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Chromosome Races in Sarcobatus (Sarcobataceae, Caryophyllales)

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Chromosome Races in Sarcobatus (Sarcobataceae, Caryophyllales) Great Basin Naturalist Volume 59 Number 4 Article 1 10-15-1999 Chromosome races in Sarcobatus (Sarcobataceae, Caryophyllales) Stewart C. Sanderson Shrub Science Laboratory, Rocky Mountain Research Station, Forest Service, U.S. Department of Agriculture, Provo, Utah Howard C. Stutz Brigham Young University Mildred Stutz Brigham Young University Richard C. Roos Waste Management Federal Services, Inc., Northwest Operations, Richland, Washington Follow this and additional works at: https://scholarsarchive.byu.edu/gbn Recommended Citation Sanderson, Stewart C.; Stutz, Howard C.; Stutz, Mildred; and Roos, Richard C. (1999) "Chromosome races in Sarcobatus (Sarcobataceae, Caryophyllales)," Great Basin Naturalist: Vol. 59 : No. 4 , Article 1. Available at: https://scholarsarchive.byu.edu/gbn/vol59/iss4/1 This Article is brought to you for free and open access by the Western North American Naturalist Publications at BYU ScholarsArchive. It has been accepted for inclusion in Great Basin Naturalist by an authorized editor of BYU ScholarsArchive. For more information, please contact [email protected], [email protected]. The Great Basin Naturalist PUBUSHED AT PROVO, UTAH, BY M.L. BEAN LIFE SCIENCE MUSEUM BRIGHAM YOUNG UNIVERSm ISSN 0017-3614 VOLUME 59 31 OCTOBER 1999 No.4 Great Basin Naturalist 59(4), '01999, pp. 301...,')14 CHROMOSOME RACES I SARCOBATUS (SARCOBATACEAE, CARYOPHYLLALES) Stewart C. Sandersonl , Howard C. Stutz', Mildred Stutz2, and Richard C. Roos3 ABSTRACf.-SarcobautS Nees" a genus of North American halophytic shrubs, consists of 2 species: S. oormi<;ulatus (Hook.) Torr. (n = 18.36), which is widespread in western North America, and S. baileyi Cov. (n = 54), endemic to Nevada. Within S. vennic-'Ulattl$, populations of n = 36 are widely distributed, whereas populations ofn = 18 are found only in the Sonoran Desert, northern California, and northwestern Great Plains, locations at the periphery ofthe species range. Although the chromosome Dumber ofn = 18 is apparently tetraploid, failure to form an n = 27 race intermediate to those ofn = 18 and n = 36 suggests that n = 18 S. vermiculatw is ofsignificant age and that it behave.\i chromosomal­ ly as a diploid. Sarcobatu3 has a long fossil pollen record and endured Pleistocene climatic extremes with little range displacement. Key words, Sarrohatns, polyploidy, autoploidy, diploidizaticn, Plei.rtocene distributions. Sarcoba:tu8 Nee,. is an endemic North Amer­ wise known to date only from the Amaran­ ican genus consisting of spiny shrubs thai are thaceae and Aizoaceae (Rodman 1994). Sarro­ monoecious and wind-pollinated, with reduced, ba:tu8 propaguJes, fonned from the winged calyx fleshy leaves. It has historically been placed in offemale flowers, are adapted for wind disper­ the Chenopodiaceae (order Caryophyllales). sal across barren surfaces such as mud flals or Like members ofsubfamily Salsoloideae (Spiro­ sand dunes (Danin 1996). The segments ofthis lobeae) in the Chenopodiaceae, the embryo of calyx are accrescent and enclose the seed in Sat'cobatus is elongate and coiled in seed, en­ fruit (Standley 1916, Welsh et aI. 1993), some­ abling rapid extension during gennination. In what like that ofsome chenopods. common with the chenopod genera Spinacia, In spite of these similarities to the Cheno­ AtTiplex, and Ceratoides, plants of Sarcobatus podiaceae, other Sarcobatus characteristics are contain 6-oxygenated flavonoids (Sanderson et discordant within that family. Macromolecular al. 1988). Of 12-plus families in the Caryo­ and sieve-tube plastid evidence suggests a link­ pbyUales, 6-oxygenated Ilavonoids are other- age of the genus to other families, particularly IShnJb Sclenoes L.abonfQry. Rocky Moonratn Researcb StatilJ.... fbrest Setvlee, u.s. De{:wtUleflt ofAgri"<:o.,ltute, 735 N. 500 E:.tst, P'n:M>, UT~. AutlWlr to whom I~ for reprinlf ~bou1d be Iel\t. tDeprutment of Bob.ny and RAn~..e Sdenc:e, Brigham YQU~ University,1't'oYo. UT84flO2. 3l,Vaste MW-igement Federa.I SeNioes, Inc.. Northw~~t OperaHom. 345 "HIs St.. Richland, WI!. 9Ii052. 301 302 CHEAT BASlN NATURALIST [Volume 59 the Nyctaginaceae and Phytolaccaceae (Behnke MATERIALS AND METHODS 1993, 1994, Downie et al. 1997). A feature of Sarcohatus not duplicated in Chromosome counts, leaf-flavonoid content, any related f~unily is the male inflorescence in and morphological measurements of Sa.rcoba­ which numerous, asepalolls flowers are borne tus were obtained Ifom sites throughont its in a conelike structure formed from peltate range, and representative voucher specimens scales. In view of the distinctiveness of Sarco­ have been deposited at BRY. Cytological and batus, a separate family, Sarcobataceae, has chemical methods have evolved somewhat recently been created for it (Behnke 1997). during accumulation of data for this report Fossil pollen of Sarcohatus has been found (1982-1996). The following procedures are in Eocene, Oligocene, and latcr strata from currently used. localities in Washington, Oregon, IdallO, Wyo­ Chromosome Counts ming, and Colorado (Leopold and Macginitie 1972, Leopold and Denton 1987); the genus We have found the use of5% acetic acid or was present in late Miocene in the area of an equivalent strength ofhousehold vinegar to Jackson Hole, Wyoming, before the present be safer fi)r field use and to give result,; com­ Teton Mountains were uplifted (Bamosky 1984). parable to those of alcohol-acetic acid in fixa­ Because floras during the early and mid-Ter­ tion of meiotic m.aterial for taxa of the Chen­ tiary were considerably more tropical and pre­ opodiaceae and Sal'cohatus (Stutz and Sander­ ponderantly arboreal (Leopold and Macginitie son 1983, Sanderson and Stutz 1994). We 1972), SarcohatuB may have had somewhat dif~ determined chromosome counts from pollen ferent environmental tolerances at that time mother cells ofmale flower huds fixed in vine­ than it does at present. However, it would gw' (5% acetic acid strength) and refrigerated likely have grown on the margins of dessicat­ (2°C) or Ifozcn (-20'C) for up to several months ing lakes rather than within forests. before examination. Anthers were squashed in Sarcoha.tus at present consists of 2 species, acetocarmine stain, which was concentrated to S. verm;'cul"tus (Hook) Torr., ofwide distribu­ supersaturation by means of evaporation on tion in western North America, and S. haileyi the microscope slide. Preparations were pre­ Cov., which is limited to the central and western served by replacement of the dye solution Palts of Nevada in the Great Basin. Sm'cohatu8 under the cover slip with 45% acetic acid, and verrnk'tdatus occupies saline bottomlands, playa then with corn syrup, thinned as necessary dunes, and badlands, while S. hai.ley;' is found with vinegar, which retarded mold growth in on arid slopes, usually in monotypic stands or the syrup solution better than did 5% acetic in association with Atriplex confertifolia. acid. A few additional counts were made from The only published chromosome count for root tips using methods described by Stutz Sarcobatus of which we are aware is n = 18, and Sanderson (1983). from a collection of S. verrniculatus near Med­ Flavonoids icine Hat, Alberta (Bassett and Crompton 1970). The Chenopodiaceac and Phytolac­ Determination ofaglycone moieties offoliar caceae have hase chromosome numbers ofx =: flavonoids was carried out upon either 25 cc of 9 (Turner 1994). Although the Nyctaginaeeae crushed air-dried leaves or a similar volume of has a variety of chromosome numbers, Turner 5% acetic acid-preserved leaf material. Sam­ (1994) suggests its base may be x = 10. Never­ ples were hydrolyzed for 60 min in IN HCI theless, x=:9 would be a plausible alternative. over a boiling water bath to remove glycosidic Sarcohatus is somewhat isolated pbylogeneti­ sugars, then ground in 85% aqueous methanol, cally, but it seems likely, in view ofits relation­ filtered, and washed with additional 85% ships with these f,unilies, that x = 9 is its base methanol. The combined filtrate and washings number as well. (50 mL total) were mixed with an equal amount It is our purpose to report on the distribu­ of water and then extracted with 50 mL of tion <md morphological characteristics of the ethyl acetate, and again afterwards with a small chromosomal races we have encountered rinse. The combined ethyl acetate extract and within taxa of the new f11milv, Sarcobataceae, rinse was blown down with compressed air and to present available evidence regarding and the residue extracted with 1-2 mL 45% their origin and genetics. acetic acid, which was then applied to half~ 1999] SARCOBATUS CHROMOSOME RACES 303 sheets of Whatman 3M chromatography paper. individuals to parental octoploid plants, were Chromatography was carried out as reported encountered in 6 additional instances. How­ previously (Sanderson and Stutz 1994), and ever, no plants ofhigher chromosome numbers sheets were thoroughly air- 01' oven-dried be­ were identified within populations of tetra­ tween 1st- and 2nd-dimensional chromatogra­ ploid S. vermiculatus, nor in S. baileyi. phy. Most flavonoid compounds occurring in Octoploid populations of S. vermiculatus Sarcobatus have been previously isolated and were found to occupy the majority of the cbemically identified (Sanderson et al. 1988). species range, from Montana and the Dakotas to northern Arizona, and from Colorado and Morphology Nebraska westward to California (Fig. 1). Tetra­ Morphological characteristics of mature ploids were limited in the north to Alberta, Sarcobatus plants in natural populations
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