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 plant height and width, leaf coastal areas with shallow water tables, upon length and width (measured from a large leaf partially stabilized sand dunes and at the edge on the branch nearest the investigator), and of salt pans. Our samples showed that these male inflorescence length (excluding pedun­ plants are tetraploid. A few small populations cle). Male inflorescence width could not be of similar tetraploid plants are found near the included because inflorescences in S. baileyi Gila River in southern Arizona, growing on had already shed most of their scales at the sandy sites in areas unmodified by agriculture. season in which measurements were taken. Except for tbe Sonoran Desert plants and a few rare octoploids at northern extremes or RESULTS higher elevations, Sarcobatus is otherwise absent from the Chihuahuan, Sonoran, and Original cbromosome counts representing Mojave walm deserts of the southwestern 224 localities (Fig. 1, Appendix) were obtained United States and Mexico. in this study. Sarcobatus chromosomes are Tetraploid and octoploid populations grow­ small, but large enough forJlgure-8 bivalents ing in close proximity were encountered in to occur at a moderate frequency (Fig. 2). the upper Pit River drainage, at Goose Lake on Meiosis appeared to be regular in both diploid the California-Oregon border, and at sites in (n = 18) and octoploid (n = 36) S. vermicult.­ Montana and North Dakota (Fig. 1, Appendix). tus plants as well as in I2-ploid (n = 54) S. In a mixed-ploidy site near Goose Lake, bailey" Abnormalities such as multivalents west of Davis Creek, California, chromosome were observed only rarely, and spontaneous counts of 17 plants showed that, although in increase to higher ploidy appears to be rela­ close proxiInity, the tetraploid and octoploid tively rare. plants formed discrete patches (Fig. 3). While Except where there was geographic overlap all were browsed severely by cattle at that site, of races of different ploidy levels, populations those in the octoploid patch appeared taller containing individuals of a higher ploidy and more vigorous) as contrasted with the occurred in only a few cases (Appendix). In 2 smaller, more heavily damaged plants in the of 166 apparently octoploid populations of S. tetraploid patches. We found no plants at any verrniculntus that were sampled, a single 12­ of the mixed-ploidy sites with intermediate ploid individual was also encountered. One chromosome numbers suggestive of interploid population was located northwest ofTonopah, hybridization. Nevada, and the other near Willard, New Mex­ Because individuals ofS. vermiculatus show ico. Individuals of intennediate chromosome considerable vegetative plasticity, it is often numbers of approximately 9-ploid to lO-ploid, difficult to distinguish tetraploid from octo­ apparently representing backcrosses of I2-ploid ploid plants in the field. However, as shown in 304 GREAT BASIN NATURAUST [Volume 59

** *

0 o 0 t! o 0 0 0 e 00 0 0 0 0 0 8 0 0 0 0 'It, 0 0 0 0 0 a 0 0 0 0 0 0 0 0 fO 0 0 <;l 0 ~ 00 0 00 o 0 0 *~ 0 0 0 0 * o 8 0 0 0 000 0 0 0 0 0 0 a o a °0 0 0 0 0 c90 0 a

0 'C 0

0 Q:> 0 0 0 '.~ 0 ,

* S. vermiculatus n = 18 o S. vermiculatus n = 36 * S. bai/eyi n = 54

Fig. 1. Geographic distribution ofchromosome counts for Sarcobatus baileyi and S. vermWtilatus.

Table 1, the leaves of tetraploid plauts are sta· ranges. Plants from northern Oregon and Wash· tistically shorter and narrower than those of ington (the "northwestern 8x" group) were octoploids. numerically tallest, though this was not statis­ Populations ofoctopJoid S. vermictJatt.. may tically significant. differ dramatically in stature (Roos 1984; Figs. Inflorescence length showed few differences 4, 5, Table 1), differences which are largely hetween ploidies or by geographic region in S. maintained in the common garden (Stutz un· oermiroWws. However, temporal variation was puhlished data). However, only slight variation observed which at a particular seasou gave the was encountered across larger geographical impression of geographic differences (data not 1999J SARCOBATUS CHROMOSOME RACES 305

Fig, 2. Chromosomes of Sarcobatus baileyi, n ~ 54 Qefi), and S. vermIc«Iatus, n = 36 (center) and n = 18 (light). Scale bar = to 11m.

t N n= 18 (5 plants .0llIllcd)

; n=36 n= 18 ![5 plan" ""'"'ted) (4 plam1l COOI1tcd) .. ------_.------Fence

Roadway

Fig. 3. Distribution ofcllromosome COWlts of Scweobatu,s verrnict.Jatw from 3.5 km west of Davis Creek. California, made in 1995. nuee additional counts of 8~ 4x, and 4x (not shown), made the year previously, came from somewhere in the combined arel.l ofthe 8x patch and the berm. shown). Since we observed many populations Also, male inflorescences in some octoploid both in early summer when meiotic buds were populations of northern Arizona and southem being collected, and later at the time when Utah were observed to be of normal size in morphological measurements were taken, it was early sununer, but by measurement time were possible to compare inflorescences produced sholter (and much less abundant) than tbose at these different times and observe differences ofnorthern populations. that may have resulted from changes in season Male inflorescences of S. bailey; plants are or weathet: In early summer 1995, inflores­ consistently shorter than the shortest lengths cences of octoploid populations in northern of those of S. vermiculatus (Table 1). This WAS California were very short compared to those observed to be true both for young inflores­ observed elsewhere. However, as shown in cences, in which pollen mother cells were Table I, by late summel; at the time when undergoing meiosis, and also in matnre inflo­ measurements were taken, inflorescences were rescences after pollen had been shed. The as long as or longer than those of other areas. plants of S. baileyi are generally also shorter 306 GREAT BASIN NATURALIST [Volume 59

TABLE 1. Average of population means for Sarcobatus baileyi (SABA) and S. vermiculaw.s (SAVE) for morphological chm"acters, measured in late summer, with st

than those of octoploid S. vermiculatwJ, but axils ofwoody branches. Ifconditions are favor­ are not significantly shorter than tetraploids able, a 2nd crop ofinflorescences may be pro­ using the present data set. Leaves of S. baileyi duced on the apices of elongate shoots. In S. are shorter than those of either ploidy of S. vermiculatus, all flowering is on elongate shoots vermi(:ulatus, while not any narrower. and may continue all summer when moisture AltllOugh S. baileyi has been suspected to is adequate (whether this relates to lowering intergrade with S. vermicttlatus (McMinn 1939), of water tables through the season or to tbe we fouud no morphological or cytological exhaustion of surface moisture would be a evidence for hybridization. S. baileyi was n = matter for further study). Individual female 54 throughout its range, but S. vermiculatus, flowers are usually located immediately below even when growing nearby, was always n = male inflorescences in both species. 36. In locations where S. vermicu1atus and S. As previously reported (Sanderson et al. baileyi come into contact, S. baileyi plants are 1988), the flavonoid complement ofSarcobatus often taller than usual, probably because of species evidences the presence ofan enzymatic moister conditions, and conversely, S, vermic­ activity for flavonol 6-oxygeuation, with indi­ ulatus plants become smaller and more spiny vidual plants showing some or all of the 6­ in drier circumstances, as they come near pop­ methoxy f1avonols (6-methoxy kampferol, pat­ ulations of S. baileyi. In spite of this conver­ uletin, and spinacetin), as well as the corre­ gence in stature, any confusion about their sponding 6-unsubstituted f1avonols (kampferol, identity can be resolved when male inflores­ quercetin, and isorhamnetin). In addition, 3­ cences are present, and foliage characteristics O-methylated f1avonols were often observed. are also often helpful. Besides the difference S. baileyi plants from southeastern Nevada already noted in leaflength, leaves of S. baileyi had f1avonoids in proportions similar to those are earlier-deciduous in the late summer than seen in S. vermicuJl1tus, but samples from the those of S. ve17niculatus, and the time ofappear­ western part of its range (western Nevada) ance ofnew leaves in tl,e spring usually differs, showed a quantitatively different and more vari­ with S. baileyi leafing out first. able flavonoid profile, often with only small Sarcobatus baileyi flowers most profusely at amounts of 6-methoxy compounds. We have leaf-flush, with male floral buds appearing ter­ not been able to discover morphological traits minally on new short shoots arising from the that correlate with this biochemical difference. 1999] SARCOBATUS CHJIOMOSOME RACES 307

Figs. 4-5. Plants from t.aJl~ and shorHtatured populations ofoctoploid (n = 36) Sarcobatus vcnnicul.atus: 4 (top), planl 2.5m in height, about 10 mi north of Monticello. Utah; 5 Oower), plants 0.25 In in height, gmwing 10 mi north of Milford. Utah. Card is 20 X 20 em.

DISCUSSION that did not shift extensively during these time periods. Late Pleistocene and Holocene Although long-distance eastward transport DistIibutiOD PattenlS ofsmall amounts ofSm·cobatus pollen has been Changes in altitudinal or geographic range noted (Maher 1964, Birks 1981), more abun­ were widespread among plants of western dant stratigraphic OCCUlTences would indicate North America in response to fluctuating cli­ that the genus was present locally (Tbompson mates during the Pleistocene and Holocene 1992), especially in the western or upwind parts (Betancourt et al. 1990). However, becall~e of of its species range. The presence of Sarcoba­ its apparentlimilation to saline soils, Sa1wha­ tus pollen at near-full glacial or earlier is docu­ tus may be an example ofa taxon with a range mented by longer pollen records from the 308 GREAT BASIN ATURALIST [Volume 59

Columbia Basin in Washington (Bamosky 1985), Adaptive Advantages ofPolyploidy eastern Idaho (Beiswenger 1991), northwest­ The absence of tetraploid S. vermiculatlls ern Wyoming (Whitlock 1993), central Nevada within the Great Ba.,in or other parts of the (Thompson 1992), Owens Valley, California central species range (Fig. 1) might suggest (Koehler and Anderson 1994), and northern that the tetraploid race in these areas has heen Arizona (Anderson 1993). In addition, pollen replaced hy competition from octoploids. In or macrofossil records document the presence several othcr apparently autoploid complexes, of SarcobNew Mexico, and Utah widespread than diploids (Manton 1934, Zohary (Betancourt 19W). Early Holocene records of and Nur 1959, Stutz et al. 1975, Stutz 1983), S

(4x + 8x = 12><) and apparent backcrosses of Department of Agriculture of any product or such to the parental chromosome number. service. Given sufficient selective value, a 12-ploid race We wish to thank those who have given like that of S. bailey might be expected to encouragement or suggestions relating to this eventually appear in S. vel'miculatus. study. Since the tetraploid chromosomal race should give rise to hexaplOid plants by unre­ LITERATURE CITED duced gametes in the same manner, the exis­ tence of an octoploid chromosomal race in the Ar..-DERSON. R.S. 1993. A 35,000 year vegetation and cli­ matic history from Polato Lake, Mogollon Rim, Ari­ absence of any hexaploid race is therefore un­ zona. Quaternary Research 40:351-359. expected. This anomaly could be best explained BABCOCK, E.B., A."'lD C.L. STEBBINS, JR. 1938. The Ameri­ if the tetraploid race were relatively old and can species ofCrepis. Carnegie Institl,lle ofWashing­ had therefore become diploidized. In that case too, Washington, DC. hexaploid individuals that might be formed BARNOSKY. C.w. 1984. Late Miocene vegetational and cli­ matic variations inferred from a pollen record in would be functionally eqUivalent to triploids, northwest Wyoming. Science 223:49-51. and sterile. Therefore, ifa race ofhigher ploidy __. 1985. Late Quaternary vegetation in the south­ was produced, it would have to be at the octo­ western Columbia Basin, Washington. Quaternary ploid level or higher, as is observed. Research 23:109-122. BASSETT, LJ., AlI.""D C.W CRQ;\1PTO~. 1970. In: IOBP chro­ Similar reasoning could be applied to poly­ mosome number reports XXXVII. Taxon 19:437-442. ploids in other taxa, for instance Atriplex BEHNKE, H.D. 1993. Further studies of the sieve-element ca.nescens and A. cOllfertifolia (x = 9) of the plastids of the Caryophyllales including Barbeuia, Chenopodiaceae. Both taxa exhibit wide­ Corrigiola, Lyallia, Microreo, Sarcobatus, and Tele­ spread tetraplOid races (Stutz and Sanderson phium. Plant Systematics and Evolution 186:231-243. ----;c. 1994. Sieve element plastids: their signjficance 1979, 1983, Sanderson et at. 1990, Sanderson for the evolution and systematics of the order. Pages and Stutz 1994) and have formed polyploids at 87-121 in H.D. Behnke and T.]. Mabry. editors, hexaplOid or octoploid levels, which would Caryophyllales: evolution and systematics. Springer­ provide an indication of whether the respec­ Verlag, Berlin. tive tetraploids are dip]oidized or not. In S. _-,-,' 1997. Sarcobataceae-a new family ofCaryophyl­ lales. Taxon 46:495-507. venlliCtlla.tus the absence of diploids and the BEISWENCER, J.M. 1991. Late Quaternary vegetational failure of the tetraploid race to polyploidize to histOly ofGray's Lake, Idaho. Ecological Monographs the hexaplOid level both suggest that the tetra­ 6U65-I82. ploid race is ancient, probably dating back BETANCOURT, J.L. 1990. Late Quaternary biogeography of the Colorado Plateau. Pages 259-292 in j.L. Betan­ into the Pleistocene. court, T.R. Van Devender, and P.S. Martin, editors, Success of octoploid S. venniculntus in the Pachat middens: the last 40,000 years of biotic relatively southern parts of the species range, change. University of Ariwna Press. Tucson. in comparison to tetraploid populations located BETANCOURT, J.L., T.R. VA'::'l DEVENDER, AND P.S. MARTIN. along the u.S.-Canada border and in north­ 1990. Packrat middens: the last 40,000 yem'S of biotic change. University ofArizona Press, Thcson. eastern California, may suggest that octoploids BIRKS, H.].B. 1981. Long-distance pollen in late Wiscon­ are better adapted to warm, arid climates. Ifthis sin sediments of Minnesota, U.S.A.: a quantitative is the case, their ascendency over the tetraploid analysis. New Phytologist 87:630-661. race may therefore likely have occurred dur­ CRO:"lQI.:IST, A. 1988. The evolution and classification of ing the Holocene. The Sonoran tetraploid race flowering plants. 2nd edition. New York Botanical Garden, Bronx, Y. occurs in a warmer climate, but it appears to DA.'llN, A. 1996. Plants of desert dunes. Springer-Verlag, be ecologically divergent and somewhat inde­ Berlin. pendent from the evolutionary trends of the DOWNJE, S.R., D.S. KATZ-DoWNIE, AND K,-J. CHO. 1997. remainder ofthe species. Relationships in the Caryopb}'lIales as suggested by phylogenetic analyses of partial cWoroplast DNA ORF-2280 homolog sequences. American Journal of ACKNOWLEDGME?>JTS Botany 84:253-273. EZCURRA, E., R.S. FELGER, A.D. RUSSELL, AND M. EQt:I­ This research was supported by BHP-Min­ HVA. 1988. Freshwater islands in a desert sand sea: erals Inc. and ROCk)' Mountain Research Sta­ the hydrology, flora, and phytogeography of the tion, Forest Service, U.S. Department ofAgri­ Gran Desierto oases ofnorthwestern Me:

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Tit VillI Devender, and ES. Martin, edi­ RODMAN, J.E. 1994. Cladistic and phenetic studies. Pages tors, Packrnt middens: the last 40,000 years of hiotic 279-301 in H.D. Belll1ke and TJ. Mabry, editors, change. University of AJ:i:r.ona Press, 'fucson, Cary{)phyllal(~s; evolution nncl systematics. Sl?rtn~cr· __. 1992. Late Quaternary environments in Ruby Verlag, Berlin. Valley, Nevada. Quaternary Research 37:1.-15. Roos, RC. 1984. Heril:nble variation in Sarcob(ttu$ vcr­ TlJRNEH, B. L. 1994. Chromosome numbers und their miculotus. Master's thesis. Brigham Young UnivC!'· phyletic ioterpretation. Pages 27-74 in RD. Behnke sHy, Provo, UT. and 1:J. Mabry. editors, Caryophyllules: evolution SANDKKSON, S.C., C.I,. Cnu. E.D. McARTHUR, ANI) RC. and systemlltic<;. Springer·Verlag, Berlin. SnJTz;. 1988. Evolutionary loss of flavunoids and WELSH. S.L., N.D. AlWOOD, S. CooDRICH. AND L.G. HlG­ other chemical chunl.cters in the Chenopodiaceal.'. CINS. 1993. A Utah £lora. 2nd edition. Print Setvices, BiochcmiCecies. P:tges 23-SO in A. Wallilt.'e. logical Monographs 63:173-198. E.D. McArthur. and M.lt Haferkamp, editors. Pro­ WOODELL,. S.R.J., AND D.H. VALENTINE. 1961. Studies in cecclings-symposium on shrub ecophysiology and the British primulas. IX. Seed inco-mp,IUbility in htott.-chnology. USDA Forest Service, Gcnt::ral Tech· diploid-autotetraploid crosscs. New Phytologist 60: nical Hepor! INT·256. Ogden, UT. 282--294. SANOGIiIiON, S.C., ANI) H.C. STUTZ. 1994. High polyploidy ZOHARY, D., ANJ) U. NUR, 1959. Natural triploids in the in Mojavean and Sonomll Desert Atriplex cCt1wscens. orchard grass D£U:tylis glomerata L, polyploid com­ American Journul ofBotany 81:1045~1053. plex and their signifieance for gene flow from diploid SANJJmt'"oN, S,C., !'LG, STUTZ, AND E.D. McAlrnnm. to tetraploid levels. Evolution 13:311-317, 1990. Geohr~ruph;c differentiation in Atriplex c.on!erti. folia. American Journal of Botany 77:490-498. Received 23 .lanuo.ry 1998 SAS INsTrruTli:. 1989. SASISTAT user's guid~. version 6. Accepted 26 Octooc-'t' 1998 4th edition, vuh.me 1. SAS Imiitute.lnc., Cary, NC. 1999J SARCOBATUS CHROMOSOME RACES 311

AwENDIX. Chromosome CQWll localities for Sarcobatus bai1.eyi and S. oenniculatus. Counts not showing a dH.le for meiotic bud oollection were made from root lips. Plants Dale of meiotic Ploidy C(lunted Nation, state, county: location bud collection Sarcohatus baU6yi 12x (1) USA, N~ Churchill Co.: roi 48, US 50,1 mi E of Sand Mountain turnoff II APR 96 12, (1) USA, NY, Churchill Co., mi 76, US SO, due NW of Eastgate 11 APR 96 l2x (1) USA, NY, Esmer.>kIa Co., oounty Iioe, US 95, S ,ide ofTooopa 15 APR 92 l2x (1) USA, NY, Lander Co., mi 1, NV 376, Big Smoy£y Valley, 15 mi S ofUS SO 15 APR 92 12K (2) USA, NY. Lander Co., mi 11, NV 376, Big Smokey Valley 25 APR 95 12x (1) USA, NY, Lyoo Co., mi 11, NV 208, 5 mi E ofSmith 12 APR 96 l2x (1) USA, NY, Lyoo Co.' roi 27,15 tni N ofYeriDgloo 12 APR 96 l2x (1) U.SA,NY, LyooCo., S'ideofSilverSprings 12 APR 96 12x (2) USA, NY, Mioernl Co., 5 mi S ofMioa 24 APR 86 12x. (1) US~ NY, Nye Co.: 5 mi N of Queen City Summit, NY 375 14 APR 92 12x (1) USA, NIT, Nye Co., tni 17, NV 379, S ofDuckwater 11 MAY 88 12, (3) USA, NIT, Nye Co., NV 267-US 95 jet, Scotty', Juoction 9 MAR 95 12x (1) USA, NY, Per,hiog Co., 10 mi W ofSulphur II APR 91 12x (2) USA, NY, Pershing Co., Thulon e,it, 1-80,10 mi S of Lovelock 12 APR 96 l2x (3) USA, NY, Washoe Co., S-S ranch turnoff, N ,ide ofWadsworth 12 APR 96 Sarcobattu venniculalus 4x (3) CAN, ALTk 1.5 mi N ofWarner 21 JUN 89, 29 JUN 91 4x (2) CAN, ALTA, 1 mi N ofAden 4JUL89 4x (1) CAN, ALTk 2 mi E ofMilk River 29 JUN 91 4x (1) CAN, ALTk 5 roi N ofMilk River 29JUN 91 4x (1) CAN, ALTA: Diuosaur Proviucial Park 5JUL83 4x (I) CAN, ALTA, N bank ofSouth Saskatchewan River, Suffield Mil. Res. 4, (2) CAN, ALTA O.ion JUL85, 4 JUL 89 4, (1) CAN, AI:fA, Peodant Orielle 4 JUL 89 4x (2) CAN, ALTA: Red Door River coulee, N of Patricia 18AUC96 4x (3) CAN, ALTA: S of41Ajct on Prov. hwy41, 5 mi E of Medidne Hat 1 AUG 95 4, (1) CAN, ALTA" Sandy Point 4x (I) CAN, ALTA: South Saskatchewan River, t~n of Bow River 5 JUL 83 4x (3) CAN, SASK, 1 km E ofMeleval, Provo hwy 13 26JUL96 4x (3) CAN, SASK: Big Muddy Valley, S ofBeogough 27 JUL96 4x (5) CAN, SASK: Cypres, Lake, Provo hwy 21 5 JUL 89, 1 AUC 95 4x (3) CAN, SASlC W Pente'" acces" Prov. hwy 13 26 JUL96 4x (2) MEX, SON, RR km 193, SO km W ofPuerto Penasco 7 SEP92 4x (4) MEX, SON, RR km222, 20 km W ofPuerto Petiasoo 13 SEP95 8x (1) USA, AZ, Apache Co., M,ny Farm' 22JUL84 8x (3) USA,. AZ, Coconino Co.: Fredonia 26 MAY 95 4x (5) USA, AZ, Maricopa Co.: Riggs Rd (Beltline) & 16th, 10 mi S of Phoenix 25AUC93 8x (2) USA, AZ, Nav,jo Co,: Holhrook 9JUN 95 4x (5) USA, AZ, Pioal Co., tni 140, AZ 287, 10 mi NW ofCoolidge 13 SEP 95 8x (3) USA, CA, Inyo Co., 10 nti NWofBishop 18 AUG 94 8, (3) USA, CA, Inyo Co.: Big Pine 18 AUC 94 8, (1) USA, CA, luyo Co., Deep Springs Valley 18 AUG 94 4, (3) USA, CA, Lassen Co., rni 13, 0.5 nti SW ofBieber 17 AUC 94, 9 JUN 95 8, (I) USA, CA, Lasseo Co., mi 89, US 395, 20 roi N ofLitchfield 16AUC94 ax (3) USA, CA, Las,en Co., Standish 16AUC94 4x (3) USA, CA, Modoc Co.: 0.5 mi WofLikely 16AUG 94 4x (11) USA, CA, Modoc Co., 2 mi W o£Davis Creek 17 AUG 94, 5 AUC 95 8, (6) USA, CA' Modoc Co., 2 nti W of Davis Creek 17 AUG 94, 9 JUN 95 8x (2) USA, CA, Modoc Co" 7 mi W ofAlturas 9 JUN 95 4, (2) USA, CA, Modoc Co., Canby 17JUL94 4x (2) USA, CA, Modoc Co" Ceoterville rd, 5 roi E ofCanby 17 AUC94 8x (3) USA, CA, Modoc Co., E ,ide ofAlkali Lake, CA 299, Cedarville 17jUL94 8, (3) USA, CA, Modoc Co.: rni 13.75, US 395, 2 rni S ofAlturas 10 JUN 95 8x (3) USA, CA, Modoc Co., Modoc Wildlife Refuge, US 395, 5 mi S ofAlturas 16 AUG 94 4x (2) USA, CA, Mono Co.: Bridgeport 18AUC94 8, (2) USA, CA, Mono Co., N side ofMono Lake, CA 167 IS AUC 94 8x (1) USA, CA, Mono Co.: S ofBeoton 27 JUL84 8x (3) USA, CA. Siskiyou Co., 0.5 mi E ofCA 161, US 'd7 jet, 3 mi NE ofDonis 17JUL94 8x (2) USA, CA, Sisl.;YOu Co.: 3 tni S ofGranada 9JUN 95 312 GRE",r BASIN NATURALIST [Volume 59

8x (3) USA, CA, Siskiyou Co.: 4 mi E ofGmnada 9JUN 95 8x (2) USA, CA, Siskiyou Co.: 5 mi S ofDorris 17jUL94 8x (2) USA, CA, Siskiyou Co.: Louie !ld., 0.5 mi E of1-5.5 !hi SW of Big Spring 9jUN 95 cH.9x (I) USA, CA, Siskiyou Co.: Louie .Rd., 0.5 mi E oft..!:), 5 mi SW ofHig Spring 9 JUN ur> Bx (I) USA, CO, Almnosa Co.: Alamosa 9AUG88 C".l. 10;< (I) USA. CO. Alamosa Co.: 51lli W ofAlamosll 9 AUG 8S 8x (3) USA, CO, Garfidd C.o.: erit 87, I-70, Rifle 27 JUL95 8x (3) USA. CO, Jackson Co.: 1 mi W ofWalden 28jUL95 8x (3) USA, CO, Jackson Co.: 2 mi N of Ilebron 28jUL95 8x (3) USA. CO, Las Anima~ Co.: TrinkL'\d 7 MAY9G 8x (2) USA.. CO, M.esa Co.: 4 mi W of Mack 27)];L95 8x (8) SA. CO, :\ioffat Co.: CO 318-<.'Ollnly rd 21 jet, Little Snake River 10 jUL 95 8x (1) USA, CO, Montezuma Co.: Aztt::c Wash 18 J'.'IAY 88 8x (3) USA, C-O, Rio Blanco Co.: 5 mi SW of M'C('ker, CO 13 27 jUL 95 8x (3) USA, 10, llannocl: Co.: Hildreth St. hrravel pit, Pocatdlu 12 jUL95 8x (1) USA, 10. Glister Co.: mi 126, US 93,15 mi N ofMackay 7jUL92 8x (2) USA, ID, Elmorc Co.: ]D 51 at N banl: ofthe Snake Hivet 13jUL9.o I" (I) US}\., MT, lleaverhc.1d en.: 6 mi E ofDell 14jUN B8 ca. lOx (J) USA, \ff, Beavcrhead Co.: 6 mi E of Dell 14 jUl' B8 4, (J) USA., Ml; Blaine Co.: Ft. Beln."1p, S !iide of ~1ilk River 5]UL89 8x (1) USA, ~n: Carbon Co.: mi 18, US 310, JO mi S ofBridger J5JUNSS 8.< (I) liSA, MT. Carfer Co.: 10 mi \V of Alz.< (5) USA, ~l), Billings C...o.: 7 mi W ofFryburg ·30 JUI, 95, ~5 JliL 00 8x (1) USA~ 'D. Billings Co.: 7 Illi W of :Fl)'burg 30JULM 4x (I) USA, ND, Billings Co.: l1ulCx!ore Roo~evelt Nl:tlll':'trk S Unit 22 JUL 87 8x (2) USA, NO, Billings Co.: Theodore Roosevelt Nat!. Park. S Unit 22. JUL 87 8x (4) USA, ND, McKenzie Co.: 0.5 mi N ofl1leo. Roos~1t NI~ N Unit entrance, "L"S 85 3OJUL95 4x (4) USA, ND, McKenzie Co.: 10 mi W ofGrassy Butte 3OJUL95 8x (3) USA, NO, McKenzie Co.: mi ]24, US 85, 4 mi S ofTheo. Ruosevelt entrdIlcc 25 JULOO 8x (I) USA, NM, Bernalillo Co.: Prince & Rio .Brdvo, Albuquc:rq\lfJ 22 MAY 96 8x (2) l:SA, NM, McKinley Co.: E side ofCaUup 26 MAY 95 cll.9x (I) USA, NM, McKinlcy Co.: E sid~ ofGallup 26 MAY 95 8x (2) GSA, N\;f, San Juan Co.: Navajo Mine ii rUN &j, 2..') JUL 85 8, (3) USA, NM, Sandoval Co.: San Ysitlm 22 MAY 95 8x (1) US."" NM, Taos Co.: Oju Calient 10 MAY 00 8x (2) USA, NM, Torranl'e Co.: Laguna tid Perro, 5 mi E 0fWillard 9 MAY 00 1999] SARCOBA'fUS CH80MOSOME RACES 313

l2x (I) USA, NM, Torrance Co.: Laguna del Perro, 5 nil E ofWillard 9 MAY 96 8x (3) USA. NM, Valencia Co.: Grants 26 MAY 95 8x (3) USA, NM, Valencia C..o.: Laguna 26 MAY 95 8x (1) USA, NY. ChurchiU Co,: Middlegatc 27 JUL 84 8x (2) USA, NY, Douglas OJ.: mi 39, US 395, Carson Valley 5AUGn5 8x (I) USA, NY, E:lko Co.: 5 mi E ofG.'1rJin 23jUN 82 8x (2) USA, NV, Elko Co.: 9 mi N of MonteUo 13 JUN 90 8x (2) USA, NV, Eureka Co.: 20 mi W of Eureka II MAY 89 8x (I) USA, NV, Eureka Co.: 8.5 OIi W ofNY 278, mi 40, 40 mi N of Eureka 1.5AlIG 94 8x (1) USA, NY, Eurcka Co.: mi 45, NV 270. 40 mi N of Eureka (short statured) 13 JUL 95 8x (2) USA, NV. Humboldt Co.: 16 mi W ofWiunemucca., NV 49 1.5 JUN gO 8x (3) USA, NY, Humboldt CO.: Denio .1.7 jULH4 8x (1) USA, NY, Humboldt Co.: OH stat.e lil1{~. McDermitt 1.4 .rUN 1J() 8x (1) USA, NY, Lander Co.: 10 mi W ofAustin II MAY 89 8, (1) USA, NY, Lander Co.: Battle Mountuin 10 MAY 89 8x (2) USA, NV, Lincoln Co.: Caliente 30 MAY 86 8x (2) USA, NY. Lincoln Co.: Dry lake, 20 mi W ofCaliente ZOAPRR9 8x (I) USA, NY, Lincoln Co.: PanaCtl 27 .rUN 86 8x (I) USA, NY, Lincoln Co.: Rachael 30 MAY 89 8x (3) USA, NV, Lyon Co.: mi Ii, NV 208, 5 mi B ofSmith 6 AUG 95 8x (I) USA. NY, Mineral Co.: 15 mi E ofHawthorne 23AP886 8x (2) USA, NY, Nyc Co.: 10 m.i N ufBentty 9 JUL82 8, (2) USA, NV, Nye Co.: 20 mi NE of lone 16jUN 90 ca.12x (1) USA, N'l, Nyc Co.: 20 mi NW ofTonopah towards Gabbs 21 APR 89 8, (1) USA, NV, Nye Co., Gabb,; 27 jUL84 8x (2) USA, NV, Nyc Co.: rni 103.5. 15 nil SWofCurrant llMAY88 8x (3) USA, NV, Washoe Co.: Wadsworth 6AUG95 8x (2) USA. NV, White Pine Co.: Cheny Creek 16 MAY 85 ca.9x (I) USA, NV, White Pine Co.: Cherry Creek 16 MAY 85 8x (I) USA, NY, White Pine Co.: US 6;50, Spring Valley 1.6 MAY 85 8x (2) USA, OR, Baker Co.: Bakel' J.4 JUL 94 8x (3) USA, OR, Grant Co.: 1 mi E ofDllyvill('1 3AUG95 8x (3) USA, OR, Grant Co.: mi 113, OR 19, 20 mi NW of Dayville 2 SEP fl4 8x (3) USA, OR, Grant Co.: mi 37, US 395, 1 mi S of Silvies 4 AUG 95 8x (2) USA. OR, Harney Co.; 1 mi E of Hurns, OR 78 4 AUG 95 8x (3) USA, OR, Harney Co.: Fields 18JIIL94 8x (3) USA. OR, Jefferson Co.: Hay Creek, 2 Ill! S ofWillowdale 16JUL94 8x (2) USA, on, Klamath Co.: N side of Klamath Falls J.7 JIIL94 8x (3) USA, OR, L'tke Co.; Alkali Lake 17jUL94 8, (3) USA, OR, Lake Co., 'Oi 88, US 395, NE of Valley Falls 18 jUL94 4x (3) USA, OR, Lake Co.: OR 140, W !tide ofLakeview 4 AUG 95 8x (3) USA, 01\, Lake Co., Plu,b 17 JUL94 8x (3) USA, OR, Malheur Co.: 5 mi Wo{Vale 19 MAY 92 8x (I) USA., OR. Malheur Co.: Owyhee Hiver Canyon, 5 mi W ofAdrian 13 SEP 94 8x (3) USA, OR, Malheur Co.: ROlTlc 18 JUL 94 8x (I) USA, OR, Morrow Co.: 1 mi SE ofLexington I SEI' 94 8x (3) USA, OR, Morrow Co.: 0.5 mi NE of Ruggs Oarge slatured} 3 AUC 95 8, (2) USA, on, Umatilla Co.: 2 mi W ofStanfield 14JUL94 8x (2) USA, OR, Union Co.; Ladd Refuge, US 30. La Grande 14jUL94 8x (2) USA, OR, Wheeler Co.: 0.5 ill; E ofOH 207-0H 19 jet E ofSpmy 3 AUG 95 8, (2) USA, SD, Fall River Co.: Hat Creek, near Ardmore 6 ]UN 89 8x (I) USA. 5D, Lyman Co.: Cedar. Cmek, mi 146, SD 1806,25 mi SE of Ft. Pierre 24 JUL96 8x \'11, USA, SD, Lyman Co,: Lower Brule 7 JUN 89 8x (I) USA, SO. Pennington Co.: 10 mi S ofVV

8. (I) USA. UT, MiUanI Co., 13 roi S ofDeserot 30 MAY 89 S. (I) USA. UT, Millard Co., 25 roi N ofMaforo (short stature