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The Plant Press THE ARIZONA NATIVE PLANT SOCIETY Volume 37, Number 2 Fall 2014 In this Issue: Interdisciplinary Botany 1 Arizona Native Food Plants for a Dry Future 6 Oaks, Cycads, Orchids, and Conservation in the Sierra Huérfana, Sonora 8 Special Soils and their Role in Plant Evolution, Distribution, and Regional Diversity 13 Characterization of the Selaginella arizonica x eremophila Hybrid Zone in Southern Arizona & Our Regular Features 2 President’s Note 11 Flora of North America Honors AZNPS 15 Who’s Who at AZNPS 16 Spotlight on a Native Plant: Mexican Tansyaster 17 Book Review: Trees of Western North America 18–19 Announcements ©2014 Arizona Native Plant Society. All rights reserved. continued next page Wild sunflower, Rio Rico, Arizona, 23 August 2014. Photo by Susan D. Carnahan. While learning some of the intricacies of Western Apache knowledge of food plants and historic landscapes, I wondered: How different would it be if they or other Native Americans had been able to teach invasive Euro-Americans about native food resources and greater respect for local ecosystems? The Sonoran Desert Region is rich in plant diversity, and rich in plants used for indigenous cuisine. Dozens of such wild plants are suitable for agricultural crops and home gardens. Fit the crop to the land, not the land to the crop: the concept includes no- tillage land use and native plants that have been wild-harvested since ancient times. The implications for energy and water conservation are huge. Some years ago I had the privilege of searching the world for new aridland/salt-tolerant food crops and now I find some of the best ones right here in our backyard. Among the Arizona Native Food Plants for a Dry Future by Richard Felger 1 1 University of Arizona Herbarium and Sky Island Alliance, Tucson.

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Page 1: The Plant Press - Arizona Native Plant Society · The Sonoran Desert Region is rich in plant diversity, and rich in plants used for indigenous cuisine. Dozens of such wild plants

The Plant PressTHE ARIZONA NATIVE PLANT SOCIETY

Volume 37, Number 2 Fall 2014

In this Issue:Interdisciplinary Botany1 Arizona Native Food Plants

for a Dry Future6 Oaks, Cycads, Orchids, and

Conservation in the SierraHuérfana, Sonora

8 Special Soils and their Rolein Plant Evolution,Distribution, and RegionalDiversity

13 Characterization of theSelaginella arizonica xeremophila Hybrid Zone inSouthern Arizona

& Our Regular Features2 President’s Note

11 Flora of North AmericaHonors AZNPS

15 Who’s Who at AZNPS16 Spotlight on a Native Plant:

Mexican Tansyaster17 Book Review: Trees of

Western North America

18–19 Announcements

©2014 Arizona Native PlantSociety. All rights reserved.

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Wild sunflower, Rio Rico, Arizona, 23 August 2014. Photo by Susan D. Carnahan.

While learning some of the intricacies of Western Apache knowledge of food plants andhistoric landscapes, I wondered: How different would it be if they or other NativeAmericans had been able to teach invasive Euro-Americans about native food resourcesand greater respect for local ecosystems?

The Sonoran Desert Region is rich in plant diversity, and rich in plants used forindigenous cuisine. Dozens of such wild plants are suitable for agricultural crops andhome gardens. Fit the crop to the land, not the land to the crop: the concept includes no-tillage land use and native plants that have been wild-harvested since ancient times. Theimplications for energy and water conservation are huge.

Some years ago I had the privilege of searching the world for new aridland/salt-tolerantfood crops and now I find some of the best ones right here in our backyard. Among the

Arizona Native Food Plants for a Dry Futureby Richard Felger1

1University of Arizona Herbarium and Sky Island Alliance, Tucson.

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President’s Note by Douglas Ripley [email protected]

2 The Plant Press Arizona Native Plant Society Fall 2014

As 2014 comes to a close I wish to thank all members fortheir fine support of the AZNPS during the past year. Ithink we can look back on the year with satisfaction for themany significant AZNPS accomplishments to which manyof you contributed. I am also very grateful to our Board ofDirectors, members of which have consistently providedsupport, encouragement, and new ideas for the successfulmanagement of the Society.

During the past year we continued to sponsor a wide rangeof activities and programs in support of native plantawareness and conservation. Habitat restoration projects,Citizen Science programs in support of the Plant AtlasProject of Arizona (PAPAZ), plant family taxonomyworkshops held in cooperation with the University ofArizona Herbarium, financial support for native plantresearch and publications through various chapter grantprograms, and our annual Arizona Botany Conference are afew of the more significant endeavors accomplished thispast year. We definitely plan on continuing, and possiblyexpanding, these activities in 2015. I am particularlyimpressed with the potential to expand our Citizen Scienceefforts by engaging more of our members, expanding ourcooperation with local universities, and possibly partneringdirectly with Arizona’s Natural Heritage Program andNatureServe, the non-profit conservation organizationwhose mission is to provide the scientific basis for effectiveconservation action. NatureServe and its network of naturalheritage programs and conservation data centers are theleading source for information about rare and endangeredspecies and threatened ecosystems and are actively seekingto partner with native plant societies to support theirCitizen Science efforts.

In 2015 we will hold the Arizona Botany Meeting incooperation with the Museum of Northern Arizona,Flagstaff. Please mark your calendar for 26-27 April 2015.

Full details regarding the theme and a call for papers will beforthcoming soon.

Although we have many worthwhile projects and activitiesplanned for 2015, we could certainly do more. I thereforeask that you consider expanding your participation in theSociety and to let us know of new efforts the Society couldinitiate or support.

The Plant Press continues to serve as our semiannualjournal for the dissemination of information on Arizona’snative flora. As such it offers an excellent forum forindividuals to publish the results of their botanical researchor other experiences with native plants. I ask all membersto please consider submitting native plant relatedinformation, results of research, or any other items ofbotanical interest for inclusion in future issues.

This issue of The Plant Press contains selected papers basedon presentations at the 2014 Arizona Botany Meeting, thetheme for which was “Interdisciplinary Botany: HowRelated Scientific Fields Support Our Understanding ofNative Plants.” Consequently these papers reflect a diversityof subjects, ranging from a discussion of the potential todevelop new native food plants; the results of aninvestigation of the evolutionary genetic processes ofhybridization and polyploidization among species of theprimitive vascular plant genus Selaginella in the SonoranDesert; and, a description of soils and their role in plantevolution, distribution and regional diversity from theUpper Verde River, Arizona. Finally, we present a report ofan April 2014 Madrean Archipelago BiodiversityAssessment (MABA) Expedition to the Sierra Huérfana,Sonora, Mexico, that resulted in over 1,000 biologicalobservations which helped provide documentation of theunique biodiversity of that important Sky Island.

a

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several hundred wild food plant species in the SonoranDesert Region, several dozen provided Native Americanswith staples. Wild regional staples include agaves, saguaroand organ pipe, mesquites, sunflowers, a richness of grasses,even native palms, and many more. Some of these wild foodplants are promising “new” food crops for home gardens aswell as large- and small-scale agriculture. Certain SouthAmerican mesquites, or algarrobas, are gaining majorsignificance in their native lands, and our native mesquitesare likewise gaining prominence as local food resources.

Driving from Tucson to New Mexico in late summer Inoticed the native sunflower, Helianthus annuus, along theroadside, mile after mile. A few weeks later I noticedabandoned farmland had become a jungle of thesesunflowers reaching above head height. Although no-tillagefarming usually implies use of perennials, this sunflower andsome of its relatives are annuals that can naturally reseedyear after year without plowing. These hardy plants thrive indisturbed habitats, although best at elevations above thedesert or in home gardens. The wild sunflower is lateripening, in fall after most summer crops have beenharvested. Native Americans made extensive use of the“seeds,” actually an achene, which includes the husk and itsenclosed seed.

This wild sunflower, native across North America, is theprogenitor of domesticated, agronomic varieties, with a verylarge and often single seed head, larger seeds, and differentcultivars bred for thinner shells and high seed oil. Russia andthe Ukraine produced 16 million tons of sunflower seeds lastyear—that’s 32 billion pounds. The wild sunflower differs inpart by having smaller but numerous flower heads, smallerachenes/seeds, and a much wider range of ecological

tolerance. Simple commercial and artisanal methods ofharvesting and cleaning are well known. In early October,New Mexico Gila River farmer Eric Leahy collected wildsunflowers from an abandoned field and showed us how toseparate the achenes from the chaff using screens, andfurther winnowing using a fan. Apaches and others groundthe whole achene, seed and husk together, for an esteemedfood resource. Some people boiled the achenes whole and,after cooling, the oil would be skimmed off, providingsunflower oil. Why not farm these plants? It would be morelike managing a sunflower-filled land.

The Sonoran Region native bounty extends to grasses, too.Civilizations co-evolved with grasses: wheat, rice, corn, oats,barley, sorghum, and a richness of others. But these familiarcrops are tropical and temperate-climate plants—requiringextensive irrigation in dry regions. They are also annuals,which require annual replanting and tillage (plowing). Whatif we developed arid-adapted no-tillage grain crops? Oneintriguing candidate is nipa (Distichlis palmeri) at theColorado River delta, one of the very few grasses endemic tothe Sonoran Desert. Nipa was a staple of the Cocopahpeople, thrives with pure seawater, and produces prodigiousamounts of grain comparable to that of early strains of rice.

Most grass grain (technically a caryopsis, a seed fused to thefruit case, or pericarp) are edible, although many are toodifficult to prepare or process. (However, avoid ergot-contaminated or otherwise infected grain, which will standout as discolored and much larger than uninfected grain.Also, ensure proper storage to preclude the growth of fungithat produce mycotoxins.) Many Sonoran region grassesmake up for being small by producing large quantities of

Arizona Native Food Plants for a Dry Future continued

Opposite: Wild sunflower, Rio Rico, Arizona, 23 August 2014. Photo by Susan D. Carnahan. Above, left: Eric Leahy harvesting wild sunflowersnear the Gila River in New Mexico, with a sheet over the hood of the pickup, 15 October 2014. Center: Initial screening of sunflower achenes(“seeds”) by Eric Leahy and Gregg Dugan, 15 October 2014. Right: Winnowing freshly screened sunflower achenes, 14 October 2014.

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grain and, like many arid land plants, pack a lot of energyinto their seeds. The Comcaac (Seri people) even harvestedthe minute grain of the six-weeks grama grasses (Boutelouaaristidoides and B. barbata var. barbata), which can carpetthe desert after summer rains.

For home gardens and agriculture, we turn to someinteresting perennial grasses. Sporobolus species are unusualamong grasses because the actual seed ripens free from thesurrounding pericarp and chaff (glumes, lemma, and palea),thus giving rise to the common name “dropseed grasses.”Perennial dropseed grasses, such as S. cryptandrus andespecially S. giganteus, have seeds so minute as to be likeflour. It is certainly possible that these seeds could be usedlike flour, without milling, because the surface-volume ratiois already very high.

Twila Cassadore, native food expert of the San CarlosApache Tribe tells us the grain of Panicum bulbosum(Zuloagaea bulbosa) is the easiest to harvest and best tastingof the wild grasses. The Apache name translates as RedGrass, a more enticing name than bulb panicgrass; let’s call itApache Red Grass. Indeed the ripening panicles are usuallyreddish and the freshly harvested grain has a pleasant taste.

Apache Red Grass is a large perennial grass native tomountains in the Southwest and northern Mexico. The grainis like that of the related Common Millet, Panicummiliaceum, albeit smaller. You can easily strip off the ripeRed Grass grain by hand into a paper bag. Like other grainsand seeds, it is essential to get them dry as soon aspossible—all moldy seeds and grains need to be avoided.You can prepare it as you would any millet or small grain.Like many wild grasses and millets, Apache Red Grass grainis gluten-free.

Some Apache Red Grass plants have bluish-green foliagewhile others have yellow-green or bright green foliage,making them attractive landscape plants as well as a tastyfood resource. Red Grass can be grown from seed ordivisions of the short rhizomes of established plants. To getgood seed set and ensure cross-pollination you should havemore than one clone. Plant it along a walkway, as an accentplant, or backdrop screen, or in an experimental plot forgrain production. The above-ground parts are frost-killedbut remain as fall color in your garden, or cut the above-ground part and this perennial will resprout in spring.

The coconut, oil palms, and date palm are among the moresignificant human food resources. The date, Phoenixdactylifera, is the desert palm of Egypt, but what about ourdesert palms? Washingtonia filifera, in desert oases andcanyons in Arizona, California, Nevada, and Baja California,is the only palm native to the western United States. Its onlyclose relative, W. robusta, is native to Sonora and the BajaCalifornia Peninsula. These fan palms are grown across theSonoran Desert Region and arid/subtropical regions aroundthe world. The fruit, produced in prodigious quantity, issimilar to that of the date palm but much smaller, often only6 mm long. Palm expert Don Hodel of the University ofCalifornia Cooperative Extension found that a single W.robusta produced approximately 572,000 fruits weighing 101kilograms (223 pounds). The Cahuillas of Palm Springs,California, made extensive use of the sweet fruit of W.filifera, fresh or dried, for beverages, or the whole fruit andseed ground into flour. Fine preserves and especiallybeverages might be the best modern-day applications, or aslocal food expert Carolyn Niethammer suggests, perhaps forbaked goods instead of sugar or corn syrup, or a source for agranulated natural sweetner. Try experimenting with thesefan palm fruits.

Palms require water, but like many other large monocots, theroot system is not extensive. You only need to provide waterclose to the trunk, and poor-quality water will suffice, suchas gray water, or water too alkaline or saline for other plants.You will need an extension pole, bucket lift, or some otherway to harvest high-hanging fruit.

Any new agronomic crop faces challenges — observations,experimenting, serendipity, perseverance, and creativity cangive results. Incentives for our own arid-adapted food crops

Arizona Native Food Plants for aDry Future continued

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Gregg Dugan harvesting Apache Red Grass, Pinos Altos Mountains,New Mexico, 10 September 2014. Photo by Jarrod Swackhamer.

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include adapting to a warming and drying climate,water shortages, energy conservation, moresophisticated consumers, and improved nutrition ineveryday diet.

aReferences

Bean, Lowell John and Katherine S. Saubel. 1972.Temalpakh: Cahuilla Indian knowledge and usage ofplants. Reprinted 2001, Malki-Ballena Press, Banning,CA.

Buffalo Bird Woman’s Garden Recounted by Maxi'diwiac(Buffalo Bird Woman) of the Hidatsa Indian Tribe,originally published as G.L. Wilson, 1917, Agriculture ofthe Hidatsa Indians: An Indian interpretation.University of Minnesota Studies in the Social Sciences 9.digital.library.upenn.edu/women/buffalo/garden/garden.html#III

Felger, Richard S. 2014. Perennial native grasses: Food for adry world. Newsletter of the Native Plant Society of NewMexico 39 (4): 1, 6–7, 10. www.npsnm.org

Hodgson, Wendy C. 2001. Food plants of the SonoranDesert. University of Arizona Press, Tucson.

Niethammer, Carolyn and Paul Mirocha. 2011. Cookingthe wild Southwest: Delicious recipes for desert plants.University of Arizona Press, Tucson.

Pearlstein, S.L., R.S. Felger, E.P. Glenn, J. Harrington, K.A.Al-Ghanemd, and S.G. Nelson. 2012. Nipa (Distichlispalmeri): A perennial grain crop for saltwater irrigation.Journal of Arid Environments 82: 60–70.

Acknowledgements

I thank the many friends and colleagues who have contributedto the new crops work, especially co-investigator Gregg GrahamDugan, as well as Elizabeth “Betsy” Arnold, Susan DavisCarnahan, Twila Cassadore, A.T. Cole, Alicia Edwards, AnnLane Hedlund, Donald R. Hodel, Kelly Kindscher, RussellKleinman, Eric Michael Leahy, Jim Malusa, Gary PaulNabhan, Carolyn Niethammer, William Norris, Seth Pilsk,Joseph “Joe” Ronald Runyan, Silke Schneider, Carolyn RuthSmith, Douglas Arthur Smith, Jarrod Swackhamer, James “Jim”Thomas Verrier, Caleb Weaver, and Robert Hazard Winston.For financial support I thank New Mexico Native Plant Society,Southwestern Foundation for Education and HistoricalPreservation, and Agricultural Development and PromotionFunds Program of New Mexico.

Arizona Native Food Plants for a Dry Future continued

Desert fan palm, Washingtonia filifera. Arizona StateUniversity, Tempe. Photo by Les Landrum (from SEINet).

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The southwestern most Sky Island in Sonora is 80 km westof Hermosillo, a city of about a million inhabitants. It iscalled the Sierra Mazatán by the residents of Mazatán (froma Nahuatl Indian word meaning “place of deer” – the sameas Mazatlán). But most of the mountain is in the ComunidadAgraria de Pueblo de Álamos, where it is Sierra Huérfana —or Orphan Mountain — a good name for a Sky Island! It issurrounded by tropical foothills thornscrub just east of theSonoran Desert, with oak woodland on top in a landscape ofwhitish granite boulders.

The flora of the Sierra Huérfana, with reliable summer rainsand granitic soils that hold moisture very well, is unique.Sanchez-E. et al. (in 2005) reported 357 species in apreliminary flora. Documentation of the flora is still inprogress and a more complete flora will be published soon.The oak woodland is dominated by Chihuahua andwillowleaf oaks, often festooned with ball moss (Tillandsiarecurvata). Various thornscrub plants, including papelío(Jatropha cordata), kapok/pochote (Ceiba acuminata), torotes(Bursera fagaroides, B. laxiflora), and tree ocotillo(Fouquieria macdougalii) reach their upper elevational limitsin this woodland.

In May 1957, Howard S. Gentry, the pioneering botanistwho worked in the Río Mayo Region of tropical southernSonora, collected a cycad in the Sierra Huérfana. In 1997, it

was described as Dioon sonorense, a species endemic toSonora and northern Sinaloa. Palma de la vírgen or peine(the serrated leaf edge looks like a comb) occurs in 5-6populations in Sonora, and has a Protegida status under the2010 Mexican endangered species law. There are 14 speciesof Dioon from Nicaragua northwest into Mexico. The SierraHuérfana D. sonorense are the northernmost cycads in theworld. During the Triassic and Jurassic geologic periods 200to 145 million years ago, cycad trees and shrubs along withconifers and ginkos dominated archaic dinosaur forests.Although cycads were once widely distributed over theEarth, including Alaska and Antarctica, the familyZamiaceae today only contains eight genera in Africa,Australia, and North and South America.

Other unusual plants in the Sierra Huérfana include theendemic Mazatán rock daisy (Perityle reinana, named inhonor of Ana Lilia Reina-Guererro), coral root orchids(Hexalectris spicata var. arizonica, H. warnockii, onlySonoran locations), a white morning glory/trompillo(Ipomoea scopulorum), and tiger flower (Tigidia pavonia).The red-flowered claret cup cacti in the Sierra Huérfana aremostly Echinocereus santaritensis var. bacanorensis, asouthern, summer-flowering variety of the Santa RitaMountains claret cup. There is also a record of E. klapperi, arare species only known from here and near the El Novilloreservoir.

Oaks, Cycads, Orchids, and Conservation in theSierra Huérfana, Sonoraby Thomas R. Van Devender1, J. Jesús Sánchez-Escalante2, and Ana L. Reina-Guerrero1

Left: A view of Sierra Mazatán. Right: Chihuahuan oak. Photos by Thomas R. Van Devender.

1 Sky Island Alliance, PO Box 41165, Tucson, AZ 85717. [email protected]. 2Departmento de InvestigacionesCientíficas y Tecnológicas, Universidad de Sonora, Av. Rosales y Niños Heroes s/n, Hermosillo, Sonora 83000, México.

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The Comisión Nacional de Áreas Naturales Protegidas(CONANP, the Mexican Park Service), is working onestablishing the Sierra Huérfana as a federally-protectednatural area. Pueblo de Álamos is a comunidad agraria, afederal designation that recognizes historical residence andland rights in an area. Comuneros use and manage the land,but cannot sell it. Otherwise the prime woodlands of SierraHuérfana, so close to Hermosillo in the Sonoran Desert,would have been developed long ago.

In April 2014, 55 botanists, entomologists, herpetologists,and ornithologists, as well as agency biologists,photographers, journalists, and college students from theUnited States and Mexico joined local residents on aMadrean Archipelago Biodiversity Assessment (MABA)Expedition to the Sierra Huérfana. Over 1,000 biologicalobservations were made and added to the MABA database

Above, left: Dioon sonorense, detail of serrated leaf edge. Photo by by Ana L. Reina-G. Center: Hexalectris spicata and Right: H. warnockii,coral root orchids. Photos by Michael F. Wilson. Below: Evening meeting of Pueblo de Álamos residents, CONANP personnel, and MABAparticipants to discuss the proposed Sierra Huérfana Protected Natural Area. Photo by James Rorabaugh.

(Madrean.org), providing documentation of the uniquebiodiversity of this Sky Island (Isla Serrana in Spanish). Thegroup’s exuberant appreciation of natural history wascontagious, and in July the residents of Pueblo de Álamosvoted in favor of the new Áreas Naturales Protegidas.

We thank the Greater Good Foundation for support for theMABA Expedition.

a

Above: White-flowered morning glory (left) and Tiger flower (center). Photos by Erik Enderson. Right: Bacanora claret cup cactus. Photoby Stephen L. Minter.

Oaks, Cycads, Orchids continued

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Geology and soils have been cited as second only to climateas the primary determinants of plant distribution anddiversity (Cain 1944). The stark contrast between plantcommunities of distinct geologic formations has longfascinated naturalists, ecologists and evolutionary biologists.Theophrastus (ca. 380-287 B.C.), the “Father of Botany” wasone of the first to record a singular vegetation type growingon limestone bedrock in ancient Greece. In some of the firstpublished articles on natural selection, Wallace (1858) spokeof the high degree of selection imposed on plants by theirsubstrata. Throughout the globe, managers and researchershave become increasingly interested in the effect of geologyon plant distribution, endemism, and regional diversity.

Infertile soils in particular have drawn attention. They oftenform islands with distinct plant communities, host disjunctspecies, and drive the evolution of rare and endemic species.Major examples include serpentine, which is low in nutrientsand high in toxic metals, and the highly basic and lownutrient limestones, dolostone (a magnesium limestone) andgypsum. Other examples include mine tailings, variousigneous and sedimentary rocks, sand, clay, and saline soils.On top of harsh chemistry, the physical nature of some ofthese soils can create stressful conditions, beingimpenetrable to plant roots, and steep or rocky.

Edaphic endemics have morphological and physiologicalfeatures that allow them to cope with harsh conditions.Among these are perennial growth form, small stature, lowspecific leaf area, high root to shoot ratio, low growth rates,stress-related leaf characteristics (glaucousness, pubescence,sclerophylly) and mutualistic relationships with soilorganisms. These traits allow them to tolerate drought andthrive with relatively low amounts of nutrients (Kruckeberg2004).

The presence of disjuncts and endemics on edaphic islandstell stories of the biogeographical and evolutionary history ofa region. Two main paths to endemism have been identified;paleoendemics are relicts of older, more widespread specieswhile neoendemics have diverged more recently.Throughout North America since the late Tertiary andPleistocene, plant migrations and radiations have trackeddramatic climatic and topographic shifts (Axelrod 1958,Betancourt et al. 1990). Over time the ranges of species lessfit for the changing climate retracted as they wereoutcompeted by the incoming dominant vegetation. Plantsthat were poor competitors but were able to survive oninfertile islands were released from competition and able topersist, while they were extirpated on surrounding soils.These refugial populations exist today as disjuncts,

Special Soils and their Role in Plant Evolution,Distribution, and Regional Diversity:Examples from the Upper Verde River, Arizonaby Francis S. Coburn1

Left: Polygala rusbyi, a central Arizona endemic growing on dolostone. Center: Upper Verde Riverflowing through Cambrian Tapeats sandstone below Devonian Martin dolostone. Right: Salviadorrii ssp. mearnsii, a Verde Valley and Upper Verde River endemic growing on dolostone. Photos byFrancis S. Coburn.

continued next page1 School of Life Sciences, Arizona State University

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populations at the periphery of their ranges and in the caseof plants that have been extirpated everywhere else,paleoendemics. Some populations that persisted on theedaphic islands became more adapted to the soilenvironment, forming ecotypes, and over time becameincreasingly isolated, leading to speciation and theneoendemics seen today (Stebbins and Major 1965,Kruckeberg 2004).

Other neoendemics arose when individuals of speciesgrowing off of infertile soils possessed traits allowing themto cope with infertile soils and colonized an edaphic island.Many of these transitions were associated with hybridizationand polyploidy events and coupled with other geneticchanges such as shifts in flowering time, drivingreproductive isolation and speciation in species with thesame ranges (sympatric speciation) (Rajakaruna 2004).These are a sample of the processes that have led to the highamount of endemism on infertile soils.

Geologic formations and their endemic inhabitantscontribute to regional diversity in many parts of the world.They contribute significantly to the high rates of endemismin “biodiversity hotspots” such as the California and CapeFloristic Provinces. In the arid Southwest United States andNorthern Mexico, geologic richness has given rise tosubstrate-specific floras and edaphic endemics includingthose of gypsum and limestone in the Chihuahuan Desert(Van Devender et al. 2013, Escudero et al. 2014), the fresh-water limestones of the Sonoran Desert (Anderson 1996,2011) and plants of various sedimentary formations on the

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Colorado Plateau. Many of the Southwest’s rare endemicspecies occur on specific substrates and the presence ofgeological diversity significantly increases the plant speciesdiversity of the region’s deserts.

The Upper Verde River

The region surrounding the Central and Upper Verde Riverof Arizona is yet another example of an area rich ingeological and botanical diversity. Abundant faultingresulted in uplifts of ancient igneous formations andPaleozoic sedimentary formations. Volcanos dotted thelandscape and lava flows formed mesas and plugs thatdammed streams, resulting in large freshwater lakes thatpersisted for thousands of years in which layers of freshwaterlimestone accumulated. This formed the patchy mosaic ofgeology and scenic landforms that can be seen today.

As part of a Master’s Thesis at Arizona State University, Ihave been conducting a floristic inventory of the UpperVerde River region. Over 12 geologic formations exist in thearea and sharp distinctions in plant species have beenobserved between the limestones (Devonian MartinFormation Dolostone [440-400 Ma], Paleozoic Redwall,Supai Group [380-260 Ma] and Tertiary Verde formation [7-2 Ma]) and a small patch of Paleozoic sedimentary TapeatsSandstone (700-400Ma) and two very old igneousformations (Gabbro [>1500Ma] and VerdeGranite[1700Ma]). Many species on these formations showpatterns of disjunction and endemism classically associatedwith edaphic islands.

Special Soils and their Role continued

Left: Escobaria missouriensis, a disjunct primarily from the Great Plains growing on dolostone. Right: Layers of the Supai group on topof cliffs of Redwall limestone at Perkinsville, Arizona. Photos by Francis S. Coburn.

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To date, 55 species of the approximately700 species in the Upper Verde River areaare found to grow exclusively onlimestones. Of these, 29 are populations atthe periphery of their ranges, 15 aredisjuncts, and 10 are regional endemics.The limestones, particularly the Devonian Martin Dolostoneand Tertiary Verde Formation, host many peripheralpopulations and disjuncts, primarily from the northernGreat Basin Desert and Colorado Plateau with a few speciesfrom the southern Chihuahuan Desert. Notable examplesfrom the north include dwarf sand verbena, Abronia nana(Nyctaginaceae), a disjunct primarily from the Great BasinDesert with ca. 100 miles to its nearest population in theGrand Canyon, and Missouri foxtail cactus, Escobaria(Coryphantha) missouriensis (Cactaceae), a disjunct mostcommon in the Great Plains of Wyoming and disjunct ca.125 miles from populations on the Arizona Strip. Thelimestones and Martin formation Dolostone host at least 10central Arizona endemics, five of which are US ForestService Sensitive and one, Purshia subintegra (Rosaceae),federally Endangered. These contribute to the area’s havingone of the highest densities of endemics anywhere inArizona (Salywon et al. 2013). Notable sensitive endemicsinclude Mearns’ Sage, Salvia dorrii subsp. mearnsii(Lamiaceae), an endemic of the Verde Valley and UpperVerde River, Rusby’s milkwort, Polygala rusbyi(Polygalaceae), an endemic spanning from the Verde Valleynorthwest along the Upper Verde River to the HualapaiReservation and Pediomelum verdiense (Fabaceae), a recentlydescribed endemic to the Verde Valley and Upper VerdeRiver.

To date, 32 species from the Upper Verde River are found togrow exclusively on the Verde granite, gabbro, and Tapeatssandstone, of which 14 are marginal populations, 17 aredisjuncts and one a central Arizona endemic: Mt.Dellenbaugh sandwort, Eremogone aberrans(Caryophyllaceae). Peripheral populations and disjunctsgrowing on these formations, as opposed to the northernaffinity of limestone plants, have centers of distribution insoutheastern Arizona and northern Mexico, with multiplespecies disjunct from the Sonoran and Mojave Deserts.Examples include harlequin spiralseed, Schistophragmaintermedia (Plantaginaceae), a disjunct with populationsnear Payson, ca. 80 miles to the southwest but otherwise

with a center of distribution in southeast Arizona, 240 milesaway, and royal sandmat, Chamaesyce dioica(Euphorbiaceae), a disjunct primarily from 200 miles to thesouth.

The limestone affiliated species represent >10% of the flora,and the contribution from other edaphic formationscontribute another 5%. Further analysis will bring to lightthe contributions of basalt formations, which have not yetbeen analyzed. Clearly, distinct geological formationscontribute significantly to the species richness of the region.

The geologic diversity and its relationships to plant speciesdiversity and distributions along the Upper Verde Riverundoubtedly hold evidence of the historic biogeography andplant evolution in the region. Since multiple speciesassociated with a particular rock type have centers ofdistribution in common, it is plausible they may be relicts ofpast migrations and dominant vegetation types, since thereis a low probability each species distributed to the soil typeindependently over time. On the other hand, on the scale ofthousands of years, species may have had the time enoughdisperse, one by one to these islands and form similarassemblages to their centers of distribution. Populationgenetic and phylogeographic studies combined with toolssuch as climate and species distribution (niche) modelingcould help to clarify the causes and history behind thesepatterns.

The patterns seen here of endemism are also in need offurther study. Although many groups inhabiting both theserpentine of California and gypsum of the ChihuahuanDesert floras are becoming well studied, little has been doneto address the systematics, adaptation, speciation, andbiogeography of edaphic endemics on limestone anddolostone in central Arizona. It is likely most of the

Special Soils andtheir Role continued

Left: Chamaesyce dioica, a disjunct from southern Arizona andNorthern Mexico growing in granite. Right: Eremogone aberrans,a central-northern Arizona endemic growing on granite. Photosby Francis S. Coburn.

continued on page 12

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The following information from the FNA website(floranorthamerica.org) describes the scope of this majorbotanical undertaking:

The Flora of North America project builds upon thecumulative wealth of information acquired sincebotanical studies began in the United States and Canadamore than two centuries ago. Recent research has beenintegrated with historical studies, so that the Flora ofNorth America is a single-source synthesis of NorthAmerican floristics. FNA has the full support of scientificbotanical societies and is the botanical community’svehicle for synthesizing and presenting this information.

The Flora of North America Project will treat more than20,000 species of plants native or naturalized in NorthAmerica north of Mexico, about 7% of the world’s total.Both vascular plants and bryophytes are included.

Species descriptions are written and reviewed by expertsfrom the systematic botanical community worldwide,based on original observations of living and herbariumspecimens supplemented by a crucial review of theliterature. Each treatment includes scientific and commonnames, taxonomic descriptions, identification keys,distribution maps, illustrations, summaries of habitat andgeographic ranges, pertinent synonomy, chromosomenumbers, phenology, ethnobotanical uses and toxicity,and other relevant biological information.

AZNPS members may wish to order copies of individualvolumes of FNA which are now available from the OxfordUniversity Press at discount rates. You can order fromwww.OUP.com/US or telephone (800-451-7556, using thepromo code 33009 to get the discount). Also, individualsponsorships for new illustrations may be purchased atfloranorthamerica.org.

Flora of North America Honors the Arizona Native Plant Society

The Flora of North America (FNA) project recently honored the Arizona Native Plant Society bydedicating a beautiful illustration of Chamaebatiaria millefolium, Rosaceae, (Fern Bush) to the Society.Accompanying the notice of the dedication was an archival quality print of the illustration by the giftedbotanical artist Marjorie Leggitt. The illustration will appear in the upcoming Volume 9 of the FNA.

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12 The Plant Press Arizona Native Plant Society Fall 2014

endemics of the region are neoendemics since manybelong to widespread, diverse, rapidly radiating lineages(e.g. Eriogonum and Salvia) and paleoendemics typicallybelong to geographically restricted groups with lowdiversity. Phylogenetic and population genetic studiescould resolve relationships within lineages, particularlybetween relatives growing on and off of limestone. Thiswould provide insight into the timing of niche shifts, thespeciation of endemics, and the climatic and geologicfactors that led to their diversification and contribution tospecies diversity in the region.

aReferences

Anderson, J. 1996. Floristic patterns on late Tertiarylacustrine deposits in the Arizona Sonoran Desert.Madroño 43:255–272.

Anderson, J. 2011. Further floristics on Late Tertiarylacustrine deposits in the Southern Arizona Deserts.Madroño 43:255–272.

Axelrod, D.I. 1958. Evolution of the Madro-Tertiary geoflora.The Botanical Review, 24(7), 433-509

Betancourt, J.L., T.R. Van Devender, and P.S. Martin, (Eds.).1990. Packrat middens: the last 40,000 years of bioticchange. University of Arizona Press.

Cain, S.A. 1944. Foundations of plant geography.Foundations of plant geography. Harper & BrothersPublishers.

Escudero, A., S. Palacio, F.T. Maestre, and A.L. Luzuriaga.2014. Plant life on gypsum: a review of its multiple facets.Biological Reviews.

Kruckeberg, A.R. 2004. Geology and plant life: the effects oflandforms and rock types on plants. University ofWashington Press.

Rajakaruna, N. 2004. The edaphic factor in the origin of plantspecies. International Geology Review 46: 471-478.

Raven, Peter H. 1964. Catastrophic selection and edaphicendemism. Evolution: 336-338.

Salywon, A., W. Hodgson, M. King, and K. McCue. 2013.Arizona’s endemic plants -more than just a list. ArizonaNative Plant Society’s 10th Arizona Botany Meeting.Desert Botanical Garden, Phoenix, Arizona.

Stebbins, L.G. and J. Major.1965. Endemism and speciation inthe California Flora. Ecological Monographs 35:1–35.

Van Devender, T.R., A.L. Reina-Guerrero, and J.J. Sánchez-Escalante. 2013. Flora of Chihuahuan Desertscrub onlimestone in Northeastern Sonora, Mexico. In: Gottfried,Gerald J., Peter F. Ffolliott, Brook S. Gebow, Lane G. Eskew,and Loa C. Collins, comps. 2013. Merging science andmanagement in a rapidly changing world: Biodiversity andmanagement of the Madrean Archipelago III; 2012, May 1-5, Tucson, AZ. Proceedings. RMRS-P-67. Fort Collins, CO:U.S. Department of Agriculture, Forest Service, RockyMountain Research Station.

Wallace, A.R. 1858. On the tendency of varieties to departindefinitely from the original type. J. Proc. Linn. Soc. Zool.3:53–62

Special Soils and their Role continued

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continued next page

Introduction

Botanists have long recognized the dynamic nature of plantgenomes. Hybrid and polyploid taxa have been describedfrom artificial crosses, morphological traits, cytology,isozymes, and more recently the incongruence betweennuclear and plastid gene trees. However, the contribution ofhybridization and polyploidization (aka whole genomeduplication) to plant diversity has long been debated. Thespectrum of opinions ranges from an ephemeralevolutionary dead end (Wagner 1970) to an omnipotentevolutionary process involved in adaptation to newenvironments (Lewontin & Birch 1966). At its core thisdebate reflects the tension between gradual versuspunctuated evolutionary change.

At the University of Arizona in the Barker Lab we investigatethe evolutionary genetic processes of hybridization andpolyploidization through comparative genomic methods. Weare currently generating a model system with naturalpopulations of hybrid and polyploid desert adaptedlycophytes in the genus Selaginella (Selaginellaceae). Theseplants commonly form large mats in association with rockoutcrops, and they possess a complex integrated phenotypethat allows them to resurrect from metabolic dormancy forshort intervals following rainfall events.

The Selaginella arizonica x eremophila Hybrid Zone

In the Sonoran Desert of southern Arizona and northernMexico two closely related species, Selaginella arizonicaMaxon and Selaginella eremophila Maxon are found. Thesetaxa are morphologically, ecologically, and for the most partgeographically differentiated. However, in the transitionzone between the Arizona Upland and Lower ColoradoValley vegetation divisions (Shreve & Wiggins 1951) they doform fertile hybrids. This area corresponds with themountainous areas east and west of Arizona State Highway85, which runs from the U.S. Mexico border at Lukeville toBuckeye west of Phoenix.

Hybridization between S. arizonica and S. eremophila washypothesized decades ago based on morphologicalintermediates in the field and mysterious mixed collectionson herbarium sheets. This hybridization hypothesis wasfurther supported as an example of homoploid hybridspeciation from biparental isozyme expression patterns(Therrien 2004).

Currently, based on work by members of the Barker Lab andothers (Windham & Yatskivetch 2009; Felger et al. 2013),several putative diploid hybrid populations as well as oneputative allotetraploid population have been locatedthroughout the hybrid zone. In order to complement ourevolutionary genetic studies, an ecological characterizationof the Selaginella arizonica x eremophila hybrid zone wasperformed from field surveys and herbarium specimens.These characterizations include a quantification ofmorphological differences, climatic niche differences, as wellas predicted occurrences of parental and hybrid taxa insouthern Arizona.

Methods and Results

Morphology

A morphological analysis was performed in order toquantify inter- and intraspecific variation of the two parentaland hybrid taxa. The morphological traits examined includeterminal and axillary seta shape, texture and length;microphyll width and length; internode length; rhizophoredensity; and strobili length. At least five measurements weremade per herbarium sheet for each trait under a dissectingmicroscope. All available voucher specimens deposited atthe University of Arizona and Arizona State Universityherbaria found within the vicinity of the hybrid zone, as wellas a random subset of three parental samples outside of thehybrid zone were examined. These were supplemented byfield observations and collections from 2012-2014.

Of the morphological traits examined, hybrids consistentlyhave intermediate character values for adaxial microphylllength and terminal seta traits such as shape, and texture

Characterization of the Selaginella arizonica xeremophila (Selaginellaceae) Hybrid Zone inSouthern Arizonaby Anthony E. Baniaga1 and Michael S. Barker1

1Department of Ecology & Evolutionary Biology, Universityof Arizona, Tucson, AZ 85712

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(Table 1). These terminal seta characters are delicateand best observed in newly formed microphylls (Tryon1955; Yatskivetch & Windham 2009). Interestingly,unique morphological variation is found in the hybridsfrom the southern populations of Ajo, Cabeza Prieta,and Organ Pipe Cactus National Monument. In theseindividuals, the terminal seta is often forked at the distaltip. This character is not observed from hybridindividuals around Phoenix.

Hybrid Climatic Niche and Occurrence Prediction

All available records throughout the entire range of S.arizonica, S. eremophila, and S. arizonica x eremophilawere collated from SEINet, the California Consortiumof Herbaria, and the Global Biodiversity InformationFacility. These records were manually verified,georeferenced, and the associated 19 bioclimaticvariables at 30 arc-second resolution (Hijmans et al.2005) were extracted and compared.

Of the bioclimatic variables included in the analysis,annual mean temperature and winter precipitation weresignificantly different between the hybrids and bothparents. Based on a Wilcoxon rank sum test, hybrids arefound in locations that are on average warmer and drierthan either of the parents (Figure 1).

These bioclimatic variables were also used in a MaxEntframework to predict regions with a high probability ofoccurrence of hybrids. Four different types of modelpredictions were made; S. arizonica, S. eremophila, S.arizonica + S. eremophila (parents lumped into onedataset), and all known localities of S. arizonica xeremophila.

The four different models revealed a high probability ofoccurrence of hybrids in the known hybrid zone, andoverlapped with all known localites of hybridpopulations. The prediction models did highlight a highprobability of hybrids in the Batamote Mountains eastof Ajo, and the northern Maricopa Mountains of theSonoran Desert National Monument of which there areno known observations of any Selaginella taxa.Interestingly, these models also highlighted a high

14 The Plant Press Arizona Native Plant Society Fall 2014

Table 1. Descriptions of important morphological differencesbetween Selaginella taxa.

Taxon Adaxial microphyll Terminal setae length (mm) shape and textureS. arizonica Maxon (1.7) 1.9-2.3 (2.8) Stout, straight; serrate, usually persistentS. eremophila Maxon (0.8) 1.3-1.5 (1.7) Delicate, pigtailed to contorted; smooth, deciduous with ageS. arizonica (1.1) 1.6-2.0 (2.6) Slender, sigmoidal; x eremophila usually smooth, often forked at tip; +/- persistent

Selaginella arizonica continued

Figure 1: Hybrid climatic niche differentiation. Annual meantemperature (°C) (p < 0.001), and winter precipitation (mm)(p < 0.001), Wilcoxon rank sum test. Notches representmedian values and asterisks represent significantlydifferent comparisons to parental taxa (p < 0.05).

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probability of hybrid occurrence outside the known hybrid zonein an unexpected region to the northwest. Only S. arizonica isknown from this general region, which includes the Kofa, LittleHorn, Harquahala, and Harcuvar Mountains.

Summary

The Selaginella arizonica x eremophila hybrid zone can becharacterized in three main ways. First, hybrids can berecognized via a combination of intermediate morphologicalcharacters related to adaxial microphyll length, and seta shapeand texture. Setae traits appear less labile to environmentaleffects and are the single best indicator of hybrid taxa. Second,the hybrid climatic niche is significantly warmer and drier thaneither of the parents. Finally, the hybrid zone is associated withthe transitional zone between the Arizona Upland and LowerColorado Valley vegetation divisions which corresponds withArizona State Highway 85 and may extend northwestward alongthe Interstate Highway 10 corridor.

Whether hybrids and polyploids are found in marginal habitatsrelative to the parents or have adapted faster to a new habitat isan open and important question. In addition to field surveys, weare testing hypotheses concerning the multiple origins of hybridand polyploid taxa, as well as measuring relevantecophysiological traits in this system.

aReferences

Felger, R.S., S. Rutman, J. Malusa, and T.R. Van Devender. 2013.Ajo Peak to Tinajas Altas: A flora of southwestern Arizona: Part3: Ferns, lycopods, and gymnosperms. Phytoneuron 37: 1–46.

Hijmans, R.J., S.E. Cameron, J.L. Parra, P.G. Jones, and A. Jarvis.2005. Very high resolution climate surfaces for global land areas.International Journal of Climatology 24:1965-1978.

Lewontin, R. C. and L.C. Birch. 1966. Hybridization as a source ofvariation for adaptation to new environments. Evolution 20:315-336.

Shreve, F. and I. L. Wiggins. 1951. Vegetation and flora of theSonoran Desert, Vol. I. Carnegie Institution of WashingtonPublications 591: 1-192.

Therrien, J. P. 2004. Phylogeny of Selaginella subgenusTetragonostachys based on nuclear and chloroplast DNAsequence data. PhD Dissertation, University of Kansas.

Tryon, Rolla M. Jr. 1955. Selaginella rupestris and its allies. Annalsof the Missouri Botanical Garden 42: 1-99.

Wagner, W.H. 1970. Biosystematics and evolutionary noise. Taxon19: 146-151.

Yatskievych, G. and M. D. Windham. 2009. Vascular plants ofArizona: Selaginellaceae. Canotia 5: 39-48.

Selaginella arizonica continued Who’s Who at AZNPSBOARD OF DIRECTORS

Cass Blodgett Director, Phoenix Chapter Co-President

[email protected]

Suzanne Cash Director, State Secretary

[email protected]

Wendy Hodgson Director, Education, Outreach and

Conservation Committee Chair [email protected]

Diane Kelly State Treasurer

[email protected]

Andy Laurenzi Vice President

[email protected]

Ries Lindley Director, Tucson Chapter President

[email protected]

Valerie Morrill Director, Yuma Chapter President

[email protected]

Mike Plagens Director at Large, Phoenix Chapter Co-President

[email protected]

Doug Ripley President

[email protected]

Andrew SalywomDirector at Large

[email protected]

Sue Smith Director, Prescott Chapter President

[email protected]

AZNPS COLLABORATORS Cass Blodgett Claire McLane Web Editor Administrative Assistant [email protected] [email protected]

Sue Carnahan Julie St. John Happenings Editor The Plant Press [email protected] Layout Editor [email protected]

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16 The Plant Press Arizona Native Plant Society Fall 2014

SPOTLIGHT ON A NATIVE PLANT

Mexican Tansyaster(Psilactis gentryi)by Bob Herrmann, Cochise Chapter, Arizona Native Plant Society

The Mexican Tansyaster (Psilactis gentryi) isa beautiful native wildflower that reaches thenorthern end of its range in the HuachucaMountains, its only known occurrence inArizona. In Mexico it occurs as far south asthe State of Durango. In the Huachucas it’san occasional annual that grows up to 2 feettall and produces beautiful 3/4 inch blue topurple flowers. The leaves are alternatelanceolate which distinguishes the species from otherblue-flowering asters in the Huachucas. It prefers moisthabitats in canyon bottoms and along rocky streambedsranging in elevation from six to nine thousand feet. The

plant has hairy, slightly glandular stems and leaves, startsgrowing in June, and flowers from September throughNovember.

The genus Psilactis consists of six species, five of whichoccur in the southwestern United States. Two of the fiveU.S. species occur in Arizona: P. gentryi and P. asteroids.The genus Psilactis was described by Asa Gray in 1849,its name being derived from the Greek psilos meaningnaked and aktis, meaning ray, referring to the fact thatthe ray akenes lack a pappus. The species name honorsHoward S. Gentry, a U.S. Department of Agriculturebotanist who collected extensively in northwesternMexico and later was a research botanist with the DesertBotanical Garden in Phoenix.

Caught in this photograph sharing a sip of MexicanTansyaster nectar is a European immigrant, the WoolCarder Bee (Anthidium manicatum) and an Arizonanative, the Gray Bee Fly (Anastoechus melanohalteralis).The Wool Carder Bee is a solitary bee, so named becausethe female bee collects plant hairs and makes her nestappear to be wool lined. The Gray Bee Fly is named forits color and bee-like appearance. The female fly seeksout grasshopper egg-pods that her maggots then feedupon. I have photographed several other varieties of beesand butterflies pollinating the flowers of the MexicanTanseyaster. As only an occasional annual it may takesome time to find it but the search is well worth theeffort.

a

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BOOK REVIEW by George M. Ferguson, University of Arizona Herbarium and Arizona NativePlant Society, Tucson Chapter

Trees of Western North America by R. Spellenberg, C.J. Earle, and G. Nelson, edited by A.K. HughesPrinceton University Press. 560 pp. July 2014. ISBN 978-0-691-14580-8. Paperback Flexibound: $29.95

New in the series of Princeton Field Guides, Trees ofWestern North America is a most definitive compactbook, designed for easy use in the field. It complements aparallel release: Trees of Eastern North America byanother combination of the same authors at the sameprice (for paperback with a firm binding). Both are alsoavailable in hardcover. For anyone interested in tree-likeplants in the West, the Trees of Western North America isit, covering 630 species west of the 100th meridian, andrightly proclaiming itself to be themost comprehensive, bestillustrated, and easiest-to-use bookof its kind.

The prize is in the detail. Simplyarranged to keep you from gettinglost in the forest of information,each tree entry is presented intaxonomic order. If you think youknow the plant family, there is noneed to search the index ofcommon and scientific namescombined. But, to facilitatedetermination of a species, apictorial leaf-key is presented at thefront of the book, designed for youwith leaf-in-hand, to find a speciesmatch within the book. Atop everypage of tree entries is a color-codedheader for the family in both Latinand vernacular, e.g. “Fabaceae: Bean or Pea Family.”Subsequent genera and species are covered alphabeticallyby scientific name within the family. Pages opposite thetree entries contain meticulous color drawings by DavidMore, depicting key features of tree habit, leaf shape, fruitand flower as well as the bark. Subtleties of texture andcolor shade are superbly conveyed. On the same pagewith the descriptive text, are precisely detaileddistribution maps for each species. These are reminiscentof the USDA maps based on those by arborist Elbert L.Little, Jr.

Descriptions for every native or naturalized tree-likespecies in the West are accurately written for even themost demanding botanist, yet in layman’s terms.Wonderful panels with expert botanical summaries startthe description of each family and each genus. Tree taxonentries are standardized. Thus consistent terms are usedand appear in the same order throughout, beginningwith a single, best, common name, followed by thescientific name, then “a.k.a.” alternate common names, if

any. Next, a “Quick ID” sentence inboldface summarizes traitsuniquely delineating the species inthe region it occurs. Followingthis, an easy to read paragraphdescribes the bark, twig, leaf, fruitor cone, and for flowering plants,the flower, all in metric units. Stillnot convinced of your ID? There’smore.

In compiling this book, the authors’expertise shines through in the“Similar Species” paragraphs and“Notes” sections of each tree entry.Fine details for determiningwhether you have the right speciesare found there, as well asexplanations of up-to-datenomenclature, including differingscientists’ views on the species, if

any, or evidence of intergradation and hybridization.There is also information about the plant’s uses, andwhether it is considered poisonous. A most usefulstatement under “Habitat/Range” is provided for eachentry, beginning with either “Native” or “Introduced”(and from where). It includes elevations in meters, inwhich States the species occur, including Alaska andCanadian Provinces (even if just a handful tree speciesgrow there), and whether that taxon extends beyond themap boundary to the eastern U.S. or Mexico.

continued next page

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18 The Plant Press Arizona Native Plant Society Fall 2014

Trees of Western North America continued

What’s a tree? The authors state there is no scientific differencebetween a tree and a shrub. Though a tree should have a single woodystem and a crown of branches, a broader definition of a tree is takenhere. It includes plants generally taller than a human, often withmultiple woody trunks, and plants that may be thought of as shrubs.So, sagebrush, cholla and prickly pear cacti, nolinas and yuccas areincluded in the book as well as many small overlooked species usuallyconsidered shrubs. Likewise, species, local or endemic to narrowregions, are not neglected, e.g. Oregon coast trees, Channel Islandnatives, borderlands plants of San Diego County, and rare species ofsouthern Arizona, or Big Bend and the lower Rio Grande Valley ofTexas.

Complex and difficult genera are treated in this book so they appeareasy; such as the willows, with nearly 30 species in the West;hawthorns, another 25 to 30 species, some of which are hybrids; notto mention the oaks, comprising about 48 species in the West. And,seven species of introduced tamarisk are covered. The boomingbusiness of nomenclature change is addressed effectively with newfamily alignments, and generic rearrangements adopted. The authorsrelied on these references: Flora of North America (eFloras.org), thePLANTS Database (USDA) as well as recent technical literature. Forexample, the former New World Acacia genus in the book is split intothree other genera, Mariosousa, Senegalia, and Vachellia, and some ofthe genus Rhamnus species are now represented in this book asFrangula.

Despite all the wonderful aspects of this magnificent new book,perhaps wishful improvements would include the fascinatingdiversity, foreign and native, of Hawaii. At least some indication isgiven if a species occurs in Mexico, otherwise it stops at the border!While coverage of many landscape shrubs and trees is welcomed, afew native trees were left out, such as species of tree-like Berberis,Coursetia and Erythrina in Arizona. Perhaps some more commonornamental trees, such as other Senna and Pinus halepensis shouldhave been included. Several of the native pines are poorly portrayed.A few typographical errors exist, e.g. “mm” instead of “cm” in a fewentries; obvious portions of ranges missing from some maps, thoughstated correctly in the text; and, in a few, probable measurementdiscrepancies such as tree height.

Overall, this book makes the grade “A” in nearly every aspect ofpresentation; it is tremendously useful, navigable, readable, highlyaccurate and immensely informative. The added perks, such as thefamily and genus panels, glossary, essays on The Biggest Trees, PineEcology, Forests and Climate Change, the 20 page introduction toTree Biology, Taxonomy and Names, Forest Structure and theexpertly illustrated leaf keys only increase its usefulness, functionality,and value.

a

Many Thanks to Marilyn HansonLong-time Tucson Chapter member andAZNPS webmaster Marilyn Hanson recentlyturned her webmaster duties over to CassBlodgett, co-president of the PhoenixChapter. For many years Marilyn did asuperb job of maintaining the Society’swebsite as well as performing numerousother duties for the Tucson Chapter. Shehas also had a profound positive impact onnative plant conservation by serving as theindefatigable volunteer coordinator for theSonoran Desert Weedwackers, the AZNPSpartner organization responsible foreradicating bufflegrass and other noxiousspecies from many critical Sonoran Deserthabitats. We sincerely thank Marilyn for hermany contributions to the Arizona NativePlant Society and look forward to hercontinued efforts to help achieve theSociety’s plant conservation goals.

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www.aznativeplantsociety.org The Plant Press Arizona Native Plant Society 19

Please plan to attend The Arizona NativePlant Society’s 2015 Arizona Botany Meeting,which will be held in the beautiful andhistoric Branigar/Chase Discovery Center atthe Museum of Northern Arizona, Flagstaff.

The theme for the conference is:“Ethnobotany: Past, Present and Future.”Presentations and poster sessions will bedelivered at the Museum on Saturday, April25, and several field trips will be conductedon Sunday, April 26. Please consult the AZNPSwebsite for the formal call for presentationsand posters, meeting schedules, andregistration information.

Announcing the 2015 Arizona Botany MeetingApril 25-26, 2015 — Museum of Northern Arizona, Flagstaff

Congratulations to Cass BlodgettIn the summer of 2014 Cox Enterprises and The Trustfor Public Land (TPL) selected AZNPS Phoenix Chapterco-president Cass Blodgett as a finalist for the CoxConserves Heroes Award in recognition of hisvolunteer efforts to create, preserve or enhance theshared outdoor places in local communities. Cass wasselected for his outstanding work as a volunteer citizenscientist on the North Mountain Plant InventoryProject. Through that on-going work he collects anddocuments North Mountain’s diversity of flora, teachesthe public about the native flora, and leadsinterpretative hikes. As a long-time member of theAZNPS, Cass has made many other contributions tothe Society by serving as co-president of the PhoenixChapter, helping to organize monthly meetings,leading field trips, recently becoming the AZNPSwebmaster, and participating in many other efforts toraise the awareness of native plants and theimportance of their conservation. Cass’ nominationincluded a $2,500 award which he designated to theAZNPS. Thank you Cass!

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THE ARIZONANATIVE PLANT

SOCIETYPO Box 41206

Tucson AZ 85717www.aznativeplantsociety.org

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Enclosed: m $15 Student m $75 Commercialm $30 Individual m $100 Plant Loverm $35 Family m $500 Patronm $50 Organization m $1,000 Lifetime

Mail to: Arizona Native Plant SocietyPO Box 41206, Tucson AZ 85717

New Members Welcome!People interested in native plants areencouraged to become members. People mayjoin chapters in Flagstaff, Phoenix, Prescott,Cochise County (Sierra Vista), Tucson, orYuma, or may choose not to be active at achapter level and simply support thestatewide organization.

For more information, please write toAZNPS (see return address above), visitwww.aznativeplantsociety.org, or contactone of the people below:

Cochise: Doug Ripley,[email protected]

Flagstaff: Dorothy Lamm, 928.779.7296

Phoenix: Cass Blodgett, 480.460.0725, andMichael Plagens, 602.459.5224

Prescott: Sue Smith, 408.507.7706

Tucson: Ries Lindley,[email protected]

Yuma: Valerie Morrill, [email protected]