting this description are popular campground, were appar- can make a persuasive argument for the ones between the Seminoe and ently native due to their size and potential nativity. Ferris Mountains, where P. hay- maturity. It was later determined Blowout penstemon (Penstemon denii has been found in recent sur- from local historians that the trees haydenii) is a native species long veys. Thus, Hayden’s collections were actually planted in the 1930s thought to be endemic to the Ne- of his namesake plant were proba- by the Civilian Conservation Corps, braska Sand Hills. In 1996, BLM bly taken in both Wyoming and and were thus not native to the area. range ecologist Frank Blomquist Nebraska, with Pennell having Location data from type speci- discovered a small population of mistakenly presumed the Wyo- mens can help inform whether a blowout penstemon in high, shifting ming reports were in error (Fertig species should be considered native. sand dunes at the west end of the 2001). Pineapple weed (Matricaria matri- Seminoe Mountains in Carbon Western larch (Larix occiden- carioides) occurs in disturbed habi- County, Wyoming, approximately talis) is a native species of north- tats over much of North America 300 km west of the nearest known western North America that was and is cited as non-native in most population in Nebraska. Immedi- discovered growing on the west eastern floras. The earliest speci- ately after this discovery, a promi- mens of this plant were taken by nent researcher in Nebraska ques- Lewis and Clark in June 1806 ―along tioned whether the Wyoming occur- Below: Russian-thistle (Salsola tra- the banks of the Kooskoosky‖ in rence was native or possibly intro- gus) is not a true thistle (Cirsium), but Clearwater County, Idaho (Phillips duced from seed he supplied to an did arrive in North America from 2003), suggesting it is native at least Eurasia. Once here, it hybridized unspecified Wyoming gardener in with other Salsola taxa to form a new west of the Rocky Mountains. Other the early 1980s. To complicate mat- North American form in California, S. weedy species considered native to ters, blowout penstemon is listed as ryanii. Does this make S. ryanii native western North America but adven- Endangered under the US Endan- to California? Photo by Al Schneider tive elsewhere include prostrate pig- gered Species Act. Legal protection (www.swcoloradowildflowers.com). weed (Amaranthus blitoides) with a under the Act would be limited only type from Iowa, windmill grass to the Nebraska populations if the (Chloris verticillata) first collected Wyoming plants were found to be by Nuttall from Arkansas, and introduced. blackcreeper sedge (Carex phaeo- Fortunately, historical data from cephala) with a type from San herbarium specimens and 19th cen- Diego, California. tury government research reports There are important exceptions to have helped shed light on this case. the rule that type localities infer lo- In his original description of the cal nativity. Aven Nelson described species, Watson (1891) reported P. Salsola pestifer from western North haydenii in Wyoming based on a America under the mistaken as- specimen collected by Ferdinand sumption that the Linnaean name S. Hayden from ―the Laramie Moun- tragus was misapplied. Salsola pes- tains‖ deposited at Harvard’s Gray tifer is now considered synonymous Herbarium. Unfortunately, Hayden with S. tragus and accepted as ad- did not include a collection number ventive from southern Europe. or date on the specimen label. Fran- Other Salsola taxa have been de- cis Pennell believed this specimen scribed from North America, includ- was a duplicate of another unnum- ing the recently named S. ryanii bered and undated Hayden collec- from California, which is believed to tion of P. haydenii labeled ―Loup be a recently derived fertile hybrid Fork‖ collected during an 1857 expe- between S. tragus and S. australis dition in the Nebraska Sand Hills (Hrusa and Gaskin 2008). Whether and deposited at the Missouri Bo- S. ryanii should be considered na- tanical Garden. Hayden and his tive presents an interesting conun- team of surveyors and geologists drum. were in Wyoming in August 1877 At least one putative native spe- and crossed Sandy Creek Pass along cies has a foreign type locality. Bi- the west side of the Seminoe Moun- ennial wormwood (Artemisia bien- tains on the 28th while traveling be- nis), a weedy forb often found in tween Rawlins and Casper. Hayden disturbed habitats, is usually re- notes in his Eleventh Annual garded as native to western North

6 Sego Lily September 2011 34 (5)

America but introduced elsewhere. native species if assessed only on ing some authorities to consider it Confusion over this plant’s nativity their habitat preferences. Many introduced. Mossgrass has since stems from the type specimen being native ―weeds‖ were originally been documented from scattered taken from a garden plant grown in adapted to disturbed sites such as sites in Washington, southern Brit- New Zealand. The source of the buffalo wallows, landslides, ish Columbia, and most recently in seed for this plant is thought to have burned areas, shifting river the Northwest Territories. All popu- come from northwestern North courses, and other early seral habi- lations have been associated with America. An unusual, multi- tats and have prospered over the ephemeral wetlands, islands, or branched form of this species does last three centuries following shorelines with fluctuating water have a type from North America. It European colonization. Such spe- levels. Catling (2009) argues that was first collected and named by cies have been called ―expansive‖ the species should be considered a Robert Dorn from Sweetwater by Pysek et al. (2004) to differenti- rare native because of its restriction County, Wyoming in 1980. Ironi- ate them from non-indigenous to unusual natural habitats shared cally, the Wyoming population has invaders. Fossil evidence can cor- by other rare species, such as the not been relocated since (and the roborate the post-contact expan- Columbia River endemic Columbian type locality significantly altered by sion or increase in abundance of yellowcress (Rorippa columbiae). construction of a power plant), but such native weeds as ragweeds the variety is still known from Boul- (Ambrosia spp.) or marsh-elder Geographic Distribution der Mountain in southern Utah. (Iva xanthifolia) (McAndrews Plant species with localized There is a possibility that the Wyo- 1988). Other common expansive ranges (endemics) or widespread ming population was established species include small-flowered and continuous distributions may be naturally by long-distance dispersal butterfly-plant (Gaura parvi- invasive or expansive at the leading from Utah or through human facili- flora), leathery knotweed (Poly- margins of their range, but are gen- tation but failed to persist due to the gonum achoreum), buffalo-bur erally not considered introduced marked differences in habitat (Solanum rostratum) and cut-leaf unless they somehow jump a large (Fertig 2008). nightshade (S. triflorum). geographic barrier, i.e. exhibit a sig- One species commonly pre- nificant disjunction. Range disjunc- Habitat sumed to be non-native is garden tion is a relatively uncommon phe- Webb (1985) notes that non- sorrel (Rumex acetosa) based on nomenon in most floristic regions. native plants tend to grow in hu- its long history of use as an edible Globally, Thorne (1972) estimated man-influenced habitats (including species and its typical distribution the percentage of disjunct species in cultivated gardens, agricultural in weedy sites in North America a typical flora to be about 1% (by fields, and disturbed sites), while and its native Europe. In the comparison, about 2% of the flora of native species are more likely to oc- northern Rocky Mountains, how- Utah consists of disjunct taxa and cupy ―natural‖ habitats that are not ever, R. acetosa is found in just 4% of the flora of Wyoming). dominated by humans. Some intro- ―natural‖ habitats such as alpine Several case studies have already duced crop and ornamental species, and subalpine meadows, fellfields, been discussed that illustrate the such as hollyhock (Alcea rosea), and coniferous forests. This habi- difficulties in differentiating be- asparagus (Asparagus officinalis), tat preference has led some re- tween naturally occurring disjunc- and white poplar (Populus alba) searchers to question whether tions (arising by vicariance or long- have barely escaped from cultiva- these populations might not actu- distance dispersal) and those that tion, and tend to occur only in dis- ally be native. Mosyakin (2005) are human-mediated. Disjunct spe- turbed sites near human dwellings. presents morphological evidence cies frequently co-occur in similar Many others have naturalized and that the high elevation material habitats or geographic areas, sug- can spread into both disturbed and from North America should be gesting that such patterns represent relatively pristine sites. Examples of recognized as a distinct, native natural phenomena. An unusually this latter group include some of our species called R. lapponicus. high concentration of disjunct spe- more pernicious weed species, like Native disjunct species that oc- cies from the taiga and deciduous cheatgrass (Bromus tectorum), salt- cupy early successional or ephem- forests of northern Canada and east- cedar (Tamarix chinensis), and eral wetland habitats are also fre- ern North America occur together in spotted knapweed (Centaurea quently mistaken for non-native the Black Hills of Wyoming and maculosa), as well as widespread species. One such species is moss- South Dakota. Thermal wetlands in species such as Kentucky bluegrass grass (Coleanthus subtilis), an in- Yellowstone National Park support a (Poa pratensis), redtop (Agrostis conspicuous, annual of vernal wet- small suite of disjuncts from compa- stolonifera), and red clover lands and gravel bars at scattered rable areas of Kamchatka (Agrostis (Trifolium pratense) that have es- locations in central Europe, north- pauzhetica) and the New World sentially become integrated into ern Asia, and northwestern North tropics (Myriophyllum quitense) natural systems and are not consid- America. The first record of this (Couch and Nelson 1988; Tercek et ered noxious. species in North America was al. 2003). Several groups of native plant made on an island in the Columbia species can be confused with non- River of Oregon in 1880, prompt-

7 Utah Native Plant Society

Frequency of Known Holmgren and Holmgren (2009) non-native haplotypes in North Naturalization hypothesized that the plant was America. Recent taxonomic work Species that have been intro- introduced and spread along the corroborates these findings and duced in one area of the world may transcontinental railroad, though three subspecies are now recognized be prone to invasion in other geo- all known S. linifolia in the west in the continent’s flora. These find- graphic locations of similar climate. have only been found since the ings are particularly important as This can be a useful clue to demon- mid 1980s. the introduced form is highly inva- strate non-native status. For exam- sive and has largely displaced native ple, dozens of introduced, noxious Genetics and Phylogeography ssp. americanus over much of the weed species in North America have Research on the degree of ge- eastern United States. also become successfully established netic differentiation between Genetic studies have helped re- and troublesome in temperate areas populations can be a useful tool to solve a long-standing controversy of southern Africa and Australia, reconstruct the evolutionary his- regarding the nativity of flaxleaf including Canada thistle (Cirsium tory of a species (phylogeography) whitepuff (Oligomeris linifolia) in arvense, ironically called Califor- or infer the age and origin of dis- the deserts of the southwestern nian thistle in Australia), bull thistle junct occurrences. Genetic studies United States and adjacent Mexico. (C. vulgare), Scotch thistle (Ono- have been used to determine the Although documented by Thomas pordum acanthium), tree-of-heaven region of origin for different geno- Nuttall in the early 1800s near San (Ailanthus alitissima), Scotch types of invasive weeds, such as Diego (and described by him as a broom (Cytissus scoparius), and cheatgrass. Such work can benefit distinct species), flaxleaf whitepuff horehound (Marrubium vulgare) resource managers searching for has usually been considered an in- (Parsons and Cuthbertson 2001). specific biological control species troduction from desert regions of Rapidity of spread can be another that might be most effective. Ge- North Africa and the Middle East. clue to suggest that a species has nome analysis has also shown that Recent genome analysis has shown been introduced. Cheatgrass be- non-native species may hybridize that significant differentiation has came established in British Colum- outside of their natal region to cre- occurred between North American bia in 1889 but by the 1930s was ate novel genotypes, as has been and Old World populations consis- widespread and dominant over most demonstrated with salt-cedar tent with establishment via a long- of western North America (Novak (Tamarix spp.) in western North distance dispersal event during the and Mack 2001). Russian-thistle America. As many as 12 of the 54 Pleistocene, rather than following and Halogeton (Halogeton glom- salt-cedar species native to Eurasia European colonization (Martin- eratus) had similarly rapid spreads and Africa may have been im- Bravo et al. 2009). A similar pattern across disturbed western rangelands ported to North America as orna- of disjunction has been shown with following establishment in the 1870s mentals and for bank stabilization, Mohave groundsel (Senecio mo- and early 1900s, respectively (Young but only two (T. chinensis and T. havensis) of desert areas of North et al. 1999). More recently, bur but- ramosissima) have become par- America and its close relative yellow tercup (Ceratocephala testiculata or ticularly invasive (Schaal et al. groundsel (S. flavus) of the Middle Ranunculus testiculatus) has 2003). Apparently, these two taxa East and between North American marched across the southwest fol- do not ordinarily cross in their na- desert Indian-wheat (Plantago insu- lowing an apparent first establish- tive range but hybridize readily in laris) and Saharan-Arabian ovate ment near Salt Lake City in 1932 North America. Genetic studies Indian-wheat (P. ovata) (Coleman (Barkworth 1982). suggest that the hybrid is the most et al. 2001; Stebbins and Day 1967). Not all introduced species, how- widespread form of salt-cedar In the case of these latter species ever, disperse widely or rapidly. In across western North America. pairs, sufficient morphologic and fact, non-native species are often Whether novel taxa derived from genetic divergence have occurred quite rare. For example, in Wyo- two introduced parents should be since the Pleistocene so that North ming, 38% of non-native species are considered ―native‖ in their coun- American taxa are now considered known from 5 or fewer extant popu- try of origin remains controversial separate, though sibling, species. lations and nearly 2/3 are known (Webb 1985). from less than 20 populations. Saltonstall (2002) used DNA Reproductive Pattern ―Rare‖ exotics can be potentially analysis to identify different ge- A high percentage of non-native difficult to recognize if they occur in netic strains of common reed. As species are annuals with ruderal life natural habitats rather than dis- previously discussed, macrofossil history characteristics that predis- turbed areas. evidence has shown that Phrag- pose them for successful invasion of One such exotic is Pin-leaf seep- mites australis was native to disturbed habitats. Several non- weed (Suaeda linifolia), known North America prior to European native species also reproduce by from scattered locations in Wyo- contact. Ecologists have debated apomixis (dandelion, Taraxacum ming, Nevada, and Utah, often in whether the species was also sec- officinale), asexual bulblets muddy riverbank habitats. This spe- ondarily introduced from Eurasia. (bulbous bluegrass, Poa bulbosa), or cies is indigenous to central Asia The genetic studies confirm the they spread vegetatively (mossy where it is not considered invasive. presence of multiple native and stonecrop, Sedum acre). Weedy

8 Sego Lily September 2011 34 (5) natives, however, can also occupy of the panicle branches and are rec- the ruderal niche and reproduce by ognized as separate subspecies apomixis (various Crepis spp.), (under P. pratensis) in the recent bulblets (alpine bistort, Polygonum Flora of North America treatment. bistortoides), or vegetative growth Historical data place P. pratensis in (numerous rhizomatous perennial Canada as early as 1749, where it grasses). Thus reproductive pat- was collected by Linnaeus’ student terns by themselves cannot reliably Peter Kalm. Traditionally, Kentucky confirm nativity. bluegrass is presumed to have been Webb noted that some intro- spread by early colonists along the duced species are incapable of re- Atlantic seaboard and the Missis- production without human assis- sippi drainage. Anecdotal evidence tance. An example of such a plant is suggests that Native Americans had maize, which occurs infrequently as no name for the species. Fossil data a waif in roadside and agricultural have not been of help, as grass pol- settings, but does not become natu- len and macrofossils are notoriously ralized because it cannot success- difficult to determine to species or fully reseed without human inter- genus. Genetic analysis would be vention. useful to resolve the taxonomy and clarify phylogeography, but unfortu- Means of Introduction nately have not been conducted on By definition, introduced species P. agassizensis. In the absence of require a means of introduction to a additional data, the case remains new location. Knowledge of the Above: Alfilaria (Erodium cicutar- unresolved. means of introduction is often a ium): historical and fossil evidence I will close with one final, enig- matter of the historical record, and provide conflicting evidence that this matic case. Alfilaria or crane’s-bill several cases were discussed previ- common weedy species might actually (Erodium cicutarium) has tradition- ously. Additional examples include be native to North America. Photo by ally been presumed to be introduced species that have escaped from gar- Al Schneider (www.swcolorado due to its abundance in weedy, dis- wildflowers.com). dens (Menzies’ baby blue-eyes, turbed habitats and its distribution Nemophila menziesii, or Missouri like blowout penstemon or white in Europe. Historical and pa- evening-primrose, Oenothera mac- spruce (Picea glauca) do not have leobotanical evidence appear to be rocarpa), plants used in highway to be explained by human inter- contradictory, however. Erodium beautification or soil stabilization vention or vicariance. cicutarium (Palmer’s penstemon, Penstemon was documented as early as 1820 by palmeri or cultivated flax, Linum Unresolved Cases the Long expedition in Colorado, usitatissimum), or intentionally in- For the vast majority of species well before the area was widely set- troduced for crops or livestock for- determining nativity is simple and tled and converted to ranching and age (smooth brome, Bromus iner- straight forward. In more prob- farming (Williams 2003). Stott et mis or nigerseed, Guizotia abys- lematic cases, researchers should al. (1998) present pollen evidence sinica). follow the advice of Webb (1985), that E. cicutarium was present in Disjunctions by native species Pysek et al. (2004), and Willis and the Santa Barbara area of California have often been confused with intro- Burks (2006) and consider multi- before Spanish missionaries arrived ductions by humans, in part due to ple lines of evidence to assign na- in 1769. But other pollen studies uncertainty over mechanisms facili- tivity. Authors of state or regional show that the species was absent in tating long-distance dispersal. Pa- floras should explicitly state their southern California before the set- leoecological evidence has long assumptions to make the process tlement era (Cole and Wahl 1999). shown that plants migrated at ex- of assigning native status more Could alfilaria have been present as ceedingly fast rates in response to transparent and defensable. an uncommon native weed (like changes in climate, despite having Despite our best efforts, some purslane and carpetweed) before limited dispersability: the so-called species remain problematic. One European contact? ―Reid’s Paradox‖. Clark et al. (1998) of the best known examples is Knowing whether a plant spe- provide theoretical and empirical Kentucky bluegrass (Poa pratensis cies is native or introduced is less evidence that such rapid migrations or P. agassizensis). Authors dis- clear-cut than it might appear. are possible if populations expand agree whether populations from Many disciplines, ranging from pa- through jumps rather than a broad, the Rocky Mountains and Great leobotany to genetics, play an im- advancing front. Such jumps are Plains are native (P. agassizensis) portant role in determining nativity. possible by the infrequent success of or introduced from Europe (P. Conservationists need to get such chance, long-distance dispersal pratensis). The two taxa have sub- determinations correct, lest we ex- events. Viewed in this light, disjunct tle morphological differences in pend time and resources on species distributions of large-seeded species leaf pubescence and the roughness protection or eradication that are unwise or unnecessary. 9 Utah Native Plant Society

References: Pimental, D., L. Lach, R. Zuniga, and Barkworth, M.E. 1982. Bur buttercup. D. Morrison. 2000. Environmental A weedy immigrant. Utah Science 43:6- and economic costs of nonindigenous 9. species in the United States. Biosci- Betancourt, J.L., A. Long, D.J. Dona- ence 50(1):53-65. hue, A.J.T. Hull, and T.H. Zabel. 1984. Pysek, P., D.M. Richardson, M. Re- Pre-Columbian age for North American jmanek, G.L. Webster, M. Williamson, Corispermum L. (Chenopodiaceae) con- and J. Kirschner. 2004. Alien plants firmed by accelerator radiocarbon dat- in checklists and floras: towards better ing. Nature 311:653-655. communication between taxonomists Byrne, R. and J.H. McAndrews. 1975. and ecologists. Taxon 53(1):131-143. Pre-Columbian purslane (Portulaca Saltonstall, K. 2002. Cryptic inva- oleracea L.) in the new world. Nature sion by a non-native genotype of the 253:726-727. common reed, Phragmites australis, Catling, P.M. 2009. Coleanthus sub- into North America. Proceedings Na- tilis (Poaceae), new to Northwest Terri- tional Academy Science, USA 99:2445- tories and its status in North America. 2449. Rhodora 111(945):109-119. Schaal, B.A., J.F. Gaskin, and A.L. Chapman, J., R.B. Stewart, and R.A. Caicedo. 2003. Phylogeography, hap- Yarnell. 1974. Archaeological evidence lophyte trees, and invasive plant spe- for precolumbian introduction of Portu- Above: Chicory (Cichorium intybus): cies. Journal of Heredity 94(3):197- laca oleracea and Mollugo verticillata Historical records confirm that this 204. into eastern North America. coffee-substitute was intentionally Stebbins, G.L. and A. Day. 1967. Clark, J.S., C. Fastie, G. Hurtt, S.T. introduced to North America. Photo Cytogenetic evidence for long contin- Jackson, C. Johnson, G.A. King, M. by Al Schneider (www.swcolorado ued stability in the genus Plantago. Lewis, J. Lynch, S. Pacala, C. Prentice, wildflowers.com). Evolution 21:409-428. E.W. Schupp, T. Webb III, and P. Wy- Stott, P., S. Mensing, and R. Byrne. ckoff. 1998. Reid’s paradox of rapid Mack, R.N. 1990. Catalog of woes. 1998. Pre-mission invasion of plant migration. Bioscience 48(1):13-24. Natural History 99(3):44-53. Erodium cicutarium in California. Cole, K.L. and E. Wahl. 1999. A late Martín-Bravo, S., P. Vargas, and M. Journal of Biogeography 25(4):757- Holocene paleoecological record from Luceño. 2009. Is Oligomeris 762. Torrey Pines State Reserve, California. (Resedaceae) indigineous to North Tercek, M.T., D.P. Hauber, and S.P. Quaternary Research 53:341-351. America? Molecular evidence for a Darwin. 2003. Genetic and historical Coleman, M., D.G. Forbes, and R.J. natural colonization from the Old relationships among geothermally Abbott. 2001. A new subspecies of Se- World. American Journal of Botany adapted (bentgrass) of North America necio mohavensis (Compositae) reveals 96(2):507-518. and Kamchatka: Evidence for a previ- old-new world species disjunction. Ed- McAndrews, J.H. 1988. Human ously unrecognized, thermally adapted inburgh Journal Botany 58(3):389-403. disturbance of North American forests taxon. American Journal of Botany Couch, R. and E. Nelson. 1988. Myrio- and grasslands: The fossil pollen re- 90:1306-1312. phyllum quitense (Halorag-aceae) in the cord. In: Huntley, B. and T. Webb III, Thorne, R. F. 1972. Major disjunc- United States. Brittonia 40(1 eds. Vegetation History. Kluwer Aca- tions in the geographic ranges of seed Fertig, W. 2001. 2000 survey for demic Publishers, Utrecht, Nether- plants. Quarterly Review of Biology 47 blowout penstemon (Penstemon hay- lands. Pp. 673-697. (4):365-411. denii) in Wyoming. Wyoming Natural Mosyakin, S.L. 1995. New taxa of Watson, S. 1891. Penstemon hay- Diversity Database, University of Wyo- Corispermum L. (Chenopodiaceae), denii. Botanical Gazette 16(11):311. ming, Laramie, WY. 40 pp. with preliminary comments of the Webb, D.A. 1985. What are the cri- Fertig, W. 2008. A new Artemisia for taxonomy of the genus in North Amer- teria for presuming native status? Utah? Sego Lily 31(4):9. ica. Novon 5:340-353. Watsonia 15:231-236. Hansen, R.M. 1978. Shasta ground Mosyakin, S.L. 2005. Rumex. In: Williams, R.L. 2003. ―A Region of sloth food habits, Rampart Cave, Ari- Flora of North America Editorial Com- Astonishing Beauty‖: The Botanical zona. Paleobiology 4(3):302-319. mittee. Volume 5 Magnoliophyta: Exploration of the Rocky Mountains. Hartman, R.L., B.E. Nelson, and K.H. Caryophyllidae, part 2. Flora of North Roberts Rinehart, Lanham, MD. 209 Dueholm. 1985. Noteworthy collections: America North of Mexico. Oxford pp. Wyoming. Madrono 32(2):125. University Press, New York. Pp 489- Willis, K.J. and H.J.B. Birks. 2006. Holmgren, N. and P. Holmgren. 2009. 533. What is natural? The need for a long Suaeda linifolia (Chenopodi-aceae) in Novak, S.J. and R.N. Mack. 2001. term perspective in biodiversity con- Utah. Sego Lily 32(2):10-11. Tracing plant introduction and spread: servation. Science 314:1261-1265. Hrusa, G.F. and J.F. Gaskin. 2008. Genetic evidence from Bromus tecto- Young, J.A., P.C. Martinelli, R.E. The Salsola tragus complex in Califor- rum (cheatgrass). Bioscience 51 Eckert, Jr., and R.A. Evans. 1999. nia (Chenopodiaceae): Characterization (2):114-122. Russian-thistle – a preview to the and status of Salsola australis and the Parsons, W.T. and E.G. Cuthbertson. spread of Halogeton. Pp 9-13. In: th autochthonous allopolyploid Salsola 2001. Noxious Weeds of Australia, Halogeton: A history of mid-20 cen- ryanii sp. nov. Madrono 55(2):113-131. second edition. Csiro Publ., Colling- tury range conservation in the inter- Jackson, S.T. 1997. Documenting wood, Australia. 436 pp. mountain area. Miscellaneous publi- natural and human-caused plant inva- Phillips, H.W. 2003. Plants of the cation US Department of Agriculture # sions using paleoecological methods. In: Lewis and Clark Expedition. Moun- 1553, Washington, DC. J.O. Luken and J.W. Thieret (eds.). As- tain Press Publ., Missoula, MT. 278 sessment and Management of Plant In- pp. vasions. Springer-Verlag. Pp. 37-55. 10 Sego Lily September 2011 34 (5) Utah Botanica Odds and Ends from Utah Botany

A New Sandwort for Utah: Ode to the Staminode, In recent years taxonomists have Part II: In the July issue of the resurrected the genus Eremogone Sego Lily, Peter Lesica answered for species with narrowly linear the age-old question of ―What leaves and 6-toothed capsules that good is a sterile staminode‖ in the were formerly included in the genus genus Penstemon Scrophulari- Arenaria (sandworts) of the pink aceae or Plantaginaceae*). Alert family (Caryophyllaceae). While reader and retired pollination bi- working on their treatment of ologist Vince Tepedino noted Eremogone for the upcoming final some recent papers that shed ad- volume of the Intermountain Flora, ditional light on why beardtongues Noel and Pat Holmgren of the New have a beardtongue. York Botanical Garden examined In the case of the Blowout pen- hundreds of specimens in the Erem- stemon (P. haydenii, also dis- Above: Blowout penstemon (Penstemon haydenii) from sand dunes of central ogone kingii/fendleri complex and cussed on page 6 of this issue), the became convinced that collections Wyoming. Photo by W. Fertig. removal of the staminode had dif- traditionally identified as Arenaria ferent effects on pollinating bees kingii var. uintahensis were suffi- depending on their size. Larger marked reduction in the amount of ciently distinct to warrant recogni- insects, such as bumblebees tion as a full species. Unfortunately, pollen deposited on the stigma by (Bombus) and masarid wasps visiting bees, but did not seem to the name uintahensis turned out to (Pseudomasaris vespoides) were be unusable, because the type collec- affect total pollen removal. How- completely unaffected by elimina- ever, the researchers found that tion of the variety was actually a tion of the staminode. The native specimen of A. kingii var. glabres- both large and small-bodied bees bee Osmia brevis, which normally removed much less pollen when the cens. With no legitimate name straddles the staminode to reach available, the Holmgrens were staminodes were gone, while me- the pollen-bearing anthers, com- dium-sized bees were apparently forced to come up with a new name pensated in altered flowers by of their own. unaffected (accounting for the total merely stretching its legs to still amount of pollen removal appearing Enter ―Arnow’s sandwort‖ or gain access to pollen. Only the Eremogone loisiae, named to com- unaffected). smaller sweat bees (Dialictus prui- Interestingly, pioneer ecologist memorate longtime Garrett Herbar- nosus) were befuddled by the lack ium curator, Lois Arnow. In addi- Frederic Clements conducted ex- of a staminode and frequently left perimental work with staminode tion to her work building the Uni- the flowers without picking up any versity of Utah’s herbarium, Arnow removal in the 1920s and also found pollen. Overall, fruit set and seeds that the absence of the bearded also wrote the Flora of the Central per fruit differed little between P. Wasatch Front, Utah and resolved tongue affected pollinators differ- haydenii flowers with or without ently depending on their size - W. many taxonomic problems in the staminodes. In those penstemon Poaceae (grass family). Fertig populations in which the larger Eremogone loisiae can be distin- pollinators were common, the sta- References: guished from E. fendleri by its more minode of P. haydenii is probably Clements, F.E. and F.L. Long. 1923. ovate sepals and from E. kingii var. truly vestigial, but where small Experimental pollination: An outline of glabrescens by its larger sepals (4-7 pollinators are important, the sta- the ecology of flowers and insects. Car- mm long vs. 2.2-5 mm). Arnow’s minode is still essential for sexual negie Institue of Washington Publ. No,. sandwort is most similar to E. east- reproduction (and thus evolution- 336, Washington, DC. woodiae, but has longer and more arily adaptive). Dieringer, G. and L. Cabrera R. 2002. The interaction between pollinator size flexuous basal leaves (2.5-5 cm Dieringer and Cabrera R. long), and larger styles and seeds. and the bristle staminode of Penstemon (2002) also found a link between digitalis (Scrophulariaceae). American This new species is restricted to the pollinator size and staminode Journal of Botany 89(6):991-997. Wasatch Range and vicinity, from function in Foxglove beardtongue Hawk, J.L. and V.J. Tepedino. 2007. southern Idaho to central Utah (Penstemon digitalis) of open The effect of staminode removal on fe- (Cache and Rich to Sanpete coun- woods and prairies of eastern male reproductive success in a Wyoming ties) and Uinta County, Wyoming. North America. Removal of the population of the Endangered Blowout bristly staminode resulted in a penstemon, Penstemon haydenii Holmgren, N.H. and P.K. Holmgren. (Scrophulariaceae). Madrono 54(1):22- 26. 2011. A new species of Eremogone (Caryophyllaceae) from northern Utah Lesica, P. 2011. What good is a sterile *I still can’t get used to that change. and southeastern Idaho, USA. Brittonia staminode? Sego Lily 34(4):9. 63(1):1-6. 11 Utah Native Plant Society

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