A Checklist of the Alpine Vascular Flora of the Teton Range, Wyoming, with Notes on Biology and Habitat Preferences

Great Basin Naturalist

Volume 41 Number 2 Article 11

6-30-1981

A checklist of the alpine vascular flora of the Teton Range, Wyoming, with notes on biology and habitat preferences

John R. Spence Utah State University

Richard J. Shaw Utah State University

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Recommended Citation Spence, John R. and Shaw, Richard J. (1981) "A checklist of the alpine vascular flora of the Teton Range, Wyoming, with notes on biology and habitat preferences," Great Basin Naturalist: Vol. 41 : No. 2 , Article 11. Available at: https://scholarsarchive.byu.edu/gbn/vol41/iss2/11

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]. A CHECKLIST OF THE ALPINE VASCULAR FLORA OF THE TETON RANGE, WYOMING, WITH NOTES ON BIOLOGY AND HABITAT PREFERENCES

Shaw^ John R. Spence''^ and Richard J.

Abstract.— A checkHst of the vascular flora of the alpine zone (treeless vegetation above 9500 feet or 2900 m) of the Teton Range is presented. For each of the 216 species, si.x attributes are listed: flower color and shape, pollination mode, life form, habitat preference, and whether each species is found in the Arctic. White and yellow flowered species are most common, and zoophilous species greatly predominate over anemophilous and apomictic species. Perennial/biennial herbs are the most common life form. Common habitats in the alpine zone include dry and wet meadows, bogs, debris accumulations, and cliffs and rock faces. Arctic species account for 25.9 percent of the flora. The 216 species are distributed among 111 genera and 36 families. The largest families, in order of size, are Aster- aceae, Poaceae, Cyperaceae, Brassicaceae, Rosaceae, and Scrophulariaceae.

The Teton Range, located for the most flora of the Teton Range and provide general on floral characteristics, pollina- part inside Grand Teton National Park, is a information in the alpine zone. typical fault block range. The mountains are tion modes, and habitats about 60 km long and average 6 to 9 km Phytogeographical relationships will also be In this paper, the alpine zone is wide. The major peaks in the center of the mentioned. elevation areas of treeless range and to the north are composed of Pre- defined as high lower limits arbitrarily cambrian gneisses, schists, and granites. The vegetation, with the southern peaks are capped by Paleozoic sedi- set at 9500 feet (2900 m). Although tim- Picea mentary rocks, and the divide to the west of berline (composed of Pinus alhicaulis, is general- the main peaks consists of Paleozoic and Me- engelmannii, and Abies lasiocarpa) to 10,500 sozoic rocks. The range has been extensively ly found at elevations of 10,000 in Tetons, glaciated in the past, and several small cirque feet (3050 to 3200 m) the many typical alpine spe- glaciers of the Neoglacial age exist in shel- areas below this support tered areas of the range (Love and Reed cies. Such areas include many of the cirques neoglacial deposits be- 1968, Reed and Zartman 1973). The climate in the range, and the on the floor of Jackson Hole to the east of the low the present glaciers.

Tetons is continental, with long, cold winters Methods and a short growing season. Annual mean temperature is low. Precipitation is also low, The checklist was collected from studies falling mostly as snow in the winter months by Merkle (1962, 1963), Scott (1966), Shaw (Reed 1952, Shaw 1958). (1976), Spence (1980), and Hartman and The vascular flora of the Tetons has been Lichvar (1979), supplemented by personal well documented over the last 30 years observations and herbarium materials. All (Shaw 1958, 1968, 1976). Approximately 150 species on the list can be found on deposit at species were listed as occurring in the alpine the Intermountain Herbarium (UTC) at Utah zone (Shaw 1976, unpublished data). Recent State University, the Rocky Mountain Her- intensive collecting in portions of the alpine barium (RM) at the University of Wyoming, zone (Spence 1980) and reexamination of and the Moose Herbarium in Grand Teton herbarium collections (Hartman and Lichvar National Park. Polunin (1959) and Hulten 1979) have added more than 50 additional (1968) were consulted for those species found species. in the Arctic. For each species six character- The purposes of this checklist are to in- istics are noted. An asterisk {") before the clude all the recent information on the alpine name of the species indicates it is found in

'Department of Biology, UMC 45, Utah State University, Logan, Utah 84322. 'Present address: Department of Botany, University of British Columbia, Vancouver. B.C., Canada V6T 1 W5.

232 June 1981 Spence and Shaw: Teton Plants 233

the Arctic; a dagger (f) indicates the species the world (Wardle 1978). One possible reason is characteristically svibalpine. Flower color is for the commonness of white and yellow listed as w = white, y = yellow, v = violet, flowers in the alpine zone could be that the p = pink, b = blue, r = red, br = brown, g most common and important pollinators of- = green, and o = orange. Only species with ten are generalist fly and bee species, which conspicuously colored flowers are indicated. frequently prefer such colors (Percival 1965, is = Flower shape listed as a actinomorphic, Moldenke 1976, Wardle 1978). z = zygomorphic, and is listed only for those Some difficulty was encountered with the species with conspicuously colored flowers. category Pollination Mode. Table 2 shows Pollination mode is listed as Z = zoophilous, that 75.2 percent of the species are charac- A = anemophilous or apomictic. Fryxell teristically animal pollinated. This should be (1957), Pojar (1974), Ostler and Harper considered as a maximum value, because (1978), and Swales (1979) were consulted for many of the species included in this category aid in determining pollination mode. Life are probably autogamous, especially in the form is listed as s = shrub, p = per- Brassicaceae. Not enough information on ennial/biennial herb, g = perennial /biennial breeding systems of alpine sp'^ -es is available graminoid, and a / annual herb or graminoid. yet to state definitely whethe a species out- Major habitat types are listed as 1 = dry meadows, 2 = wet meadows, 3 = bogs, 4 = crosses or is predominantly autogamous. The debris accumulations, 5 = cliffs and rock category Anemophilous/ Apomictic species faces, 6 = neoglacial deposits. For more in- includes most monocots, Artemisia, Oxijria formation on the habitat types see the dis- digyna, and the known apomicts Taraxacum cussion and Table 3. officinale and Polygonum viviparum (Fryxell 1957, Swales 1979). Some of the grasses, par- ticularly the Poa species, are probably par- Results and Discussion tially or wholly apomictic also. In the category Arctic and Alpine species Table 1 lists the 216 species of vascular 56, or 25.9 percent, are included. This can be plants and presents for each species the bio- compared with the Beartooth Range 150 km logical attributes and habitat preferences by to the north. There, 47 percent of the species family in alphabetical order within the major occur in the arctic as well (Johnson and Bil- categories Ferns, Fern Allies, and Gymno- lings 1962). The lower value for the Tetons is sperms; Dicots; and Monocots. The species possibly due to the somewhat drier condi- are arranged in alphabetical order within tions found there compared with typical each family. Table 2 is a statistical summary Rocky Mountain ranges like the Beartooths of the species and their attributes. or the Wind River Range to the southeast More than 50 percent of the species belong (Mahaney 1980). The value of 25.9 percent is to the six largest families. In addition, 78 of intermediate between the Beartooths and the species belong to the 1 1 largest genera. It northern Great Basin Ranges to the south- is interesting to note that the six families list- west of the Tetons (Billings 1978). The some- ed in Table 2 are also the six largest families what drier conditions, in the flora of Teton County (Shaw 1976). In- perhaps accentuated the explain deed, the genera Carex, Potentilla, Poa, Salix, during Hypsithermal, probably and Erigeron are also listed as being among the absence of such widespread arctic species the largest genera in Teton County. as Koenigia islandica, Gentiana algida, and White and yellow flowered species pre- Saxifraga caespitosa in the Tetons. All three dominate (64.1 percent). This result is similar species occur in the Beartooth and Wind Riv- to data presented by Ostler and Harper er Ranges (Johnson and Billings 1962, Scott (1978) in a study of plant communities in the 1966). Wasatch Mountains of Utah and Idaho. In Certain species, such as Senecio in- that study, from 65 to 75 percent of all con- tegerrimus, Carex douglasii, Mitella pen- spicuously colored flowers in several alpine tandra, and Eriophyllum lanatum, are more

communities were white or yellow. This is a characteristic of the subalpine zone in the common feature of alpine floras throughout Tetons. The inclusion of the 18 subalpine 234 Great Basin Naturalist Vol. 41, No. 2

Table 1. A list of all species occurring in the alpine zone of the Teton Range with flower color and shape, pollination mode, life form, habitat preference, and origin noted. ° indicates an arctic species; f indicates a subalpine species. The attributes are listed as 1 = flower color (y = yellow, w = white, v = violet, p = pink, b = blue, r=red, br = brown, g = green, and o = orange), 2= flower shape (a = actinomorphic, z = zygomorphic), 3 = pollination mode (A = anemophilous or apomictic, Z = zoophilous), 4 = life form (p = perennial/biennial herb, s = shrub, g = perennial/biennial graminoid, a/annual herb/graminoid), and .5 = habitat (l = dry meadows, 2 = wet meadows, 3 = bogs, 4 = debris accumulations, 5 = cliffs and rock faces, 6 = neoglacial deposits). For more detail see Methods.

Attribute Family 12 3 4 5

Ferns, fern allies, and gymnosperms

CUPRESSACEAE - 'Juniperus communis L. var. depressa Pursh - A s 6

Lycopodiaceae - - - Lycopodium selago L. p 5

Polypodiaceae - - - ° Asplenium viride Huds. p 4 - - - Athyrium filix-femina (L.) Roth p 6 Cryptogramma crispa (L.) R.Br, ex Hook. var. acrostichoides _ _ _ (R.Br.) Clarke p 4,6 _ _ _ 'Cystopteris fragilis {L.)BeTnh. p 4,6

Selaginellaceae Selaginella densa Rydb. _ _ - p 4,5,6

Dicots

Apiaceae "Bupleuriim americantim Coult. & Rose y a Z p 1,4 Cymopterus hendersonii (Coult. & Rose) Cronq. y a Z p 4,6

ASTERACEAE 'Achillea millefolium L. ssp. lanulosa (Nutt.) Piper var. alpicola (Rydb.) Garrett w ]Agoseris aurantiaca (Hook.) Greene A. glaitca (Pursh) Raf. jAnaphalis margaritacea (L.) Benth. & Hook. Antennaria alpina (L.) Gaertn. var. media (Greene) Jeps. A. microplnjlla Rydb. A. umbrinella Rydb. Arnica latifolia Bong. A. longifolia D.C. Eat. jA. nwllis Hook. Artemisia campestris L. ssp. borealis (Pall.) Hall & Clements °A. frigida Willd. °A. norvegica Fries ssp. saxatilis (Bess.) Hall & Clements A. scopulorum Gray Aster alpigenus (T. & G.) A. Gray var. haydenii (Porter) Cronq. Chaenactis alpina (Gray) M. E. Jones Cirsium tweedyi (Rydb.) Petr. 'Erigeron compositus Pursh E. leiomerus Gray E. peregriniis (Pursh) Greene ssp. callianthcmus (Greene) Cronq. E. simplex Greene E. ursinus D.C. Eat. jEriophylhim lanatum (Pursh) Forbes var. integrifolium (Hook.) Smiley Haplopappus acaulis (Nutt.) Gray H. suffruticosa (Nutt.) Gray June 1981 236 Great Basin Naturalist Vol. 41, No. 2

Table 1 continued.

Attribute Family 3 4 Ericaceae 'Arctostaphtjios uva-ursi (L.) Spreng. w Gaultlwria humifusa (Grab.) Rydb. Kalmia microphylla (Hook.) Heller PhyHodoce empetriformis (Sw.) D. Don P. glanduliflora (Hook.) Gov. Vaccinittm scoparium Leib.

Fabaceae Astragalus kentrophyta Gray var. implexus (Ganby) Barneby 'Hedysartim boreale Nutt. H. occidentale Greene 'Oxytropis campestris (L.) DG. O. deflexa (Pall.) DG. var. foliosa Hook. Barneby

Gentianaceae Gentiana calycosa Griseb.

Grossulariaceae Ribes montigenum McGlatchie

Hydrophyllaceae Phacelia hastata Dougl. var. alpina (Rydb.) Gronq.

P. sericea (Grab.) Gray

Hypericaceae \ Hypericum formosum H.B.K. var. nortoniae (Jones) Hitchc.

Linaceae °Linum perenne L. var. lewisii (Pursh) Eat. & Wright

Onagraceae 'Epilobium alpinum L. °E. latifolium L.

Polemoniaceae Linanthus nuttaUii Gray Plilox pulvinata (Wherry) Gronq.

" Polemonium pulcherrimum Hook. P. viscosum Nutt.

POLYGONACEAE Eriogonum ovalifoliitrn Nutt. var. depressum Blank. t£. imibellatum Torr. var. subalpinum (Greene) Jones 'Oxyria digyna (L.) Hill Polygonum bistortoides Pursh "P. viviparum L.

PORTULACEAE Claytonia lanceolata Pursh C. megarhiza (Gray) Parry Lewisia pygmaea (Gray) Robins. L. triphylla (Wats.) Robins. Spraguca umbeUata Torr.

Primulaceae 'Androsace septentrionalis L. Dodecatheon conjugens Greene D. pulchellum (Raf.) Merrill Primula parryi A. Gray June 1981 Spence and Shaw: Teton Plants 237

Table 1 continued.

Attri Family 3 Vol. No. 2 238 Great Basin Naturalist 41,

Table 1 continued.

Attribute

Familv ^ bracteosa Benth. y P Pedicularis ^ , w z /.pi P. contorta Benth. "^ ^ ^''"^ 'P. grocnlandica Retz. P w z Z^ P. pcirnji Gray var. purpurea Parry p b z Z b tPensYeHion attenuatus Dougl. ssp. pseudoprocerus (Rydb.) Keck. p ^ ^ F. montanus Greene P ^ whippleanus Gray -^ P P. if J v z Z Z,t)fi ° Veroriicfl (ior(»,s/c;oWii Roem. & Schult. p

Valerianaceae w a Z 1,4 Va/fn«nfl acuHloha Rydb. p

ViOLACEAE j^ v z Z p - Viola adunca ]. E. Smith

Monocots

Cyperaceae A Cflrcv albonigra MacKenz. °C. atrata L. tC. douglasii Boott. C. ehjnoides Holm tC. geijeri Boott. C. haijdeuiana Olney fC. i//ota Bailey C. hizuUna Olney C. microptera Mack. C. nardina Fries C. nigricans C. A. Mey.

C. rioffl L. Bailey C. paysonis Clokey C. phaeoccphala Piper C. pijrenaica Wahl.

fC. ro.s.sii Boott.

JUNCACEAE Juncus drummondii E. Bey.

/. mcrtervsianus Bong.

/. parriji Engelm. Luzula piperi (Gov.) Jones °L. spicata (L.) DC.

LiLlACEAE Enjthronium grandiflorum Pursh 7 ^ '^ ^ 'Lloijdia serotina (L.) Sweet. ^ 'Zigadenus elegans Pursh ^ w a Tofielda glutinosa (Michx.) Pers.

POACEAE Hitchc. Agropt/ro»i caninum (L.) Beauv. var. latiglume (Scribn. & Smith) - var. andinum (Scribn. & Smith) Hitchc. ~ A. scribneri Vasey Agrostis humilis Vasey tA. idafwensis Nash °A. scabru Willd. A. diurberiana Hitchc. A. variabilis Rydb. R. Br. ° Calamagrostis purpurascens ^ ; June 1981 Spence and Shaw: Teton Plants 239

Table 1 continued.

Attribute Family 3 4

Danthonia intermedia Vasey A Descliampsia atropurpurea (Wahl.) Scheele °D. cespitosa (L.) Beauv. 'Festuca ovina L. Leucopoa kin^ii (Wats.) Weber 'Phleum alpinum L. Poa alpina L.

P. cusickii Vasey var. ctisickii Hitchc. var. epilis (Scribn.) Hitchc. P. inctirva Scribn. & Wnis. P. nervosa (Hook.) Vasey var. wheeleri (Vasey) Hitchc. P. pattersonii Vasey P. reflexa Vasev & Scribn. P. rupicola Nash ex Rydb.

P. sandbergii Vasey 'Trisetum spicatum (L.) Richt. 240 Great Basin Naturalist Vol. 41, No. 2 with Epilobium latifolium, and Carex, Do- stretches along the Skyline Trail in the south- decatlieon, and CastiUeja species. Seeps gen- ern Tetons, and slopes in the major cirques erally support a rich mixture of Mertensia ci- consist of this habitat type. At high eleva- liata, and Carex, Mitnulus, and Soxifraga tions, this habitat is composed of scattered species, with many mosses. Ground which is patches of plants with many bare areas be- exposed late by late melting snow usually tween the patches. The top of Prospectors supports species like Claijtonia lanceolata, Mountain and Hurricane Pass are good exam- Enjthronium grandiflomm, and Ranunculus ples of this type of vegetation. The lower ele- esdischoltzii. vation and more extensively vegetated dry Dry meadows are fairly common. Areas meadows are dominated by species of the As- like Alaska Basin, parts of Moose Basin, large teraceae and Poaceae families. Astragalus

Table 2. Statistical suinmaiv of the alpine flora of the Teton Range. The families with more than 10 species and the genera with 5 or more species are also listed.

Attribute Nimiber of species Percent of total

Life form perennial /biennial herb 149 69.0 perennial /biennial graminoid 44 20.4 shrub 21 9.9 annual herb or graminoid 2 1.0

Total 216 100.0

Flower c;olor yellow 52 33.3 white 48 30.8 violet 23 14.7 pink 20 12.8 blue 8 5.1 red 2 1.3

brown / green /orange 3 2.0

Total 156 100.0

Flower sh.\pe actinomorphic 137 87.8 zygomorphic 19 12.2

Total 156 100.0

POLLLNATION MODE zoophilous 158 75.2 anemophilous/apomictic 52

Total 210

Oricln alpine but not arctic 142 arctic and alpine 56 subalpine 18

Total 216 June 1981 Spence and Shaw: Teton Plants 241 kentrophyta, Carex nigricans, and Junciis are among the richest of the habitat types in dnimmondii are also very common. At high the alpine of the Tetons. elevations the vegetation is dominated by Rock faces and cliffs are abundant in the Hijmenoxys grandiflora, Smelowskia cahj- Tetons. The most common species found in cina, Oxytropis campestris, Lloydia serotina, this habitat type include Telesonix jamesii, and Eritrichium nanum. Oxyria digyna, and various species of Saxi- Debris accumulations include taluses, scree fraga, Arabis, and Draba. slopes, and boulder fellfields. These habitats are often physically disturbed by mass move- Acknowledgments ments. Vegetation cover is usually very low. Ivan G. Palmblad and David R. Given read Many species have adapted to the disturbed and criticized an early draft of the and often xeric conditions of debris accumu- manuscript. lations, generally by producing extensive root or rhizome systems. Common species include Dryas octopetala, Oxyria digyna, Crypto- Literature Cited

B. 1973. Chronology' of cirque glaciation, gramma crispa, Senecio freinontii, Phacelia Benedict, J. hastata, Erigeron compositiis, Senecio long- Colorado Front Range. Quat. Res. 3: 584-599. Billings, \V. D. 1978. .\lpine phvtogeographv across the ifolia, and Epilobium alpinum. Great Basin. Great Basin Nat. Mem. 2:105-117. Neoglacial deposits include the full range Fry.xell, p. a. 1957. Mode of reproduction of higher of debris accumulations, plus some unique plants. Botanical Review 23:135-233. minor habitat types (Spence 1980). Charac- Hartman, R., and R. Lichvar. 1979. Rocky Mountain Herbarium, University of Wyoming, Laramie, teristic species include Oxyria digyna, Se- Wyoming 82071. Written communication to R. necio fremontii, Poa pattersonii, Juncus drurn- J. Shaw. mondii, Carex phaeocephala, and Trisetmn Hi LTEN, E. 1968. Flora of .\laska and neighl)oring terri- spicatum. Floristically, Neoglacial deposits tories. Stanford I'niv. Press, Stanford.

Table 3. The major habitat types in the alpine zone of the Teton Range, with some physical and vegetative characteristics noted for each type.

Habitat type Characteristics

Lake and stream habitat standing or moving water present.

Bog habitat water-saturated soil throughout the growing season, sometimes flooded in early season, vegetation cover usually 100 percent, with mosses common.

Wet meadow habitat ground flat to gently sloping, .soil generally moist, obvious boulders few and scattered, usually occurs near streams, lakes, seeps, bogs, or late lying snowbanks, vegetation cover generally high (often 100

percent), usually found below 10,500 ft (3200 m).

Dry meadow habitat ground flat to gently sloping, .soil generally dry, obvious boulders few and scattered, usually occurs away from lakes, streams, seeps, bogs, or late lying snowbanks, vegetation cover moderate to high at low elevations, low at high elevations.

Debris accimivilation habitat ground flat to steeply sloping, little soil, mostly accumulations of (includes taluses, scree slopes, moderate- to large-sized rock fragments, vegetation cov^r very low, and boulder fellfields) plants usually confined to crevices between rocks.

Rock face and cliff habitat mostly bedrock, generally steep (40°-!-) sometimes with ephemeral streams in early season, plants usually growing on small ledges or in cracks in the rocks.

Neoglacial deposits flat to more often steep accumulations of glacial debris, usually near existing glaciers or permanent snowbanks, vegetation cover generally low, recently formed (100-3000 years old). See Benedict 1973, Mahanev 1975. 242 Great Basin Naturalist Vol. 41, No. 2

Reproductive Johnson, P. L., and W. D. Billings. 1962. The alpine PojAR, J. 1974. dynamics of four plant vegetation of the Beartooth Plateau in relation to communities of southwestern British Columbia. 52:1819-18.34. crvopedogenic processes and patterns. Ecol. Can. J. Bot. Monogr. 32:105-135. PoLU.NiN, N. 1959. Circumpolar .\rctic flora. Clarendon D., C. Reed. 1968. Creation of the Teton Press, Oxford. Love, J. and J. History Associ- Reed, R. E. 1973. Geochronology of landscape. Grand Teton Natural J. C and Zartman. ation Press. precambrian rocks of the Teton Range, Wyom- Maha.ney, W. C. 1975. Soils of post-.\udiibon age, Teton ing. Geol. Soc. .\mer. Bull. 84:561-582. Reed, F. 1952. Vegetation of Wildlife Glacier area. Wvoming. Arct. & Alp. Res. J. Jackson Hole 7:141-153. Park, Wyoming. Amer. Midi. Natur. 48:700-729. 1980. Department of Geography, York Univ., At- Scott, R. W. 1966. The alpine flora of northwestern kinson College. 47(X) Keele Street, Downsview, Wyoming. Unpublished thesis. University of Ontario, Canada M3J 2R7. Written commu- Wyoming, 219 pp. to R. R. 1958. Vascular plants of Grand Teton Na- nication J. Spence. Shaw, J. 1962. Ecological studies of the Amphitheater tional Park. Amer. Midi. Natur. 59:146-166. Merkle, J. and Surprise Lakes cirque in the Teton Moun- 1968. Vascular plants of Grand Teton National tains, Wyoming. National Park Service Files, un- Park, Wyoming. Sida 4:1-56. published manuscript. 2.3 pp. 1976. Field guide to the vascular plants of Grand 1963. Ecological studies in the Holly Lake cirque Teton National Park and Teton County, Wyom- of the Teton Moiuitains, Wyoming. National ing. Utah State Univ., Logan. Files, 1980. of subalpine alpine Park Service unpublished manuscript, 39 Spence, J. R. Vegetation and pp. moraines in the Teton Range, Grand Teton Na- MoLDENKE, A. R. 1976. California pollination ecology tional Park, Wvoming. Unpublished thesis. Utah and vegetation types. Phytologia 34:305-361. State Univ., 207 pp. Ostler, W. K., and K. T. Harper. 1978. Floral ecology Swales, D. E. 1979. Nectaries of certain arctic and sub- in relation to plant species diversity in the arctic plants with notes on pollination. Rhodora Wasatch Mountains of Utah. Ecology 81:367-407. 59:848-861. Wardle, P. 1978. Origin of the New Zealand mountain

Percival, M. S. 1965. Floral biology. Pergamon Press, flora, with special reference to trans-Tasman Bot. 16:535-550. London. relationships. New Zealand J.