Aquatic and Semiaquatic Vegetation of Utah Lake and Its Bays Jack D

Great Basin Naturalist Memoirs

Volume 5 Utah Lake Monograph Article 5

2-1-1981 Aquatic and semiaquatic vegetation of Utah Lake and its bays Jack D. Brotherson Department of Botany and Range Science, Brigham Young University, Provo, Utah 84602

Follow this and additional works at: https://scholarsarchive.byu.edu/gbnm

Recommended Citation Brotherson, Jack D. (1981) "Aquatic and semiaquatic vegetation of Utah Lake and its bays," Great Basin Naturalist Memoirs: Vol. 5 , Article 5. Available at: https://scholarsarchive.byu.edu/gbnm/vol5/iss1/5

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 Memoirs by an authorized editor of BYU ScholarsArchive. For more information, please contact [email protected], [email protected]. AQUATIC AND SEMIAQUATIC VEGETATION OF UTAH LAKE AND ITS BAYS

Jack D. BrothersoiV

.\bstract.— Seven aquatic and seniiaquatic comniiinities surrounding Utah Lake and its bays are described. Sim- ilarities and differences in the community types are discussed. Prevalent species in each type are given. The flora contained 48.3 species, 150 of which were prevalent enough to be included in the quantitative data analysis. Dis- tichlis stricta was the most important and widespread species. Total cover varied in the communities from 10 to 77 percent. Asexual reproduction was shown to increase in importance as moisture in the soil increased. Introduced exotic species were shown to invade most successfully those habitats that show the greatest variability in moisture and/or those that have the greatest internal variation.

Initial comments on the vegetation sin- and Murphy (1951), in conjunction with stud- rounding Utah Lake were recorded as early ies of passerine birds found in the vicinity of as 23 September 1776. Fathers Atanasio Do- the lake, studied and classified the plant com- minguez and Silvestre Velez de Escalante mimities frequented by the birds on Bird Is- and their party camped on that date adjacent land and the area from the mouth of Provo to the southeast shore of the Lake, and it was River to the south end of the Provo Munici- during their stay that they penned the first pal Airport. Barnett (1964) studied waterfowl known records concerning plant communities habitat at Powell's Slough on the east shores in the area. They recorded wide meadows, of the lake. He placed the vegetation found abmidant pasture, and marsh communities on there into four major communities based the shores of Utah Lake and noted the preva- upon habitat type and plant species present. lence of poplars, willows, flax, and hemp Christensen (1965) studied two Tamarix along the streams and east side of the lake ramosissimo-Salix amygdaloides stands near (Chevez and Warner 1976). Other early visits the mouth of the Spanish Fork River and pre- were made to the area by trappers, mountain dicted that ramosissima (which he imder- men, and explorers. However, their written stood to be T. pentrandra) as a type would records yield little information on the vegeta- eventually replace Solix amygdaloides as tion of Utah Lake that is not extractable from these trees die. Foster (1968) in a statewide the Dominguez-Velez de Escalante journals. study of the major plant communities of Utah We leam from their writings of occasional recognized four community types around bogs, communities containing reeds and Utah Lake. His plant community types are abimdant marsh grasses, infrequent patches broad in definition and based on observation of wild sage, and swamps filled with Lemna rather than analytical data. Coombs (1970) and Chum (Wakefield 1933). examined the vascular aquatic and semi- More detailed studies of the plant commu- aquatic vegetation around the lake and de- nities found in and around Utah Lake have limited 29 plant communities in 7 major been made only in the past 50 years. Cottom types. Local taxonomic and ecological studies (1926) made the first quantitative studies of (e.g.. Weight 1928, Leichtv 1952, Lawler the vegetation of the lake. He listed 11 for- 1960, Bessey 1960, Arnold 1960, White 1963, mations and 20 associations that he described Skougard 1976) have been of great value by as making up the vegetation aroimd Utah identifying many of the plant species grow- Lake and adjacent Utah Valley. Wakefield ing in and aroimd the lake. (1937) reported on vegetational changes over Even though Utah Lake and its environs is a six-year period on the lakeshore south of in many localities well studied from the natu- the present Provo boat harbor. Beck (1942) ral history and ecological points of view.

'Department of Botany and Range Science. Brigham Young University, Provo. Utah 84602.

68 1981 Utah Lake M ONOGRAPH

little has been reported in the literature with Ultimately, each stand and/or community regard to (1) man's impact on the plant com- was compared to all other stands and/or mimities since settlement, (2) the influence commimities. This process resulted in the and changes wrought by introduced exotic production of interstand or intercommunity plants, (3) species composition for the major similarity index values (Ruzicka 1958). A commimity types, (4) environmental factors matrix of similarity index values was con- influencing the distribution of major commu- structed. The similarity values were clustered nity types, (5) community diversity, and (6) by the pair-group clustering procedures de- information with regard to successional scribed by Sneath and Sokal (1973). changes and life form patterns along environ- Moisture index data were assigned to each mental gradients. stand using a modification of the methods employed by Coombs (1970). Moisture class- Methods es were set up as reported in Table 2. Floristics and nomenclature follow Forty stands of data were selected from Cronquist et al. (1977) for the mon- the hterature (Coombs 1970, Bamett 1964, ocotyledons and Holmgren and Reveal (1966) Christensen 1965) and combined with 10 for the dicotyledons. stands studied by the author in the summer of 1974. Percent sum-frequency values for each Results species (Phillips 1959), total cover information (Brown 1968), and moisture index values General Vegetation Descriptions (Coombs 1970) were then assigned to all 50 The aquatic and semiaquatic communities stands. Percent sum-frequency figures were surrounding Utah Lake form a band of vege- used to give the species data from the differ- tation along the lake shore varying in width ent sources equivalent standing. Where in- from 20 m or less on the western shore to 400 formation was questionable and /or lacking m on the eastern shore. In addition, two large (especially with respect to moisture informa- bays, Provo Bay and Goshen Bay, extend tion), supplementary field observations were away from the lake in eastern and southern made in the summer of 1976. Of the stand directions, respectively, and contain a major- data taken from the literature only those hav- ity of the land area occupied by the aquatic ing relatively complete information were and semiaquatic communities. used in this analysis. During this investigaton 483 plant species Species lists (150 total) were assembled for were found to be part of the Utah Lake vege- each stand. Importance values (Warner and tation. Of these, only 150 were of sufficient Harper 1972) were then computed for each importance to include in the quantitative species in relationship to the total vegetative data analyses. Only 13 species were included complex and the major communities found in in a prevalence list for the entire area and, as the area. From this information, prevalent can be seen from Table 1, the list is highly species tables were compiled (Tables 4-9).

The number of prevalent species included on Table 1. The prevalent species found in the vegeta- any one list was equal to the mean number of tion of Utah Lake with their importance vahie. species reported for the stands of a given Scientific name commimity. The prevalents are listed in de- creasing order of importance and are the most frequent species in the community; un- common or rare species are ignored. Diversity indices (McArthur 1972) were computed from the percent sum-frequency data using the formula:

where Di is the diversity index and pi is the relative proportion each species contributes to the overall composition of a community. 70 Great Basin Naturalist Memoirs No. 5 dominated by grasses and sedges, with Dis- exclusively by a single species or clone even tichlis stricta being the most important and though the abiotic environment is homoge- widespread species. nous. Seven major vegetative types exist around Average values for selected environmental the Lake (Tables 2 and 3), each occupying variables are given for the seven major vege- unique habitats and each showing varying de- tative types in Table 2. It will be seen that grees of internal structure with respect to the number of stands considered for each

subcommunity dominants. This subcommunity is not equal, varying from 5 to community variation is related in some de- 16. The communities vary with respect to gree to the prominance of asexual reproduc- moisture from continuous inundation to sea- tion (by rhizomes) in the dominant species. sonal inundation, and finally to those that When dominant species reproduce vegeta- never experience standing water or high wa- titively, large areas may be occupied almost ter tables. Communities on the dry end of the

Table 2. Selected environmental characteristics of major plant communities surrounding Utah Lake. 1981 Utah Lake MONOGRAPH 71

Table 4. The prevalent species found in the hnlrnsh- cattail marsh communities of Utah Lake with their importance values.

Scientific name Importance values

Typha latifolia Lemna minor 6243 Scirpus actitus 5471 Berula crcrta 3457 Eleocluiris palusths 1771 Spirodchi polyrhiza 1257 Riccia fluitans 957 Polypogon monspeliensis 657 Epilobium adenocaulon 614 Lycopus Ittcidus 314 Nasturtium officinale 3(K) Scirpus americanus 286

Table 5. The prevalent species found in the semiaquatic herbaceous meadow communities of Utah Lake with their importance values.

MOISTURE INDEX Scientific name Importance values WET DRY Eleocharis palustris 9229 Fig. 1. Variation in moisture in the communities of Carex nebraskensis 4914 Utah Lake in relationship to moisture index. DistichHs spicata 2929 Scirpus americanus 2271 scale exhibit the greatest fluctuations in mois- Trifolium hyhridum 2029 Lycopus lucidus 1629 ture (Fig. 1). The amount of exposed soil var- Scirpus validus 1429 ies among the communities from less than 15 Panicum capillare 1371 percent to slightly less than 50 percent in the Polygonum coccineum 1357 playa and beach communities. Polygonum amphibium 1100 Compositional data for the seven commu- Iva axillaris 943 Plantago major 943 nity types is given in Table 3. Each commu- Ambrosia artemisifolia 800 nity is life- dominated by a different set of Agrostis alba 771 form types, with annuals being especially Bidens eernua 714 prevalent in the playa and beach areas. In Polypogon monspeliensis 614 Xanthium strumarium 557 only two cases do particular life form types become sufficiently abundant to contribute In the cluster diagram, communities that are over 50 percent of the plant cover. General- most similar appear close together. The hori- ly, the vegetation of the communities consid- zontal line connecting any two communities ered includes species from several life form shows the degree of similarity between those classes. entities. Figure 2 demonstrates that each Diversity measurements varied from low community recognized is highly unlike all values for pond communities to high weed other commimities considered. The most sim- values for the annual herbaceous commu- ilar entities are the Spikerush-bulrush mead- nities. However, even though the diversity ows and the Grass-rush-sedge meadows, indices varied considerably, no significant which are only 25 percent similar. Other sim- correlations could be established between di- ilarity patterns exist, but the similarity per- versity and other parameters. cents are so low that the community types in- Similarities between the seven community volved can be considered essentially types are evident since some species show independent of each other. dominance in more than one type (Tables 4-9). To better understand these inter- Community Type Descriptions relationships and to assess the degree of uniqueness of the different community types Pond Weed Communities

(Tables 2 and 3), a graphical summary of in- The pond weed communities are contin-

tercommunity similarity is presented (Fig. 2). uously inundated by water. They are essen- 72 Great Basin Naturalist Memoirs No. 5

Table 6. The prevalent species found in the grass- Table 9. The prevalent species found in the annual rush-sedge meadow communities of Utah Lake with herbaceous communities of Utah Lake with their impor- their importance values. tance values.

Scientific name Importance values Scientific names

Distichlis spicata 7206 Scirpus americanus Juncus balticus Eleocharis palustris Hordeum jiibattim Carex nebraskensis Sporobolus aeroides Glaiix maritima Ambrosia ortimisifolia Potentilla anserina Lycopus hicidus Trifolitim hybridum Pohjpogon monspeliensis Ranunculus cymbalaria Phntago major Iva axillaris Aster brachyactis Suaeda occidentalis

Table 7. The prevalent species found in the lowland woody communities of Utah Lake with their importance values. 1981 Utah Lake Monograph 73

% SIMILARITY

^ SPltCERUSH- BULRUSH MEADOWS ^ GRASS RUSH -SEDGE MEADOWS „ LOWLAND WOODY COMMUNITIES J^ ^ ANNUAL HERBACEOUS COMMUNITIES

, ^ BULRUSH CATTAIL MARSHES

. o. SALINE TERRESTRIAL COMMUNITIES

, ^ PONDWEED COMMUNITIES

Fig. 2. Community similarity analysis reported as a cluster diagram based on plant composition of the Utah Lake commimities. of the lake, but reaches maximum devel- ship appears to be associated with a water opment m Provo Bay and Powell's Slough. gradient in which moisture increases as one Whether Scirpus acutiis or Typlia latifolia moves toward areas dominated by Scirpus dominates any particular marsh seems to be validus (Coombs 1970). Again one sees local largely a matter of priority, according to areas dominated by single species that repro- Cottam (1926). This observation tends to sup- duce vigorously by asexual processes. port the concept that the subcommunities de- fined by Coombs (1970) can be partially ac- Grass-Rush-Sedge Meadows counted for by patterns in asexual The grass-rush-sedge meadows inhabit the reproduction of the dominant species. largest area of any of the semiaquatic herbaceous communities described thus far. Spikerush-Biilmsh Meadows They are situated geographically much like The spikcRish-bulrush meadow commu- the spikerush-bulrush meadows, but tend to nities are generally situated in areas that are differ in at least the following ways: (1) al- inmidated in the early seasons of the year but though seasonally saturated the excess water dry by September. The soil of the community has generally drained away by late spring, (2) varies but generally consists of peaty sandy the soils generally are less peaty, and (3) the loams (Coombs 1970). Organic matter con- soils are often slightly to moderately saline. tent of the soil is high and, in places, the This community shows the greatest intercommunity occurs on peat beds that are 30 nal variation and as a result exhibits the high- inches deep. The type averaged 17 species est mean diversity value (Table 3), which is per stand; several species share dominance. exceeded only by the annual herbaceous The two most important species are Eleo- communities. The community averaged 18 charis macrostachya and Carex nebrascensis species per study unit and is the only commu-

(Table 5). The community is restricted to the nity dominated by grass (Table 6). Of the 8 eastern side of the lake extending from near most important species, 6 are considered to White Lake in Goshen Bay to the Jordan be salt tolerant. The community is extensive

River, but is best developed in Benjamin's (found throughout the study area) and often Slough and Provo Bay. The major component occupies sites lying between upland shrub species appear to distribute themselves in types and the communities already described. predictable ways in space— as subdominants There is a great deal of subdominant varia- of the community. Scirpus validiis for ex- tion within the type that appears to reflect ample, is often found in nearly pure stands patterns of asexual reproduction on the one surrounded by mixed zones of Eleochraris hand and islands of local habitat variation on macrostachya and Carex nebraskensis. These the other (i.e., pockets of peat loam soil dom- latter species generally give way to areas inated by Carex nebraskensis, etc.). Again, dominated by Distichlis stricta. The relation- the major dominants and subdominants segre- 74 Great Basin Naturalist Memoirs No. 5

gate along a moisture gradient. The sedges thus far. It is essentially confined to Ben- {Scirpus americanus, Eleocharis macro- jamin's Slough, Goshen Bay, and surrounding stachya, and Carex nehraskensis) tend to be areas. The soils vary from sandy clay loams to dominant on those areas of seasonal in- heavy clays and are generally poorly drained undation, and the grasses {Distichlis stricta, and alkaline or saline in nature. Soil erosion

Hordetim fubatum, and Sporoholus aeroides) is often evident and disturbance from several

tend to dominate the higher dryer areas. sources is generally apparent. Salt content in the soil varies greatly in both lateral and ver- Lowland Woody Communities tical space. Variation in salinity combines

The lowland woody community is a broad- with variation in soil moisture and local to- ly scattered type occupying a variety of dis- pography to produce small scale hetero- jimct sites about the lake. It is among the geneity in the vegetation. The soils in many three most extensive communities surround- areas are seasonally wet, but the communities

ing the lake and is found most often in sea- are not required to develop under water. sonally submerged sites often near flowing Small drainage basins are scattered streams. The soils are predominantly mineral throughout the type and act as receptacles of (sandy to sandy clay loams) with varying de- spring nmoff. As the trapped water evapo- grees of incorporated organic matter. The rates from these catchment basins, salts and community averaged only 9 species per stand other materials carried there by the water are left behind. Salt pans or playas develop in (Table 7) and yielded one of the lowest diver- such areas. It is around such playa areas that sity indices (Table 3). Of the woody domi- majority of the vegetational variation is nants listed, 3 are shrubs and 2 are trees. a variation is for con- There are two layers in the community, the found. This accounted by tree-shrub overstory and a grass-annual or centric rings of vegetation that surround the aquatic herb understory. The aquatic her- playas. Terrestrial saline communities are low in species diversity (Table 3) and average baceous understory is important only in areas only five species per stand (Table 8). Of the where willows are dominant. There is a high dominants listed, all are salt tolerant and two degree of subdominant variation and internal (i.e. Kochia americana and Suaedo nigra) are heterogeneity in the community. However, considered to be disturbance indicators in this case, as opposed to previous described (Coombs 1970). types, the majority of the variation is due to habitat differences rather than asexual repro- Annual Herbaceous Communities ductive patterns. The annual herbaceous type is a con- Tamarix ramosissima and Elaeagniis ang- glomeration of several terrestrial commu- ustifolia, two of the most important species nities that occupy waste places around the listed (Table 7), are exotic invaders. Since lake. These areas often have little in common they occur in the overstory and since T. and exhibit high variability in environment ramosissima is the most widely distributed and species composition. Because of great en- plant in the type, it appears that this type has vironmental variability and regular disturb- been more extensively modified by human ance, such as along beaches, seasonally in- activities than any other community consid- undated islands, and areas heavily impacted ered here. considered both Coombs (1970) by the activities of man, the communities of- species to be increasing suggested that and ten remain in early serai stages of succession. much of the woodland is in vari- community This is evidenced by the fact that most of the ous stages of recovery disturbance. If from dominant species (Table 9) are of the annual his evaluation is accurate, it appears that the life form, a life-style that permits plants to woodland community will undergo a great complete their life cycle in a few months.

deal of change in the future. Since variation is great and conditions change from year to vear, patterns in species Saline Terrestrial Community dominance also fluctuate annually. Stability The saline terrestrial community is the will only come to these communities as envi- most geographically restricted type discussed ronmental predictability increases. 1981 Utah Lake Monograph 75

Ecological Relationships reproduction and against species incapable of such reproduction methods. Total Cover Total cover in the communities surround- Intraconirnunity Similarity ing Utah Lake varies from 9.8 percent in the Earlier in this paper reference has been pondweed sites to 76.6 percent in the Grass- made of the subcommunity (within) varia- Rish-sedge meadows (Table 3). Observed dif- tions in each of the seven major community ferences appear to be related to variations in types. Such internal variations can be mea-

moisture (Fig. 3). As seen in Figure 3, the sured with similarity indices. I have com- largest cover values occur midway along the puted a similarity index matrix (Runzicka moisture gradient in communities that tend 1958) utilizing all stands in each community. to exhibit the most favorable soil moisture re- Thus, the similarity of each stand with all gimes. When there is either too much mois- other stands of a commimity is obtained. All ture (year-round inundation) or too little similarity indices in each community matrix moisture (dry upland sites), fewer plants ap- is finally averaged to obtain a mean and stan- pear to perform well, thus lowering cover dard deviation for internal similarity of each values in these areas. community type. The larger the value the

more internally similar is the community; Asexual Reproduction conversely, the lower the value the greater As previously suggested, much of the sub- the internal variability. Variation in in- community variation with the aquatic and tracommunity similarity is plotted against semiaquatic communities of Utah Lake can variation in available moisture for growth in be related to asexual reproduction by domi- Figure 5. Intracommunity variation is seen to nant species. This seems especially true in increase as moisture variability increases. those communities that are continuously or This indicates that as habitat predictability seasonally inundated for long periods. Figure decreases, the composition of communities 4 illustrates this relationship. Communities occupying such habitats also becomes more having dominant species that reproduce asex- variable and less recognizable as distinct en- ually are also those communities common to tities. the wet end of the moisture gradient. This being the case, it appears that those habitats Life Forms with the most uniform moisture conditions The relationship of plant life forms to envi- tend to select for species capable of asexual ronmental factors has been the concern of

< i o 111 ^ a.

is *°-

25 " 20. o o

MOISTURE INDEX MOISTURE INDEX WET DRY WET DRY Fig. 4. Importance of asexual reproducing species in the Utah Lake communities as moisture becomes less Fig. 3. Total living cover in the Utah Lake commu- available. nities in relationship to changing moisture conditions. 76 Great Basin Naturalist Memoirs No. 5

^ 2.0 < 30_

< i 1.0

.2 MOISTURE INDEX MOISTURE VARIATION WET DRY

Fig. 5. Variation in internal community similarity as Fig. 7. Importance of sedges in Utah Lake commu- moisture variation increases. nities n relationship to changing moisture conditions.

o iti £ 20_

MOISTURE INDEX MOISTURE INDEX WET DRY WET DRY

Fig. 6. Importance of grasses the Utah Lake com- Fig. 8. Importance of annuals in the communities of munities in relation.ship to changing moisture condi- Utah Lake in relationship to changing moisture conditions. tions.

ecologists for many years. The life form con- in habitats with moisture regimes midway cept was useful in this paper in delimiting along the gradient (Fig. 6). In contrast, the

community types (i.e., grass-ru.sh-sedge mead- sedges are most abundant at the higher mois- ows, lowland woody communities, or annual ture levels (Fig. 7). Annuals reach their great- herbaceous communities). The concept also est importance in the driest habitats (Fig. 8). helps relate environmental pattern to plant With respect to annuals, the relation.ships de- response in the habitat complex of Utah Lake picted by Figures 9 and 10 are also of inter- (Table 3, Figs. 6-10). The data demonstrate est. As shown, the annual life form does espe- that some of the life form classes exhibit cially well in habitats that are open, low in rather distinct responses to moisture patterns cover, and support a good deal of exposed around the lake. Grasses, for example, do best .soil. In such areas, interspecific competition 1981 Utah Lake Monograph 77 78 Great Basin Naturalist Memoirs No. 5

Table 10. Plant families contributing the majority of species to the flora of Utah Lake.

Family Percent of speci

Asteraceae 16.7 Poaceae 14.5 u 40 Cyperaceae 6.3 Chenopodiaceae 5.9 Cruciferae 5.9 Leguminosae 3.9 Polygonaceae 2.9 Rosaceae 2.9 Labiatae 2.5 Salicaceae 2.2 Scrophulariaceae 2.2 Onagraceae 2.0

Total 67.9

MOISTURE VARIABILITY

Fig. 12. Importance of introduced species in the com- that allow a species entering from the outside munities of Utah Lake as moisture variability increases. to become established and compete successfully. These gaps would almost certainly arise as a result of interaction between moisture

variability and the resultant effect it has on internal community structure.

30- Floristic Relationships

A total of 483 species of vascular plants, representing 275 genera, and 74 families was observed and/ or found recorded as belonging to the plant communities of Utah Lake. Of these, 67.9 percent belonged to 12 plant families (Table 10). The ecological or phyto- geographical significance of the dominance

of these families (Table 10) is not known, but .5 .1jO further investigations along such lines should hold great interest. VARIATION IN INTERNAL SIMILARITY OF COMMUNITY TYPES ACERACEAE Fig. 13. Importance of introduced species in the Utah Acer grandidentatum Nutt. Lake communities as internal community similarity in- Acer negundo L. creases. AlZOACEAE conditions surrounding the lake are consid- Sesuvium verrucosum Raf. ered, two important relationships emerge (Figs. 12 and 13). First, as shown in Figure Alismataceae Alisma triviale Pursh 12, the introduced species reach their great- Sagittaria cuneata Sheld. est development in those habitats that show the greatest variability in moisture (the most Amaranthaceae unpredictable environments); and second Amaranthus graecizans L. Amarantlnis retro flextis L. (Fig. 13), those communities having the greatest internal variation in composition Anacardiaceae tend to be the most easily invaded. Undoubt- Rhus radicans L. edly, such communities have structural gaps Rhus trilohata Nutt. 1981 Utah Lake Monograph 79

Apocynaceae Machaeranthera tanacetifolia (HBK.) Ne.ss Apoctjninn cdnncihinimi L. var. glahcrriimttn A. DC. Matricaria matricarioides (Less.) Porter Scnccio lujdrophilus Nutt. Senecio uintahcnsis (A. Nels.) Greene ASCLEPIADACEAE Solidago canadensis L. Asclepias incarnata L. Solidago occidentalis (Nutt.) Torr. & Gray Asck'pias speciosa Torr. Sonchus arvensis L. Sonchus asper (L.) Hill Stephanomeria pauciflora (Torr.) Nutt. ASTERACEAE Tanacetum vtdgare L. L. Achillea millefolium Taraxacum officinale Weber Ambrosia artemisiifolia L. Tctradymia glabrafa A. Gray Ambrosia psilostachya DC. Tctradyniid spinosa Hook. & Arn. Antheinis cotula L. Townscndia florifcr (Hook.) A. Gray Arctium minus Schk. Townsendia strigosa Nutt. Artemisia absinthium L. Tragopogon dubius Scop. Artemisia dracunctilus L. Tragopogon porrifolius L. Nutt. Artemisia hidoviciana Viguiera ciliata (Robins. & Greenm.) Blake Artemisia spinescens D.C. Eaton Viguiera multiflora (Nutt.) Blake Artemisia tridentata Nutt. Wyethia amplexicaulis (Nutt.) Nutt. Aster brachyactis Blake Xanthium strumarium L. Aster chilensis Nees ssp. adscendens (Lindl.) Cronq. Xanthocephalum sarothrae (Pursh) Shinners Aster eatonii (A. Gray) Howell Aster frondosus (Nutt.) Torr. & Gray Betulaceae Aster perelegans A. Nels. & Macbr. Alnus tenuifolia Nutt. Balsamorhiza hookeri Nutt. Betula occidentalis Hook. Bidens cernua L. Bidens frondosa L. Boraginaceae Chaenactis douglasii H. & A. Cryptantha flavoculata (A. Nels.) Payson Chrysopsis villosa (Pursh) Nutt. var. foliosa (Nutt.) Cryptantha nana (Eastw.) Payson D.C. Eaton Cynoglossum officinalis L. Chrysothamnus nauseosus (Pall.) Britt. Heliotropium curassavicum L. Chrysothamnus viscidiflorus (Hook.) Nutt. Lappida redowskii (Hornem.) Greene Cichorium intybiis L. Lithospennum ruderale Doug, ex Lehm. Cirsium arvense (L.) Scop. Plagiobothrys scouleri (Hook. & Arn.) I.M. Cirisium foliosum (Hook.) DC. Cirsium undulatum (Nutt.) Spreng. Cirsium vulgare (Savi) Airy- Shaw Cactaceae Conyza candensis (L.) Cronq. Echinocactus simpsonii Engelm. Crepis modocensis Greene Echinocereus triglochidiatus Engelm. var. melana- Crepis runcinata (James) Torr. & Gray canthus (Engelm.) L. Benson Erigeron bellidiastrum var. typicus Cronq. Opuntia fragilis (Nutt.) Haw. Erigeron divergens Torr. & Gray Opuntia polycantha Haw. Erigeron glabellus Nutt. Erigeron lonchophyllus Hook. Capparidaceae Eupatorium maculatum L. Cleome lutea Hook. Franseria acanthicarpa (Hook.) Gov. Cleome serrulata Pursh Gnapluiliimi chilense Spreng. Polanisia dodecandra (L.) DC. Gnaphalium patustre Nutt. Grendelia squarrosa (Pursh) Donal Caprifoliaceae Haplopappus lanceolatus (Hook.) Torr. & Gray Lonicera involucrata (Rich.) Banks Haplopappiis watsoni A. Gray Helenium autumnale D.C. Eaton Caryophyllaceae Helianthus annuus L. Cerastium vulgatttm L. Helianthus nuttallii Torr. Gray & Saponaria officinalis L. Helianthus petiolaris Nutt. Spergularia marina (L.) Griseb. Hieracium gracile Hook. Hymenoxys acaulis (Pursh) Parker Ceratophyllaceae Inula helenium L. Ceratophyllum demersum L. Iva axillaris Pursh Iva xanthifolia Nutt. Lactuca pulchella (Pursh) DC. Chenopodiaceae occidentalis (S. Wats.) Kuntze Lactuca scariola L. Allenrolfea Atriplex (Torr. Frem.) S. Wats Laphamia stansburii A. Gray confertifolia & Atriplex heterosperma Bunge iMyia glandulosa (Hook.) Hook. & Am. Atriplex hortensis L. Lygodesmia grandiflora (Nutt.) Torr. & Gray 80 Great Basin Naturalist Memoirs No. 5

Atriplex patula var. hastata (L.) A. Gray Stdnleyella wrightii (A. Gray) Rydb. Atriplex tridentata Kuntze Streptanthus eordatus Nutt. ex Torr. & Gray (Pursh) T. Ceratoides lanata J. Howell Tltehjpodium sagittatum (Nutt.) Endl. Chenopodiwn album L. Chenopodium chenopodiodes (L.) Aellen Cupressaceae Chenapodium fremontii S. Wats. Juniperus osteosperma (Torr.) Little Chenopodium gigantospennum Aellen Chenopodium glaucum L. Cyperaceae Chenopodium leptophyUum Nutt. Carex aurea Nutt. Chenopodiwn murale L. Carex aquatilis Wahl. Chenopodium wat.soni A. Nels. Carex atherodes Spreng. Corispermum viUosum Rydb. Carex lanuginosa Michx. Echinopsilon hyssopifolium (Pall.) Moq. Carex nebraskensis Dewey Grayia spinosa (Hook.) Moq. Carex petasata Dewey Hologeton glomeratus (Bieb.) May. Carex praegracilis W. Boott. Kochia americana S. Wats. Cyperus erythrorhizos Muhl. Kochia scoparia (L.) Schard. Cyperus strigosus L. Monolepis nuttalliana (Schult.) Greene Eleocharis acicularis (L.) Roem. & Schult. SaUcornia pacifica Standi. Eleocharis bolanderi A. Gray Salicornia rubra A. Nels. Eleocharis palustris (L.) Roemer & Scultes Salsola iberica Senner & Pan. Eleocharis parvula (Roem. and Schult.) Link. var. col- Sarcobatus venniculatus (Hook.) Torr. oradensis (Britton) Beetle Suaeda depressa (Pursh) S. Wats. Eleocharis pauciflora (Lightf.) Link. Suaeda fruticosa (L.) Forsk. Eleocharis rostellata Torr. (Raf.) F. Suaeda nigra J. Macbride Fimbristylis spadicea (L.) Vahl. Suaeda occidentalis S. Wats. Scirpus acutus Muhl. Scirpus americanus Pers. Scirpus lacustris L. CONVOLVULACEAE Scirpus maritimus L. Convolvulus arvensis L. Scirpus microcarptis Presl. Convolvulus sepium L. Scirpus pallidus (Britton) Fernald Cressa truxillensis H.B.K. Scirpus validus Vahl. Cuscuta salina Engelm. Dipsacaceae Dipsacus sylvestris Huds. CORNACEAE Cornus stolonifera Michx. Elaeagnaceae Elaeagnus angustifolia L. Shepherdia argentea (Pursh) Nutt. Cruciferae Arabis glabra (L.) Bernh. Ephedraceae Arabis holboellii Hornem. Ephedra viridis Coville Brassica campestris L. Brassica kaber (D.C.) Wheeler var. pinnatifida Equisetaceae Brassica nigra (L.) Koch Equisetum ancnse L. Camelina microcarpa Andrz. Equisettim kansunum Schaffn. Capsella bursa-pastoris (L.) Medic. Equisetum hnvigatum \. Br. Cardamine pennsylvanica Muhl. ex Willd. Equisetum palustre L. Cardaria draba (L.) Desv. Conringia orientalis (L.) Diimort EuPHORBlACEAE Descurainia pinnata (Walt.) Britt. Euphorbia glyptospenna Engelm. ex Emory Descurainia sophia (L.) Webb. Euphorbia serpyllifolia Pers. Erysimum capitatum (Dougl.) Greene

Erysimum iiicon.spicuum (S. Wats.) Mac M. FUMARIACEAE Erysimum rcpandum L. Corydalis aurea Willd. Hutchinsia procumbens (L.) Desv. Lepidium densiflorum Schrad. Gentian ACEAE Lcpidium densiflorum var. ramosum (A. Nels.) Thel Centaurium exaliaium (Griseb.) Wight Lepidium moutdiuim Niitt. Lepidium perfoliatum L. Lepidium virginicum L. Geraniaceae Erodium (L.) Malcolmia africana (L.) R. Br. cicutarium L'Her. Nasturtium officinale R. Br. in Ait. Physaria australis (Payson) Rollins Haloracac:eae Rorippa islandica (oed.) Borbas Hippurus vulgaris L. Sisymbrium altissimum L. Myrioplu/tlum spicatum L. 1981 Utah Lake MoN()(;aAPn 81

HvDlUXIlAKirACE.VK LOASACIEAE Elodcd caiuKlcnsis Midix. Mentzrha iilbirauhs Dougl. ex Hook. Mentzcha decapetala (Pursh) Urb. & Gilg. Mentzcha lacvicauhs (Dougl.) Torr. Gray IkIDAC KAK & Mentzeha niultifhra (Nutt.) \. Gray Susynnchiiun iKilopluliiiii C.reene

Lythraceae JlNCACEAK Lythrum sahcaria L. Jiinciis hdlticus W'illd. ]uu(us htifoiiiiis L. Malvaceae Jumus cnsijolius Wikstr Ahhaea rosea Cav. Juruus lou'^istylis Torr. Muha ncgU'cta Wallr. Jiiiuiis tonciji Coville Sida ludcrarcd (Dougl.) Torr. Sidalcca ncomcxicana A. Grey JUNCAGINACEAE Sidalcea oregana (Nutt.) A. Gray Triglochin mahtiina L. Sphaeralcea coccinea (Pursh) Rydb. Sphaeralcea grossiilariaefolia (H. & A.) Rydb. Labiatae Sphaeralcea munroana (Dougl.) Spach Lamium mnplexicaulc L. Lijroptis (imcriconus Miihl. ex Bart. Moraceae Ltjcopiis Iticidus Turcz. Morus rubra L. Marrubium vulgare L. Mentha arvensis L. Nyctaginaceae Mentha spicata L. Abronia salsa Rydb. Mohlaiira parviflora (Niitt.) Britt. Nepcta cataria L. Nymphaeceae Stachys pahistris L. NupJiar pohjsepahtm Engelm. Teucrium canadense L. var. occidentale (A. Gray) McClintock & Epling Oleaceae Fraxinus vehitina Torr. Leguminosae Onagraceae Astragahis argophyUus Nutt. var. argophyUus Epilobium adenocaulon Hausskn. Astragahis heckwethii Torr. & Gray Epilobium paniculatum Nutt. ex Torr. & Gray Astragahis canadensis L. Gaura parviflora Dougl. Astragahis convallarius Greene Oenothera ahjssoides Hook. & .\rn. Astragahis oophorus S. Wats. Oenothera caespitosa Nutt. Astriigahts utahensis (Torr.) Torr. & Gray Oenothera hookeri Torr. & Gray Ghjcyrrhiza lepidota Pursh Oenothera latifoha (Rydb.) Munz Hedysanim boreale Nutt. Oenothera minor (A. Nels.) Munz Lathyrus brachycahjx Rydb. Oenothera paUida Lindl. Mcdicago hiptihna L. Oenothera scapoidea Torr. & Gray ssp. utahensis Mcdicago sativa L. Raven Mehlottis alba Descr. Mehlotiis officinahs (L.) Lam. Orchidaceae Robmia pseudo-acacia L. Cypripedium calceohis L. var. pubescens (Willd.) Thcnnopsis montana Nutt. Cornell Trifohum hyhridum L. Epipactis gigantea Dougl. Trifohiini pratense L. Spiranthes romanzoffiana Cham. & Schl. Trifohum repens L. Vicia americana Muhl. var. minor Hook. Orobanchaceae Orobanche nndtiflora Nutt. Lemnaceae Lemna minor L. Papaveraceae Lemna trisulca L. Argemone munita Dur. and Hilg. Lemna valdiviana Phil Spirodela polyrliiza (L.) Schleid. Plantaginaceae Plantago lanceolata L. Lentibulariaceae Plantago major L. Utricularia minor L. Plantago patagonica Jacq.

Liliaceae POACEAE AUiiim acuminatum Hook. Agropyron cristatum (L.) Gaertn. A.^)aragus officinahs L. Agropijron dasystachyum Scribn. (Hook.) Smilacina stellata (L.) Desf. Agropyron elongatum (Host.) Beauv. 82 Great Basin Naturalist Memoirs No. 5

Agropyron intermedium (Host) Beauv. POLEMONIACEAE Agropyron repens (L.) Beauv. Collomia linearis Nutt. Agropyron smithii Rydb. Gilia aggregata (Pursh) Spreng. Agropyron spicatum (Pursh) Scribn. & Smith Gilia inconspicua (Smith) Sweet Agropyron trachycaulum (Link) Malte. Gilia leptomeria A. Gray Agrostis semiverticillata (Forsk.) Gilia tenerrima A. Gray Agrostis stolonifera L. Phlox austromontana Goville Alopecurus aequalis Sobol. Phlox longifolia Nutt. micranthum Benth. Avena fattia L. Polemonium occidentale Greene Avena sativa L. Polemonium Beckmannia syzigachne (Steud.) Fern. Bromus commutatus Schrad. POLYGONACEAE Nutt. Bromus inermis Leyss. Eriogonum effusum Eriogonum racemosum Nutt. Bromus tectonim L. Eriogonum umbellatum Torr. Calamagrostis canadensis (Michx.) Beauv. Polygonum amphibium L. Calamagrostis neglecta (Ehrh.) Gaertn. Mey & Schreb. Polygonum aviculare L. Catabrosa aquatica (L.) Beauv. Polygonum coccineiwi Muhl. ex Willd. Cenchnis tribidoides L. Polygonum convolvulus L. Dactylis glomerata L. Polygonum lapathifolium L. Deschampsia caespitosa (L.) Beauv. Polygonum pennsylvanicum L. Distichlis spicata (L.) Green Polygonum persicaria L. Echinochloa crusgalli (L.) Beauv. Polygonum ramosissimum Michx. Elymus canadensis L. Michx. Rumex crispus L. Elymus cinereus Scribn. & Merr. Rumex fueginus Phil. Wilhams Elymus simplex Scribn. & Rumex venosus Pursh Elymus tritiocides Buckl. var. submuticus Hook. Elymus virginicus L. PORTULACACEAE cilianensis (All.) Mosher Eragrostis Portulaca oleracea L. Eragrostis hypnoides (Lam.) Britton, Sterns, & Poggenb. POTAMOGETONACEAE Eragrostis orcuttiana Vasey Potamogeton crispus L. Festuca pratensis Huds. Potomogeton filiformis Pars. Glyceria grandis S. Wats. Potamogeton foliosus Raf. Hordeum brachyantherum Nevski Potamogeton nodosus Poir. ex Lam. Hordeum jiibatnm L. Potamogeton pectinatus L. leporinum Link. Hordeum Potamogeton praelongus Wulf. Leersia oryzoides (L.) Swantz Leptochloa fascicularis (Lam.) A. Gray Primulaceae Lolium multiflorum Lam. Dodecatheon pulchellum (Raf.) Merrill Muhlenbergia asperifolia (Nees & Meyen) Parodi Glaux maritima L. Oryzopsis hymenoides (R. & S.) Riker Steironeyna ciliatum (L.) Raf. Panicum capillare L. Panicum capillare L. var. occidentale Rydb. Ranunculaceae Phalaris arundinacca L. Delphinium andersoni \. Gray Phleum pratense L. Ranunculus acris L. Phragmities australis (Cav.) Trin. ex Stendel Ranunculus acjuatilis L. capdkiceus (Thuill.) DC. Poa annua L. Ranunculus circinatus Sibth. Poa navadensis Vasey ex Scribn. Ranunculus cymbalaria Pursh Poa pratensis L. Rauuncuhis macounii Britton Polypogon monspeliensis (L.) Desf. Ranunculus orcogcnes Greene Puccinellia nuttaUiana (J. A. Schuites) ,\.S. Hitchc. Ranunculus tcsticulatus Grantz Sclerochloa dura (L.) Beauv. Secale cereale L. Rosaceae Setaria glauca (L.) Beauv. Amelanchicr alnifolia (Nutt.) Nutt. Setaria viridis (L.) Beauv. Amelanchicr utalicnsis Koehne Sitanion hystrix (Nutt.) G. Smith J. Cowania mcxicana D. Don Sitanion jubafum G. Smitli J. Crataegus douglasii Lindl. var. rivularis (Nutt.) Sarg. Hparliua gracilis Trin. Potentilla anserina L. Sphenopholis obtusata (Miciix.) Scriiin. Potentilla biennis Greene Sporobolus airoides (Torn) Torr. Potentilla ghmdutosa Lindl. Sporoholtis asper (Michx.) Kunth Potentilla gracilis Dougl. var. clmcri (Rydb.) Jeps. Sporobolus cryptandrus (Torr.) A. Gray Potentilla paradoxa Nutt. Stipa coinata Trin. & Rupr. Primus americana Marsh Triticum acstivum L. Prunus virginiana L. var. mclanocarpa {\. Nels.) Sarg. Vulpia octoflora (Walter) Rydb. 1981 Utah Lake Monograph 83

Puishia tmlcntata (Pursh) DC. Ulmaceae Rosa nutkana Presl. Celtis reticulata Torr. Rosa woodsii Lindl. Ulinus americana L. Vhnus puinila L. RUBIACKAE Galium trifkhiiii L. Umbelliferae Berulu erecta (Huds.) Coville Rl'PPIACEAE Cicuta douglasii (DC>'.) Coult. & Itose Rttppia maritima L. Coniuni maculatum L. Pastinaca sativa L. Salicaceae Slum suave Walt. Populus alba L. Populus angustifolia James Urticaceae Populus dcltoides Bartr. Urtica dioica L. var procera (Muhl.) Wedd. Populus fremontii S. Wats. Urtica serra Bhune Poptilus nigra L. var. italica Muenchh. Populus trichocarpa Torr. & Gray Verbenaceae Salix amiigdaloides Anders. Verbena bracteata Lag. and Rodr.

Sa/i.v cxigua Nutt. Verbena hastata L. Salix fragilis L. Verbena stricta Vent. Salix lasiandra Benth. Salix rigida Miihl. Violaceae Viola nephrophylla Greene Salviniaceae Azolla caroliniana Willd. Zannichelliaceae Salvinia rotundifolia Willd. Zannichellia palustris L.

Santalaceae Zygophyllaceae Comandra pallida A. DC. Tribulus terrestris L.

Saxifragaceae Species included in the literature as being Heuchera pawifolia Niitt. ex Torr. & Gray present in the Utah Lake flora but for which Ribes aureiim Pursh there is not any evidence that such is the

Scrophulariaceae case. Castilleja chromosa A. Nels. Amaranthus lividus L. Castilleja exilis A. Nels. Camelina sativa (L.) Crantz. Castilleja minor (A. Gray) A. Gray Carex apcrta Boott. Collinsia grandiflora Dougl. Cenchrus tribuloides L. Cordijlanthus canescens A. Gray Erigeron anniius (L.) Pers. Mimulus glabratus HBK Gnaphalium occidentalis Nutt. Mimidiis guttatus DC. Lepidium ramosissimum A. Nels. Penstemon humilis Nutt. ex A. Gray Mirabilis linearis (Pursh) Heimerl. Verbascum thapsus L. Sagittaria graminea Michx. Veronica americana Schwein Scirpus nebraskensis L. Veronica anagallis-aquatica L. Veronica hederaefolia L. Veronica peregrina L. Literature Cited SOLANACEAE Arnold, B. B. 1960. Life history notes on the walleye, Lijcium halimifolium Mill. Stizo stedion vitreum vitreum Mitchell, in a tur- Physalis longifolia Nutt. bid water, Utah Lake, Utah. Utah Fish and Game Solanum dulcamara L. Department. Federal Aid Project F-4-r-5 T. Sokinum nigrwn L. Job 107 Solanum triflorum Nutt. pp. Barnett, B. 1964. An ecological study of waterfowl hab- Sparganiaceae itat at Powell's Slough, Utah Lake. Unpublished Sparganium emersum Rehmann thesis, Brighani Young Univ. 45 pp. Beck, D. E. 1942. Life history notes on the California Sparganium eiirycarpum Engelm. gull, No. 1 Great Basin Nat. 3:91-108. Tamaricaceae Bessey, G. E. 1960. The aquatic plants of central Utah Tamarix ramosissima Ledeb. and their distribution. Unpublished thesis, Brig- ham Young Univ. 85 pp. Thypaceae Brown, D. 1958. Methods of surveying and measuring vegetation. Typha angustifolia L. Commonwealth Agricultural Burlaux Farnham Royal, Bucks, England. Typha latifolia L. 223 pp. 84 Great Basin Naturalist Memoirs No. 5

F. A., T. 1976. Chavez, and J. Warner. The Domin- Phillips, E. A. 19.59. Methods of vegetation studv. Holt, guez-Escalante journal. Brigham Young Univ. Rinehart and Winston, New York. 107 pp. Press, Provo, Utah. 203 pp. RuzicKA, M. 1958. Anwendung Mathematisch- Christensen, E. M. 1965. Ecological observations of Statistichae Methoden in der geobotanik (Syn- peach-leaf willow in central Proc. Utah. Utah thetische bearbeitung von aufnahmen). Biologia, Acad. Sci., Arts, Lett. 43:85-88. Bratisl. 13:647-661. Coombs, R. E. 1970. Aquatic and semi-aquatic plant Skougard, M. G. 1976. Vegetational response to three communities of Utah Lake. Unpublished dis- environmental gradients in a salt plava near sertation, Brigham Young Univ. 252 pp. Goshen, Utah County, Utah. Unpublished thesis, Cottam, W. p. 1926. An ecological study of the flora of Brigham Young Univ. 75 pp. Utah Lake, Utah. Unpublished dissertation, Univ. Sneath, p. H. A., and R. R. Sokal. Numerical tax- Chicago, Chicago, Illinois. 137 pp. onomy. W. H. Freeman and Company, San Fran- Croquist, a., et al. 1977. Intermountain flora. Vol. 6. cisco. 573 pp. Columbia Univ. Press, New York. 584 pp. Wakefield, H. 1933. A study of the plant ecology of Foster, R. H. 1968. Distribution of the major plant J. Salt Lake and Utah Valleys before the Mormon communities in Utah. Unpubli.shed dissertation, immigration. Unpublished thesis, Brigham Young Brigham Young Univ. 124 pp. A. H., L. Univ. 54 pp. Holmgren, and J. Reveal. 1966. Checklist of the vascular plants of the Intermountain Region. 1937. Transect study of Utah Lake shore line U.S. Forest Service research paper INT-.32. Inter- from 19.30 to 1936. Proc. Utah Acad. Sci., Arts, mountain Forest and Range Experiment Station, Lett. 14:.39-40. H., U.S. Forest Service, U.S. Department of Agricul- Warner, J. and K. T. Harper. 1972. Understory ture, Ogden, Utah. 160 pp. characteristics related to site quality for aspen in Lawler, R. E. 1960. Observations on the life history of Utah. Brigham Young Univ. Sci. Bull., Biol. Ser. channel catfish, Ictahinis puncfatiis Rafinesque, 16(2): 1-20. in Utah Lake. Unpublished thesis, Utah State Weic;ht, K. E. 1928. The distribution, taxonomy and Univ., Logan. 69 pp. ecology of the genus Salix of Utah County, Utah. LiECHTY, W. R. 1952. A preliminary study of the genus Unpublished thesis, Brigham Young Univ. 72 pp. Carex in Utah County, Utah. Unpublished thesis, Welsh, S. L., and G. Moore. 1973. Utah plants. Brigham Young Univ. 105 pp. Tracheophyta. Brigham Young Univ. Press, Pro- MacArthur, R. H. 1972. Geographical ecology-patterns vo. Utah. 474 pp. in the distribution of species. Harper and Row, White, D. A. 1963. Ecology of .summer aquatic in- New York. 269 pp. vertebrate populations in a marsh area of Utah Murphy, J. R. 1951. Ecology of passerine birds winter- Lake. Unpublished thesis, Brigham Young Univ. ing at Utah Lake. Unpublished thesis, Brigham 36 Young Univ. 63 pp. pp.