Proceedings of the Academy of Science

Volume 92 Number Article 12

1985

The Vegetation of the Hills Landform Along the River

Jean M. Novacek Iowa State University

Dean M. Roosa Iowa Conservation Commission

William P. Pusateri

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Recommended Citation Novacek, Jean M.; Roosa, Dean M.; and Pusateri, William P. (1985) "The Vegetation of the Loess Hills Landform Along the ," Proceedings of the Iowa Academy of Science, 92(5), 199-212. Available at: https://scholarworks.uni.edu/pias/vol92/iss5/12

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Proc. Iowa Acad. Sci. 92(5): 199-212, 1985

The Vegetation of the Loess Hills Landform Along the Missouri River

JEAN M. NOVACEK1, DEAN M. ROOSA2 , WILLIAM P. PUSATERI3

The vascular flor:i o~ the Loess Hills landform in south~astern South Dakota, western Iowa, and northwestern Missouri totals 703 species. A species checklist is presented based upon an extensive field study over a four-year period, plus herbarium and literature searches. The geology of the area 1s discussed briefly, in addition to vegetational, climatic and topographic gradients. The upland prairies of the Loess Hills are s1m1lar to the mixed-grass prairies .of the Great Plains more than 70 miles to the west. The mixed-grass prairie of the Loess Hills represent .a narrow penmsular range extens10n mto an. area which otherwise typically supports call-grass prairie. At least 17 xerophytic Great Plams species reach the easternmost edge of the!f ranges on the loess prairies. In Iowa 13 of these species are restricted to the loess bluffs. Three new taxa are added to the vascular flora oflowa: Asclepias stenophylla Gray, Gaillardia pulchella Foug. and Solidago mollis Bartl. INDEX DESCRIPTORS: Iowa vascular flora, Loess Hills, prairie, Great Plains flora.

GEOLOGY Shimek (1930) maintained that the vertical distribution of fossils gives proof of slow edaphic changes in the loess. In the lower section of The Loess Hills landform, a unique aeolian deposit, is located in deep vertical cuts, he found many fossils of the "reduced forest rype" extreme southeastern South Dakota, along the western border of Iowa whose numbers gradually diminish upward until only a small number and in extreme northwestern Missouri (Fig. 1). The most prominent of the "border-type" snails remain, which, in turn, disappear com­ ridges extend three to ten miles eastward from the Missouri River pletely. This distributional pattern suggested to Shimek that when valley. the valley walls were low, they were colonized by woodlands which Loess is a elastic deposit, composed of quartz silt particles 20-50 were inhabited by the "forest-type" snail. As the loess deposits microns in diameter, which occurs as wind-laid sheets (Smalley, gradually increased, more xeric conditions followed, as evidenced by 1972). The loess deposits which make up the Loess Hills are up to 150 the "border-type" snail which inhabited woodland edges and thickets, feet thick, and, in some areas, thicknesses of over 200 feet have been until the woodlands were replaced by prairie which was not inhabited recorded (Prior, 1976). Because of this unusual thickness, and because by snails. This would not require an overall change in climate, but loess forms the actual topography of the region rather than the more only a gradual development of local modifications. usual mantling layer, the Loess Hills along the Missouri River are considered geologically unique. Loess soil is quite porous and has the CLIMATE AND TOPOGRAPHY tendency to cohere when dry; due to this cohesive property, near­ vertical slopes of loess are naturally maintained (Prior, 1976). The Loess Hills are located within the Missouri River drainage Loess was deposited with fluctuations of intensity over a period of system. The landform is oriented in a north-south direction and lies thousands of years (Handy and Davidson, 1956). There were two between 43° and 40° North latitude. The major tributary streams and major periods of ioess deposition, during the Illinoian and during the rivers follow a southwesterly course through the hills to the Missouri Wisconsinan glacial periods, with the greatest accumulation occur­ River. The average elevations of the Missouri River floodplain and the ring during the latter period (Prior, 1976). Handy (1976) states that prominent loess bluffs decrease from north to south (Table 1). the average accumulation of Wisconsin loess was 0.6 centimeters per The annual precipitation received at the northern extremity of the year over a 15,000 year interval. Loess Hills in Union County, South Dakota, is 25"inches (63.5 cm.); According to Prior ( 1976), during spring and summer, glacial in Monona County, Iowa, 30 inches (76.2 cm.) per year is received; meltwaters formed large rivers carrying heavy sediment loads, one of and the yearly amount received in Holt County, Missouri, at the which covered most of the Missouri River floodplain. After winter southern extremity of the Loess Hills, is 38 inches (96. 5 cm.) (Lawson and colder temperatures arrived, the meltwaters abated and exposed et al., 1977). Climatological data (U.S. Department of Commerce, vast quantities of fine silt and sand which dried out in the arid winter 1973-1983) for Iowa substantiate that the northern Loess Hills receive climate. The prevailing westerly winds picked up the dry, loose significantly less precipitation annually than the southern Loess Hills material forming huge clouds of windblown silt, which, after encoun­ (Table 2). Additionally, the average annual temperature is higher in tering the rough valley walls, was deposited in massive quantities. the southern part than in the northern part, and the growing season The Loess Hills have been modified by subsequent erosion, causing length in the southern Loess Hills is longer than in the northern hills even sharper distinctions in the topography. Shimek ( 1931) pointed out that during the period of deposition, an Table 1. Average elevations of the prominent bluffs of the anchorage for the loess was necessary to prevent it from either blowing Loess Hills landform, the Missouri River floodplain and the or washing away. In his opinion, an abundant vegetation must have difference between them in feet (m.). provided this anchorage. Shimek's ( 1930) most compelling evidence of an extensive community is the presence of fossil forest­ COUNTY BLUFFS FLOODPLAIN DIFFERENCE dwelling snails, which he further contended indicate the type of Union 1470 (448) 1120 (341) 350 (107) vegetation present at the time of loess deposition. He pointed out that Plymouth 1450 (442) 1110 (338) 340 (104) the fossil shells appear in colonies similar to those of present-day Woodbury 1400 (427) 1085 (331) 315 ( 96) woodland snails and that the intervening loess between woodland Monona 1350 (411) 1045 (318) 305 ( 93) groves lacks fossils, indicating that the loess was deposited upon open Harrison 1300 (396) 1025 (312) 275 ( 84) prairie. Pottawattamie 1250 (381) 1000 (305) 250 ( 76) Mills 1225 (373) 960 (293) 265 ( 80) Fremont 1200 (366) 925 (282) 295 ( 84) 'Botany Department, Iowa State University, Ames, Iowa 50011 Atchison 1150 (350) 900 (274) 250 ( 76) 2State Preserves Advisory Board, Iowa Conservation Commission, Des Moines, Iowa Holt 1000 (305) 880 (268) 220 ( 37) 50319 377 West Post Rd., N.W., Cedar Rapids, Iowa 52405

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200 PROC. IOWA ACAD. SCI. 92(1985)

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Fig. I. The Loess Hills landform (shaded area) parallels the Missouri River in Iowa, South Dakota and Missouri. Numbers refer to counties: South Dakota: Union County (l); Iowa: Plymouth County (2), Woodbury County (3), Monona County (4), Harrison County (5), Pottawattamie County(6), Mills County (7), Fremont County (8); Missouri: Atchison County (9), Holt County (10). Scale is 1 inch to 65 miles (25 mm. to 105 km.).

(Table 2). There appears to be a two to three week difference in length. Latitude is probably the more important factor of the two. It is phenology between the northern and southern sections, with bloom­ likely that the similarity to the northern Great Plains exhibited by the ing time earlier in the southern part (Aikman, 1926; pers. observ.). northern Loess Hills is due, in part, to the increase in latitude. A Other gradational differences between north and south exist. The summary of these climatic and physical gradients is depicted in Table northern part has a greater number of sunny days, stronger and more 3. frequent winds, and a higher average daily evaporation rate than the southern portion (Aikman, 1926; Visher, 1954). During a study of a VEGETATION series of woodland transects bn the side of the Missouri River, Aikman (1926) found that humidity is consistently greater in The vegetational community of the greatest interest in the Loess the woodlands of the southern section throughout the growing season Hills is the prairie, although other community types, such as than in the woodlands of the northern part. Similarly, he found that woodland, aquatic and ruderal, are present. The aquatic community soil moisture decreases from south to north. He also pointed out that consists mainly of streamside vegetation in addition to marshy areas precipitation during the growing season is more evenly distributed in around small ponds. The ruderal element, well-represented by both the southern part, which is more conducive to tree growth. native and non-native species, occurs on disturbed sites. The wood­ While these are not all the gradients which can be measured, it is lands are basically upland; however, the floodplain woodland may be easily discerned that the environment in the northern Loess Hills is found in well-developed stream valleys. In the southern most Loess much different from that in the southern hills. The most significant Hills of Fremont County, Iowa, and Atchison and Holt counties, factors accounting for these differences seem to be latitude and altitude Missouri, the woodlands are much more diverse than in the northern which affect precipitation amounts, temperature and growing season counties. https://scholarworks.uni.edu/pias/vol92/iss5/12 2 Novacek et al.: The Vegetation of the Loess Hills Landform Along the Missouri Riv

VEGETATION OF LOESS HILLS 201

Table 2. Mean climatological data for Iowa for 1973 through 1983 arranged from north to south (Us De f C 1973-1983). ' · · pt. o ommerce, WEATHER TOTAL ANNUAL AVERAGE ANNUAL STATION PRECIPITATION LENGTH OF TEMPERATURE GROWING (coun!l'.) inches (cm.) OF (OC) SEASON* Hawarden 25.96 (65.94) 47.36 (Sioux) ( 8.53t 161 Akron 24.12 (61.26) NA (Plymouth) NA Sioux City 24.18 (61.42) 48.23 ( 9.02) (Woodbuty) 179 Castana 28.56 (72.54) 48.71 ( 9.28)b (Monona) 172 Onawa 28.70 (72.90) 50.03 (10.02)b (Monona) 177 Logan 31.39 (79. 73) 50.16 (10.09) 183 (Harrison) Glenwood 31.70 (80.52) 51.25 (10.69) 174 (Mills) Sidney 34.98 (88.85) 52.16 (11.20) 207 (Fremont)

*Number of days between last spring minimum ~28°F ( - 2°C) and first fall minimum ~28°F. •Mean based upon 9 years due to missing data. bMean based upon 10 years due to missing data.

A Great Plains vegetation map developed by Kaul (in press) places factor in the maintenance of loess prairies. Iowa in the tall-grass prairie region with deciduous woodlands located The typical upland loess prairie is dominated by two warm-season mainly along river systems. The tall-grass prairie also extends into the bunch-grasses: Andropogon scoparius Michx., which comprises up to 90 eastern one-third of Nebraska, the eastern one-fourth of South percent of the total vegetation, and Bouteloua curtipendula (Michx.) Dakota, and the northwest corner of Missouri. Mixed-grass prairie Torr. , which is found as the dominant grass in areas damaged by begins at about the lOOth meridian and extends westward until drought or overgrazing (Weaver, 1968). This short-grass component giving way to the short-grass prairie at or beyond the 102nd meridian. is represented by Bouteloua hirsuta Lag. and B. gracilis (H.B.K.) This map makes an additional distinction by dividing the mixed-grass Griffiths. There have also been reports of Buchloe dactyloides (Nutt.) prairie into a northern part, which includes Nebraska, South Dakota, Engelm. in Iowa (Pammel, 1901; Thorne, 1956) and in Missouri North Dakota and Saskatchewan, and a southern part, which includes (Goodnight, 1964; Iffrig, 1980). On the near-vertical faces of loess, , Oklahoma, and Texas. This map, however, does not recog­ Muhlenbergia cuspidata (Torr.) Rydb., and Calamovilfa longifolia nize the occurrence of mixed-grass prairie found on the Loess Hills (Hook.) Scribn. are common. which is connected to the main body of mixed-grass prairie along the A depauperate tall-grass prairie vegetation occurs on the lower Missouri River bluffs in South Dakota and Nebraska. The Loess Hills slopes and roadsides, where the soil holds more moisture. The most represent the largest relatively contiguous area of mixed-grass prairie common grasses here are Andropogon gerardi Vitman, Sorghastrum in Iowa, and the only example in Missouri (Iffrig, 1980). avenaceum (Michx.) Nash, Panicum virgatum L. and Elymus canadensis L. The prairies of the Loess Hills are characterized by a xerophytic Within the Loess Hills prairie community, only a few of the more flora. In general, the south and west slopes support the prairie com­ hardy tall-grass prairie forbs are present and, when present, are most munity, while the north and east slopes support a woodland commun­ frequently found on lower slopes and woodland edges. Thus, there is a ity. Shimek ( 1900; 1909) pointed to two dominant factors which help create the xeric conditions on the south and west slopes, an intense Table 3. Summary of the change in climatic and physical afternoon sun and hot, dcy westerly winds. These factors cause an factors along a gradient from south (40"N latitude) to north increase in transpiration and only drought-tolerant prairie have (43"N latitude). adapted to these conditions. Costello (1931) concluded that, due to prevailing westerly winds, the rate of evaporation on the southwestern FACTOR CHANGE ALONG GRADIENT slopes is almost twice as great on west-facing slopes as on protected Elevation Increases east-facing slopes. Wind• Increases In addition to the western slopes, the prairie has colonized north Sunny Days Increases and east slopes in some areas of the Loess Hills. This is a common Evaporation Increases occurrence in the northern Loess Hills in Monona, Woodbury and Precipitation Decreases Plymouth counties. According to Shimek ( 1909), this phenomenon is Temperature Decreases also a result of the wind. If the westerly wind is intercepted and Growing Season Decreases deflected by a series of ridges, this causes a division in the air current, Humidity Decreases part of which sweeps back up the leeward slope. In this situation, the Soil Moisture Decreases leeward slope is nearly as exposed to the desiccating winds as the windward slope. It must be noted that fire has also been an important "velocity and frequency

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PROC. IOWA ACAD. SCI. 92(1985) 202

( 1964), a report by Iffrig ( 1980), maps of. th.e Atlas of the noticeable lack of mesic species so common on typical tall-grass distri~ution p fora of the Great p fains by the Great Plams Flora Assooat1on (~arkley, prairies elsewhere. 1977), and personal observations. The South ~akota Loess .H1~ls fl?ra METHODS is based upon a flora of Union County by Eslick 09.62), d1stnbut1on maps (Barkley, 1977) and frequent field trips t~en smce 1?79. Other An intensive floristic survey was conducted during 1981 and 1982 plant lists and distribution reports for Iowa ~nclud~: A1km~n and in Plymouth, Woodbury and Monona coun~ies by Novacek .Bates Gilly ( 1948), Bennett ( 1948), Coffey ( 1966), ~lt~patnck and Fitzpat­ (1983); in 1983 and 1984 additional .collect10~s were made m the rick (1898), Gabel (1984), Gilley and 0 Bnan (194~), Kwang remaining Iowa loess counties. The mam collection has bee?- donated (1951), Lammers and Van Der Valk (1977; 1979), NIColson and to the herbarium at the University of Northern Iowa. During 1981, Russell(l955), Pammel(1895), Peck(l976; 1980; 1983), Pecketal. 1982 and 1983, the State Preserves Advisory Board sponsored three (1978), and Pohl (1966). . . natural history forays in the Iowa Loess Hills to obtain data on the flora One of the difficulties encountered while studymg .old records or and fauna of natural areas. The foray data supplemented the original literature reports of some species was that the preose location. of floristic study. In addition, personal observations have b~n record~d collection is not mentioned. This was most troublesome whe.n trymg during numerous field trips into the hills since 1978. Prev10~ stu~1es to distinguish whether the plant was collected. in the Loess Hills .or on of the Loess Hills and surrounding area were also consulted, mcludmg the Missouri River floodplain. If the location was not speofied, those by Carter (1960), Fay (1953), and Morrill (1953). The Missouri aquatic species were presumed to be floodi;>lain species and were n~t loess flora is based upon notes compiled by Bush (1895) on Atchison recorded as being present, nor were mesIC or low-meadow prame County loess prairie, a flora of Holt County completed by Goodmght species. However, if literature or herbarium searches revealed that either aquatic or mesic prairie species were actually collected or observed specifically within the Loess Hills landform, they were Table 4. Distribution of tree and shrub species in the Loess included in the checklist. In addition, if such species have been Hills indicates a gradient from south to north. Presence is collected within the landform by the authors, they were included. indicated by an "X". County name symbols: Union (1), Plymouth (2), Woodbury (3), Monona (4), Harrison (5), NOMENCLATURE Pottawatamie (6), Mills (7), Fremont (8), Atchison (9), Holt (10). Identifications were aided by the following references: Gillis (1971), Gilly (1946), Gleason (1952), Gleason and Cronquist (1963), 1 2 3 4 5 6 7 8 9 10 SPECIES Peck (1976), Pohl (1966), Shinners (1941)'. Van Bruggen (1976), an? x x x x x x x x x x Ce/tis occidentalis Woodson (1954a; 1954b). Nomenclature 1s based upon the determi­ x x x x x x x x x x Cornus drummondii nations of the Great Plains Flora Association (Barkley, 1977). Two x x x x x x x x x x Crategus mollis nomenclatural compromises were necessary. Based upon Shinners x x x x x x x x x x Fraxinus pennsylvanica (1941), all reports of Aster sagittifolius Wedem. ex Willd., A. Gymnocladus dioica x x x x x x x x x x drummondii Lindi. and A. cordifolius L. in the Loess Hills are included x x x x x x x x x x Jug/ans nigra in A. sagittifolius Wedem. var. drummondii (Lindi.) Shinners. Based x x x x x x x x x x Prunus americana upon Gilly (1946), all reports of Carex convoluta Mackenz. are x x x x x x x x x x Prunus virginiana included in C. rosea Schkuhr ex Willd. var. pusilla Peck ex Howe. x x x x x x x x x x Rhus glabra x x x x x x x x x x Ribes missouriense EXCLUDED SPECIES x x x x Quercus macrocarpa x x x x x x Herbarium searches at the University of Iowa and Iowa State x x x x Tilia americana x x x x x x University were undertaken for the confirmation of rare taxa or of x x x U Imus americana x x x x x x x specimens of dubious identification. Based upon these findings, the x x x x x Ulmus rubra x x x x x fullowing species are excluded from the Loess Hills flora. Agropyron x x x x x x x x Carya cordiformis x cristatum (L.) Gaertn. (including A. desertorum (Fisch.) Schult.), x x ·X x x x x x x Ostrya virginiana reported for Monona and Woodbury counties by various workers, is x x x x x x x x Quercus borealis actually A. pectiniforme Roem and Schult. Apocynum sibiricum ]acq. x x x x x x x x Staphylea trifolia reported for Plymouth County (Carter, 1960) is actually a specimen of x x x x x x x Cory/us americana Euphorbia marginata Pursh. Cornus rugosa Lam. reported for Monona x x x x x x x Morus rubra County (Carter, 1960) is actually C. drummondii Meyer, which appears x x x x x x Carya ovata to be the only species of Cornus in the Loess Hills (pers. observ.). x x x x x x Cercis canadensis Elymus riparius Wieg. reported for Woodbury County (Morrill, 1953) x x x x x x Fraxinus americana is excluded since no voucher can be located. Helianthus petiolaris Nutt. x x x x x x Quercus velutina reported from a Woodbury County loess prairie (Morrill, 1953) is x x x x x Amelanchier arborea actually H. annuus L. (H. petiolaris does grow in sandy areas of the x x x x x Platanus occidentalis Missouri River floodplain.). Petalostemon candidum (Willd.) Michx., x x Carya tomentosa reported as growing on loess bluff prairies by various workers, is actually its western counterpart, P. occidentale (Gray) Fern., which has x x Quercus imbricaria x x x x Quercus muhlenbergii sometimes been lumped with the former species (See Isely and Welsh, x x x Asimina triloba 1960.). Petalostemon multiflorum Nutt., reported in Woodbury and Plymouth counties (Carter, 1960), was based upon atypical specimens x x Phytolacca americana of P. occidentale, and it is doubtful that this species occurs in Iowa x x Quercus prinoides (Wemple, 1970). Tephrosia virginiana (L.) Pers. reported in Woodbury x Quercus stellata County (Carter, 1960) is actually a specimen of Amorpha canercens x Acer nigrum Pursh. x Quercus alba https://scholarworks.uni.edu/pias/vol92/iss5/12 4 Novacek et al.: The Vegetation of the Loess Hills Landform Along the Missouri Riv

VEGETATION OF LOESS HILLS 203

RESULTS of prairie fires has also allowed the woodland community to expand. . ~e vascular flora of the Loess Hills landform consists of 703 The woodlands along the Missouri River bluffs represent one of the mdi~~dual ~pecies from 97 families and 358 genera (Table 9). The five last .ou~ts for the eastern deciduous woodlands in the prairie region, furn.dies with the greatest number of species are: Asteraceae (96), havmg rmgrated from the east through Missouri into western Iowa Poaceae (95), Fa~eae (48), Cyperaceae (31), and Brassicaceae (30). and eastern Neb~ka (Aikman, 1926; Aikman and Gilly, 1948; Three new species have .been added t~ Iowa's state flora: Asclepias Gleason, 1922; Shimek, 1900; Transeau, 1905). The dominant trees stenophylla Gray (Asclepiadaceae), Gat!lardia pukhella Foug. (As­ throughout the Loess Hills woodlands are species of Quercus L. and teraceae), and Solidago mollis Bartl. (Asteraceae). Carya Nutt., oak and hickory. A gradient in the tree and shrub The species list (Table 10) is annotated with an asterisk (*) before composition is apparent. Not only is the areal extent of the woodland t~e bi~omial of species not native to the region. Following the species greater in the southern Loess Hills but also the number of bmom1al are symbols indicating the general habitat in which the individual species present. Table 4 graphicaily depicts this woodland plant is commonly found: P (prairie), W (woodland), A (wetland), gradient. Aikman ( 1926) noted a similar gradient in his study of the E (wooc_l~d ~ge), and R (ruderal). The county in which each species eastern Nebraska woodlands. In light of the climatic and physical gradients (Table 3), this is not unexpected. occurs 1s md1cated by numbers 1 through 10 corresponding to the numbering system presented in Figure 1. Prairie The upland loess prairies exhibit a narrow peninsular distribution DISCUSSION of mixed-grass prairie, linked to the western plains along the rugged bluffs of the Missouri River in southeastern South Dakota and Woodlands northeastern Nebraska, which extends into an area dominated by tall­ The southern Loess Hills have historically been more forested than grass prairie prior to settlement. Sixty typical upland loess prairie the northern hills (Shimek, 1900), and the same is true today. species are listed in Table 5. This composition is noteworthy because Evidence indicates that wooded areas have been increasing since the of the similarities with the mixed-grass prairies approximatley 70 or time of settlement, much to the detriment of the prairies (McComb more miles to the west. Personal observations indicate that a gradient and Loomis, 1944; Heineman, 1982). Assessing the woody plant in the number of prairie species exists from north-to-south in the Loess invasion in southern Monona county, Heineman (1982) concluded, Hills, which is opposite of the south-to-north gradient for the after examining aerial photos taken from 1940 to 1980, that the woodland community. average overall canopy coverage of woody species has increased Of the 40 Great Plains species reaching western Iowa (Table 6), 32 approximately 40 percent during this period. He found that canopy (78%) occur in Plymouth County, more than any other Loess Hills coverage has increased by 15 percent on bluff tops, 26 percent on county. Many of these Great Plains species are restricted to the loess upper slopes, and 55 percent on lower slopes. He suggested that the bluffs (Table 6), and some reach the easternmost edge of their ranges suppression of prairie fires has been a major factor in allowing the here. Novacek (this issue) discussed the phytogeographical aspects of expansion of woody species. the Iowa loess prairies and postulated that the Missouri River bluff The Loess Hills lie in a zone of transition which is climatically system is a possible migrational route for many of these Great Plains capable of supporting either woodlands or prairie. In years of abun­ species. The Plymouth County loess bluffs represent the first suitable dant rainfall, tree growth is favored and woodlands may expand; habitat for these species along the proposed migrational route, however, in extended periods of drought, trees are injured or killed perhaps offering a partial explanation of the large number of Great and prairie species are favored (Albertson and Weaver, 1945). This Plains species there. Another possible explanation. is the climatic, leads to the conclusion that long-term weather patterns may affect the topographic, and geographic similarity to the northern Great Plains. distribution of species. It is possible that the present weather cycle is Other distributional patterns of interest emerge. Some xeric Great predominantly wet which would favor woody plants. The suppression Plains species, such as Dalea enneandra Nutt. and glauca Nutt.,

Table 5. Typical upland loess prairie species which are similar to the mixed-grass prairie species occurring 70 or more miles to the west. FORBS GRAMINOIDS Agalinis aspera Delphinium virescens Penstemon grandiflorus Andropogon scoparius Ambro.ria psilostachya Echinacea angustifolia Petalostemon occidentale Aristida longiseta Amorpha canescens Erigeron strigosus Petalostemon prupureum Bouteloua curtipendula Anemone patens* Gaura coccinea Psoralea argophylla Bouteloua gracilis Asckpias viridiflora Haplopappus spinulosus Psoralea esculenta Bouteloua hirsuta Aster ericoides Helianthus rigidus Ratibida columnifera Calamrwilfa longifolia Aster oblongifolius Kuhnia eupatorioides Rosa arkansana Carex heliophila Aster sericeus Liatris punctata Senecio plattensis Dicanthelium oligosanthef' Astragalus crassicarpus Linum rigidum Sisyrinchium campestre Dicanthelium oligosanthel* Astragalus lotiflorus Lithospermum incisum Sphaeralcea coccinea* Koeleria pyramidata Astragalus missouriensis* Lygodesmia juncea Solidago missouriensis Muhlenbergia cuspidata Calylophus serrulatus Mentzelia decapetala* Solidago nemoralis Sporobolus asper Castilleja sessiliflora Microseris cuspidata Solidago rigida Sporobolus cryptandrus Cirsium flodmani Oxytropis lambertii Vicia americana var. minor Stipa spartea Datta enneandra Penstemon alibidus* Yucca glauca Stipa viridula

a var. scribnerianum b var. wilcoxianum * found in northern loess hills only

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Table 8. Plants of the southern Great Plains which reach Table 6. Great Plains species which reach western low~ their northern limits in the Loess Hills and Missouri River along the proposed Missouri Ri:ve~ system and Loess Hills migrational route (Novacek, this issue). area. Andropogon ha/Iii* Linum rigidum 41° N LATITUDE 42° N LATITUDE Aristida longiseta Lygodesmia juncea (Mills County) (Monona County) Astragalus lotif/orus Lygodesmia rostrata Callirhoe involucrata Euphorbia corollata Astragalus missouriensis* Mentzelia decapetala* foeniculaceurff' Gaura longif/ora Bouteloua hirsuta Oenothera strigosa Hedyotis nigricans Buchloe dactyloides Oxytropis lambertii Penstemon cobaea Plantago virginica Vernonia baldwini Calamovi/fa longifolia Penstemon albidus Schrankia nuttallii Callirhoe involucrata Penstemon cobaea* Cirsium flodmani Penstemon gracilis Dalea enneandra* Penstemon grandiflorus a var. daucifolium Echinacea angustifolia Petalostemon occidentale* Euphorbia hexagona Ratibida columnifera the loess prairie community. At this point, it is found quite common­ Euphorbia marginata Schedonnardus paniculatus ly growing on sandy areas of the Missouri River floodplain, where it Euphorbia stictospora* Shepherdia argentea continues northward until reaching its northern terminus in the Great Gaura coccinea Solidago mollis* Plains in North Dakota. Gaura parviflora* Sphaeralcea coccinea* Two varieties of Lomatium foeniculaceum (Nutt.) Coult. and Rose Grintklia squarrosa Stipa comata illustrate another north-south Great Plains distributional separation. Haplopappus spinulosus Stipa viridula L. foeniculaceum var. foeniculaceum is a northern Great Plains species Lomatium foeniculaceum*" Vicia americana var. minor which is fairly common westward on the high plains. This variety Lomatium orientate Yua-a glauca* occurs in Iowa only in cyon and Sioux counties just a few miles north of Plymouth County (Peck et al., 1984). A southern Great Plains * Plants restricted to the loess bluff prairies in Iowa. element, L. foeniculaceum var. daucifolium (T. & G.) Cronq. occurs only a var. daucifolium in Fremont County. This distributional pattern is noteworthy and may provide some clues for attempts at reconstructing the vegetation­ are found in all ten Loess Hills counties. Although it has not been al history and migrational routes in the Loess Hills area. quantified, it has been noted that proceeding southward the frequen­ cy of these plants becomes less (pers. observ.). Another more striking SUMMARY pattern is that some of the Great Plains species reach a southern terminus in the Loess Hills at approximately the 42nd parallel which The upland loess prairies, which are similar to the Great Plains intersects Monona County (Table 7). mixed-grass prairies, are the most intriguing vegetational community Although the predominant migrational direction in the Loess Hills of the Loess Hills. Although this prairie type is common in the for many Great Plains prairie species appears to be southward western two-thirds of South Dakota, it is quite unusual in Iowa and (Novacek, this issue), nevertheless, there are a few members of the Missouri. Vegetational gradients are noted both in the woodland and southern Great Plains flora which appear to be migrating northward prairie communities, in addition to a number of climatic and physical on the loess bluffs. Some of these southern species reach a northern gradients. Both the northern and southern Loess Hills demonstrate terminus in the Loess Hills at approximately the 42nd parallel, and affinities with the northern Great Plains; however, a few southern some near the 4 lst parallel in Mills County (Table 8). This pattern is Great Plains elements are present in the southern Loess Hills. It also noted among the woodland species, with many terminating their appears that the 42nd parallel coincides with one or more ecologically range near the 42nd parallel (Table 4). It appears, then, that the 42nd limiting factors for many species which are at the edge of their ranges. parallel coincides with one or more ecologically limiting factors for many plants which are at the edge of their ranges, regardless of ACKNOWLEDGEMENTS whether they are northern or southern elements. We thank the curators of the herbaria of Iowa State University and Some plants cope with the environmental stresses which increasing the University of Iowa, Drs. R. W. Pohl and T. Melchert, for latitude seems to bring in an unusual manner. For example, rather allowing access to their collections. We thank Drs. L. J. Eilers and than terminating its range near the 42nd parallel as some southern D. R. Farrar for their critical comments and suggestions. A special Great Plains species do, Desmanthus illionensis (Michx.) MacM. moves thanks goes to Dr. T. Van Bruggen for his valuable guidance and to another habitat. This species occurs on uplap.d loess prairie in Holt assistance during the course of this study. through southern Pottawattamie counties, but then disappears from

Table 7. Plants of the Great Plains which reach their southern limits in the Loess Hills at approximately 42"N latitude (Monona County).

Anemone patens Penstemon gracilis Table 9. Summary of the Loess Hills flora. Astragalus missouriensis Schedonnardus paniculatus Dicanthelium oligosanthe.f" Shepherdia argentea PTERI- Euphorbia stictospora Sphaeralcea coccinea DOPHYTES CONIFERS DICOTS MONOCOTS TOTAL Mentzelia decapetala Solidago mollis #Families 3 1 80 13 97 Penstemon albidus Stipa viridula #Genera 7 1 277 73 358 #Species 12 1 530 160 703 a var. wilcoxianum https://scholarworks.uni.edu/pias/vol92/iss5/12 6 Novacek et al.: The Vegetation of the Loess Hills Landform Along the Missouri Riv

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Table IO. Species checklist for the Loess Hills landform. Aralia racemosa L. W 2 4 8 9 Taxon Habitat County occurrence Panax quinquefolius L. w 8 l 2 3 4 5 6 7 8 9 10 ARISTOLOCHIACEAE (Birthwort Family) Asarum canadense L. W 6 IO EQUISETOPHYTA . ASCLEPIADACEAE (Milkweed Family) EQUISETACEAE (Horsetail Family) Asclep1as engelmanniana Woods. p 4 6 Equisetum arveme L. AW 123456789IO (A. auriculata (Engelm.) Hotz.) E X ferrissii Clute PR 12345678 IO A. lanuginosa Nutt. p 3 E. hyemale L. AW 23456789 (A. nuttalliniana Torrey) E. laevigatum A. Br. p 12345678 A. stenophylla Gray p 2 POLYPODIOPHYTA A. syriaca L. R 2 3 4 5 6 7 8 9 IO . OPHIOGLOSSACEAE (Adder's-tongue Family) A. tuberosa L. p 23456 7 8 9 IO Botrych1um sp. nov. P E 2 3 4 6 8 A . verticillata L. RP 23456 7 8 9 IO B. virginiana Sw. P W 1 2 3 4 6 7 8 9 IO A. viridiflora Raf. p 123456 7 8 9 IO POLYPODIACEAE (Fern Family) ASTERACEAE (Sunflower Family) 4 6 8 9 IO Adiantum pedatum L. w *Achillea millefolium L. PR 1 2 3 4 6 7 9 IO Athyrium fillix-femina (L.) Roth w 4 6 spp. lanulosa (Nutt.) Piper 6 8 Camptosorus rhizophyllus w 5 Ambrosia artemisiifolia L. R 2 3 4 5 6 7 8 9 IO (1.) Link A. psilostachya DC. p 2 3 4 5 6 7 8 9 IO 6 Cystopteris fragilis (L.) Bernh. w 3 4 A. trifida L. AR 2 3 4 5 6 7 8 9 IO var. mackayi Laws. Antennaria neglecta Greene E p 2 3 4 5 6 7 8 9 2 3 4 6 8 9 IO C. protrusa (Weath.) Blasd. w *Anthemis cotula DC. R 5 6 8 IO 4 Matteucia struthiopteris w *Arctium minus Schkuhr E R 3 4 6 8 IO (L.) Todaro *Artemisia biennis Willd. R 6 7 8 PINOPHYTA A. dracunculus L. E 2 3 4 IO CUPRESSACEAE (Cypress Family) (A. glauca Pall.) 7 8 9 IO Juniperus virginiana L. P W 1 2 3 4 5 6 A. ludoviciana Nutt. p 2 3 4 6789IO MAGNOLIOPHYTA: MAGNOLIOPSIDA (Dicots) var. ludoviciana ACANTHACEAE (Acanthus Family) Aster azureus Lindi. E 6 7 8 9 8 IO Ruellia humilis Nutt. E P A. ericoides L. p 2 3 4 5 6 7 8 9 IO ACERACEAE (Maple Family) A. laevis L. PW 2 3 4 5 6 7 8 9 IO Acer negundo L. EW 123456789IO A. oblongifolius Nutt. P 2 3 4 5 6 7 8 9 IO IO Acer nigrum Michx. f. w A. ontarionis Wieg. A 3 5 6 8 A. saccharinum L. w 123456789IO A. sagittifolius Wedemeyer P 5 6 7 8 9 IO AMARANTHACEAE (Pigweed Family) var. drummondii (Lindi.) Shinners Amaranthus a/bus L. R 1 9 A. sericeus Vent. P 2 3 4 6 7 8 9 IO IO A. grateizans L. R 2 3 4 6 7 8 A. simplex Willd. A 2 3 4 5 6 7 8 9 IO 9 IO A. hybridus L. R Bidens cernua L. A 2 3 4 5 6 7 8 9 IO *A. retroflexus L. R 2 3 4 6789IO B. frondosa L. A 2 3 4 5 6 7 8 9 10 8 IO A. rudis Sauer A 3 4 Cacalia atriplicifolia L. E 6 8 9 IO (A. tamariscinus (Nutt.) Wood.) C. tuberosa Nutt. P 3 4 5 6 7 8 9 IO ANACARDIACEAE (Cashew Family) *Carduus acanthoides L. R 2 3 5 6 7 8 IO RhusglabraL. EP 1 2 3 4 5 6 7 8 9IO *C. nutans L. R 2 3 4 5 6 Toxicodendron radicans E P W 2 3 4 5 6 7 8 9 IO *Centaurea cyanus L. R IO (1.) 0. Ktze. ssp. negundo (Greene) Gillis *Cichorium intybus L. R 3 4 8 3 4 5 6 7 8 9 10 T. rydbergii (Small ex E P 2 3 4 Cirsium altissimum (L.) Spreng. E 2 Rydberg) Greene C. discolor (M uh!.) Spreng. E 3 5 8 ANNONACEAE (Custard-apple Family) C. flodmani (Rydb.) Arthur p 2 3 4 6 8 9 IO p 8 9 IO Asimina triloba (L.) Dunal W C. undulatum (Nutt.) Spreng. 6789IO ( Family) Conyza canadensis (L.) Cronq. R 2 3 4 6 *Anethum gravetJlens L. R 1 (Erigeron canadensis L.) 7 2 4 6 8 *Carum carvi L. R C. ramosissima Cronq. R 6 8 IO p 2 6 7 8 9 10 *Conium maculatum L. A R 3 Coreopsis palmata Nutt. 6 8 IO Cryptotaenia canadensis (1.) DC. W 2 3 4 C. tinctoria Nutt. AR 3 4 6 8 IO R 2 4 6 7 8 *Daucus carota L. R Dyssodia papposa Heracleum sphondylium L. A E 3 5 6 (Vent.) Hitchc. p 3 4 5 6 7 8 ssp. montanum (Schleicher) Brig. Echinacea angustifolia DC. 2 (H. lanatum Michx.) var. angustifolia 8 Lomatium foeniculaceum p (incl. E. pallida Nutt. of reports) 4 6 8 9 IO (Nutt.) Coult. & Rose Erigeron annuus (1.) Pers. E 2 3 4 5 6 8 10 var. daucifolium (T. & G.) Cronq. E. philadelphicus L. W 3 4 5 6 7 8 10 Osmorhiza claytonii W 2 3 4 6 8 9 E. strigosus Muhl. P 2 3 4 5 6 7 8 9 10 (Michx.) Clarke Eupatorium altissimum L. E W 8 IO 3 4 6 7 8 IO O. longistylis (Torr.) DC. w 3 4 5 E. purpureum L. A E 3 4 5 6 8 IO 2 3 4 6 7 8 9 10 *P astinaca saliva L. R 2 E. rugosum Houtt. w 8 IO p 3 10 Sanicula canadensis L. w 2 3 4 5 Gaillardia pulchella Foug. 8 9 IO 7 S. gregaria Bickn. W 3 4 5 *Galinsoga ciliata (Raf.) Blake R 6 8 S. marilandica L. W Gnaphalium obtusifolium L. R 8 2 4 6 7 8 Tuenidia integerrima (L.) Drude W Grindelia squarrosa p R APOCYNACEAE (Dogbane Family) (Pursh) Dun. var. squarrosa IO 6 7 8 Apocynum cannabinum L. A R 1 2 3 4 5 6 7 8 Haplopappus spinulosus p 2 3 4 ARALIACEAE (Ginseng Family) (Pursh) DC.

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PROC. IOWA ACAD. SCI. 92(1985) 206

Ostrya vir.iriniana (Mill.) K. Koch E W 2 3 4 5 6 7 8 9 10 2 3 4 5 6 7 8 9 10 Helianthus annuus L. R BIGNONIACEAE (Bignonia Family) H. hirsutus Raf. W 6 8 8 10 3 4 5 6 7 8 9 *Campsis radicans (L.) Seem. E 1 H. maximiliani Schrad. P 2 10 8 10 2 3 4 5 6 7 8 9 10 *Catalpa speciosa Warder E 2 H. rigidus (Cass.) Desf. P BORAGINACEAE (Borage Family) (H. laetiflorus Pers.) *Cynoglossum officinale L. R 4 7 8 10 H. strumosus L. EW 3 4 6 7 8 *Echium vulgare L. R 4 9 H. tuberosus L. E 2 3 4 5 6 7 8 9 10 Hackelia deflexa (Wahl.) Opiz. EW 4 Heliopsis helianthoides (L.) Sw. E 2 34 6 8910 H. vir.iriniana (L.) J.M. Johnst. EW 2 3 4 6 8 9 10 var. scabra (Dun.) Fern. *Lappula ££hinata Gilib. R 2 3 4 6 7 10 Iva xanthifolia Nutt. AR 3 4 5 7 8 9 10 L. redowskii (Hornem.) Greene R 2 3 4 6 7 Kuhnia eupatorioides L. p 2 3 4 678910 Lithospermum canescens p 2 3 4 678910 var. corymbulosa T. & G. (Michx.) Lehm. Lactuca canadensis L. R 3456789 L. incisum Lehm. p 2 3 4 678910 L. floridana (L.) Gaertn. EW 2 3 4 8 9 10 L. latifolium Michx. w 6 L. ludoviciana (Nutt.) DC. p 2 3456 89 Mertensia virginica (L.) Pers. w 3 4 7 L. oblongifolia Nutt. p R 2 3 4 5 6 7 8 9 10 E 6 9 (L. pulchella (Pursh) DC.) Myosotis verna Nutt. Onosmodium mo/le Michx. E p 2 3 4 678910 *L. serriola L. (L. scariola L.) R 2 3456 8910 (0. occidentale Mack.) Liatris aspera Michx. p 2 3 4 5 6 7 8 9 10 BRASSICACEAE (Mustard Family) (L. sphaeroitka Michx.) 6 L. punctata Hook. p 2 345678910 *Alliaria officinallis Andrz. E W 3 4 5 678910 Lygodesmia juncea (Pursh) Hook. P 2 3 4 5 6 7 8 9 10 Arabis canadensis L. W 2 4 5 6 L. rostrata Gray p 2 A. hirsuta (L.) Scop. P Microseris cuspidata p 2 3 4 6 7 8 var. /l)cnocarpa (M. Hopk.) Roll. 3 4 6 (Pursh) Sch. Bip. (Agoseris A. shortii (Fern.) GI. W 2 6 cuspidata (Pursh) Raf.) *Armoracia rusticana R Prenanthes alba L. E p 8 Gaerrn., Mey. & Schreb. 9 10 P. aspera Michx. P 234567 *Barbarta vulgaris R. Br. R 10 Ratibida columnifera P 2 3 4 6 7 8 *Brassica kaber (DC.) Wheeler R 345678 (Nutt.) Woot. & Stand!. *B. nigra (L.) Koch R 2345678 R. pinnata (Vent.) Barnh. P 23456 910 *Camelina microcarpa Andrz. R 2 10 Rudbeckia hirta L. P 3 5678 10 *C. sativa (L.) Crantz. R 5 7 R. laciniata L. A E 3 8 10 *Capsella bursa-pastoris R 2 3 4 5 6 7 8 9 10 Senecio glabellus Poir. W 8 9 10 (L.) Medic. S. plattensis Nutt. P 2 3 4 6 8 9 10 *Cardaria draba (L.) Desv. R 3 10 S. pseudaur.us Rydb. W 4 *Chorispora tenella DC. R 2 3 var. semicordatus (Mack. & Bush) T.M. Barkley Dentaria laciniata Muhl. W 6 9 Silphium integrifolium Michx. P 678910 *Descurainia pinnata (Walt.) R 2 3 4 6 7 8 9 10 S. laciniatum L. P 2 3 4 6 8 9 10 Britt. var. brachycarpa (Richards.) Fern. S. perfoliatum L. E 2 3 4 6 8 10 *D. sophia (1.) Webb R 3 4 5 8 9 Solidago canadensis L. E R 2 3 4 678910 Draba reptam (Lam.) Fern. P 2 3 8 9 (S. altissima L.) *Erucastrum gallicum R 3 S. flexicaulis L. W 8 9 10 (Willd.) Schulz S. gigantea Ait. P 234 6 8910 *Erysimum cheiranthoides L. R 2 3 4 S. missouriensis Nutt. P 2 3 4 5 6 7 8 9 10 E. incompicuum (Wacs.) MacM. p 2 S. mollis Bartl. P 4 *H esperis matronalis L. R 2 3 4 8 10 S. nemoralis Ait. p 2 3 4 5 6 7 8 9 10 *Lepidium campestre (L.) R. Br. R 2 s. rigida L. p 2 3 4 5 6 7 8 9 10 *L. demiflorum Sch rad. R 2 3 4 6 7 8 10 S. speciosa Nutt. p 2 3 4 5 6 7 8 9 10 *L. vir.irinicum L. R 2 678910 S. ulmifolia Muhl. w 7 8 9 10 *Nasturtium officinale R. Br. A 7 8 *Sonchus arvensis L. R 7 Rorippa sinuata (Nutt.) Hitchc. A 3 4 678910 *S. asper (L.) Hill R 3 5 6 7 8 10 *Sisymbrium altissimum L. R 2 4 6 7 9 10 *S. oleraceus L. R 5 6 8 *S. loeselii L. R 2 3 4 8 *Taraxacum laevigatum R 2 *S. officinale (L.) Scop. R 2 4 5 6 7 8 10 (Willd.) DC. *Thlaspi arvense L. R 2 3 4 5 678910 *T. officinale Weber R 2 3 4 678910 CAMPANULACEAE (Bluebell Family) *Tragopogon dubius Scop. R 2 3 4 678910 Campanula americana L. E 1 2 3 4 5 6 8 9 10 Verbesina alternifolia (1.) Britt. W 6 8 9 10 Lobelia inflata L. E 8 (Actinomeris alternifolia (L.) DC.) L. siphilitica L. A 3 4 5 6 8 9 10 Vernonia haldwini Tott. P 678910 CANNABACEAE (Hemp Family) var. interior (Small) Schub. *Cannabis sativa L. R 1 2 3 4 678910 V. missurica Raf. P 6 8 9 10 Humulus /upulus L. E 1 2 3 4 6 8 10 BALSAMINACEAE (Jewel-weed Family) CAPPARACEAE (Caper Family) Impatiens biflora Walt. A 2 3 8 10 Cleome serrulata Pursh P R 1 2 3 4 5 6 7 8 9 I. pa/Iida Nutt. A W 2 3 4 678910 Polanisia dodecandra (L.) DC. PR 3 4 5 6 BERBERIDACEAE (Barberry Family) ssp. trachysperma (T. & G.) II tis *Berberis vulgaris L. R 8 (P. trachysperma T. & G.) Caulophyllum thalictroides w 2 3 4 CAPRIFOLIACEAE (Honeysuckle Family) (L.) Michx. Lonicera dioica L. w 3 4 5 6 8 9 Podophyllum peltatum L. E 9 10 var. glaucescens (Rydb.) Butters BETULACEAE (Birch Family) L. prolifera (Kirchn.) Rehd. E W 6 8 Cory/us americana Walt. E 4 5 6 7 8 9 10 *L. tatarica L. R 2 3 4 5 6 8

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SambtKNS canademis L. E 2 3 4 5 6 7 8 9 10 E. nutan.r Lag. (E. pres/ii Guss.) R 2 3 4 5 8 Symphoricarpos occidmtalis Hook. E P 2 3 4 5 6 7 8 9 *E. podperae Croiz. (E. esula L.) R 2 3 4 5 6 7 8 S. orbiculatm Moench E P 4 6 7 8 9 10 E. serpens H.B.K. R 2 3 5 8 9 10 L. Triosteum perfoliatum W 4 6 7 8 9 10 E. serpyllifolia Pers. R 3 5 6 7 8 L. Viburnum lentago E W 4 6 8 9 E. spathulata Lam. R 5 6 8 CARYOPHYLLACEAE (Pink Family) (E. dictyosperma Fisch. & Mey.) Cerastium nutan.r Raf. E W 9 E. stictospora Englem. P R 2 3 4 *Dianthm armeria L. R 10 FABACEAE (Legume Family) *Holosteum umbellatum L. R 3 Amorpha canescens Pursh p 1 2 3 4 5 6 7 8 9 10 *Lychnis alba Mill. R 2 10 A. fruticosa L. A 2 3 4 5 6 7 8 9 IO *Paronychia canademis (L.) Wood R 6 8 Amphicarpa bracteata (L.) Fern. E W 2 3 4 5 6 7 8 9 *Saponaria officinalis L. R 2 3 5 6 8 9 10 Apios americana Medic. E 8 10 Silene antirrhina L. R 5 7 8 10 Astragalm agrestis p *S. cucubalm Wibel R 6 Doug!. ex. D. Don *S. dichotoma Ehrh. R 8 A. canademis L. p 2 3 4 5 6 7 8 9 10 *S. noctiflora L. R 5 8 A. crassicarpm Nutt. var. p 2 3 4 5 6 8 9 S. stellata Ait. f. W 2 3 4 6 8 9 10 craJSicarpm (A. caryocarpm Ker.) *Stellaria mdia (L.) Cyr. R 3 6 7 A. lotiflorus Hook. P 2 3 4 678910 *Vatraria segetalis (Neck.) R 7 8 A. missouriensis Nutt. P 2 3 Gke. (Saponaria vatraria L.) Cassia fasciculata Michx. E R 2 3 4 678910 CELASTRACEAE (Bittersweet Family) (Chamaecrista fasciculata (Michx.) Greene) Celastrus scandms L. E 1 2 3 4 5 678910 Cercis canademis L. W 5 6 7 8 9 IO *Euonymm alatm Sieb. E 3 *Coronilla varia L. R 2 3 4 5 6 E. atropurpureus Jacq. W 3 4 678910 Crotalaria sagittalis L. p 3 CERATOPHYLLACEAE (Honewott Family) Dalea enneandra Nutt. p 2 3 4 5 6 7 8 9 IO Ceratopbyllum demersum L. A 4 Desmanthm illinoensis Michx. AP 678910 CHENOPODIACEAE (Goosefoot Family) Desmodium canademe (L.) DC. E p 5 8 Chenopodium album L. R 1 2 3 4 5 6 8 10 D. canescens (L.) DC. E 3 5 6 8 C. berlandieri Moq. R 2 3 4 5 D. cuspidatum (Muhl.) Loud. AE 8 10 C. desiccatum A. Nels. P R 2 3 4 5 6 8 D. glutinosum (Muhl.) Wood. w 2 3 4 8 9 10 (incl. C. praeterico/a Rydb. and D. illinoense Gray p 3 7 8 9 10 C. /eptopbyllum Nutt.) D. paniculatum (L.) DC. EW 5 6 7 8 10 C. bybridum L. ER 2 3 4 5 6 8 10 Gleditsia triacanthos L. EW 2 3 4 5 6 7 8 9 10 (C. gigantospermum Aellen) Glycyrrhiza lepidota Pursh AE 2 3 4 5 6 8 9 IO C. standleyanum Aellen ER 2 3 4 5 9 Gymnocladus dioica (L.) K. Koch W 2 3 4 5 6 7 8 9 IO *Kochia scoparia (L.) Schracl. R 2 3 4 5 6 7 8 9 10 Lespedeza capitata Michx. P 2 3 4 5 6 7 8 9 IO *Sa/so/a iberica Sennen & Pau R 2 3 4 5 6 7 8 9 10 *L. cuneata (Dumont) G. Don R 8 9 (S. kali L.) *L. stipulacea Maxim. R 5 6 7 8 10 CONVOLVULACEAE (Morning-glory Family) *LotUJ corniculatm L. R 2 3 4 7 8 10 *Convolvulm arvensis L. R 1 2 3 4 8 10 *Medicago lupulina L. R 2 3 4 5 6 7 8 10 C. sepium L. R 1 2 3 4 6 8 10 *M. sativa L. R 2 3 4 5 6 7 8 9 10 *Ipomota hederacea (L.) Jacq. R 8 10 *Melilotm albm Desr. R 2 3 4 ~ 6 7 8 9 IO *I. parpurea (L.) Roth R 1 5 6 8 10 *M. officinalis (L.) Lam. R 2 3 4 5 6 7 8 9 IO CORNACEAE (Dogwood Family) OX'Jtropis /ambertii Pursh p 2 3 4 5 6 7 8 9 IO Cornm drummondii Mey. E 1 2 3 4 5 6 7 8 9 10 Petalostemon occidmtale p 2 3 4 5 6 7 8 (C. asperifolia Michx.) (Gray) Fern. CRASSULACEAE (Stonecrop Family) P. parpureum (Vent.) Rydb. p 2 3 4 5 6 7 8 9 10 Penthorum sedoides L. A 1 4 8 10 Psoralea argopbylla Pursh p 2 3 4 5 6 7 8 9 10 CUCURBITACEAE (Gourd Family) P. esculenta Pursh p 2345678 9 10 Cucurbita foetidissima H.B.K. E 8 Robinia pseudoacacia L. ER 3 4 5 6 7 8 Echinocystis lobata E 2 3 4 5 6 8 10 Shrankia nuttallii (DC.) Stand!. P 5 7 p 4 5 6 7 8 (Michx.) T. & G. Strophostyles helvola (L.) Ell. p 5 6 7 8 9 10 Sicyos angulatm L. E 2 3 4 5 7 8 9 10 S. /eiosperma (T. & G.) Piper 3 ELAEAGNACEAE (Oleaster Family) *Trifolium bybridum L. R 3 4 5 6 8 9 10 *Elaeagnm angmtifolia L. R 1 2 3 4 5 6 7 8 *T. pratense L. R 2 3 4 5 6 Shepherdia argentea Nutt. P 2 3 *T. procumben.r L. R 8 EUPHORBIACEAE (Spurge Family) *T. repens L. R 2 3 4 5 6 8 10 10 Acalypha ostryaefolia Ridd. A R 8 10 Vicia americana Muhl. E 2 4 5 6 7 9 A. rhomboidea Raf. E R 2 5 6 9 10 var. americana p A. virginica L. ER 2 3 4 5 6 8 9 V. americana Muhl. 2 3 4 6 7 8 Croton capitatUJ Michx. PR 7 var. minor Hook 8 C. monanthogynm Michx. PR 4 *V. villosa Roth R 3 4 Euphorbia corollata L. E 4 5 6 7 8 9 10 FAGACEAE (Beech Family) E. cyathophora Murr. E 2 3 4 5 6 7 8 9 10 QtarrUS alba L. W 5 10 (E. heteropbylla L.) Q. borealis Michx. f. W 3 4 5 6 7 8 9 10 *E. cyparissias L. R 5 6 7 vat: maxima (Marsh.) Ashe (Q. rubra L.) E. dmtata Michx. p R 2 3 4 5 6 7 8 9 10 Q. imbricaria Michx. W 6 8 E. glyptosperma Engelrn. R 2 3 4 5 6 7 8 9 10 Q. macrocarpa Michx. P W 1 2 3 4 5 6 7 8 9 10 E. hexagona Nutt. PR 2 3 5 Q. muhlenbergii Engelrn. W 7 8 9 10 E. maculata L. R 2 3 4 5 678910 Q. prinoides Willd. W 9 10 (incl. E. supina Raf.) Q. stellata Wang. W 9 E. marginata Pursh p R 2 3 4 5 6 7 8 9 10 Q. velutina Lam. W 5 6 7 8 9 10

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p 9 10 FUMARIACEAE (Fumirory Family) Mirabilis albida (Walt.) Heimerl. C orydalis micrantha R 1 2 3 7 8 10 M. hirsuta (Pursh) MacM. P R 2 7 (Engelm.) Gray 6 8 9 10 M. linearis (Pursh) Heimerl. P 3 Dicentra cucullaria Bernh. W 2 3 4 6 8 9 10 GERANIACEAE (Geranium Family) M. nyctaginea (Michx.) MacM. R 2 3 4 5 8 10 OLEACEAE (Olive Family) Geranium maculatum L. W 5678910 HIPPOCASTANACEAE (Horse-chestnut Family) Fraxinus americana L. w 2 3 4 5 6 7 8 9 10 *Aesculus glabra Willd. W 3 7 8 9 F. penmylvanica Marsh. w HYDROPHYLLACEAE (Waterleaf Family) var. penmylvanica 2 678910 El/isia nyctelea L. E 12345678910 F. penmylvanica Marsh w Hydrophyllum appendiculatum w 4 8 10 var. subintegerrima (Vahl) Fern. ONAGRACEAE (Evening Primrose Family) Michx. 6 7 8 9 10 H. virginianum L. w 2 3 4 6 8 Calylophus serrulatus P 1 2 3 4 5 HYPERICACEAE (St. John's-wore Family) (Nuce.) Raven (Oenothera serrulata Nutt.) 2 6 8 9 10 *Hypericum perforatum L. R 2 3 4 6 10 Circaea lutetiana L. W 3 4 H. sphaerocarpum Michx. A 8 ssp. canadensis (L.) Asch. & Mag. JUGLANDACEAE (Walnut Family) (C. quadrisulcata (Maxim.) Franch & Sav.) Carya cordiformis w 2 3 4 678910 Gaura coccinea Pursh P 2 3 4 678910 (Wang.) K. Koch G. longif/ora Spach (G. biennis L.) P 6 8 9 10 C. avata (Mill.) K. Koch w 6 7 8 9 10 G. parviflora Doug!. P R 2 3 4 6 7 8 9 10 C. tomentosa N utc. w 6 8 Oenothera biennis L. R 3 4 8 10 p j uglam cinera L. w 5 6 8 0. laciniata Hill 6 7 }. nigra L. w 2 3 4 5 6 7 8 9 10 0. strigosa (Rydb.) Mack & Bush P 2 3 4 6 7 8 10 LAMIACEAE (Mine Family) ssp. canavirem (Steel) Munz Agastache nepetoides (L.) 0. Ktze. E 1 3 4 8 9 10 OXALIDACEAE (Wood Sorrel Family) A. scrophulariaefolia W 3 4 6 8 9 Oxalis dil/enii Jacq. E R 2 3 4 5 6 8 10 (Willd.) 0. Ktze. 0. stricta L. (incl. E R 2 3 4 5 6 7 8 10 *Glecoma hederacea L. R 2 3 0. europea Jard.) p Hedeoma hispida Pursh p R 2 3 4 5 6 7 8 9 0. violacea L. 2 3 4 6 7 8 10 H. pulegioides (L.) Pers. E p 8 PAPAVARACEAE (Poppy Family) lsanthus brachiatus (L.) B. S. P. p R 2 3 6 Sanguinaria canademis L. W 1 2 3 4 6 8 9 *Leonurus cardiaca L. R 2 3 4 5 6 7 8 PHRYMACEAE (Lopseed Family) Lycopus americanus Muhl. A 2 3 8 Phryma leptostachya L. W 1 2 3 4 8 9 10 L. asper Greene A 3 4 6 PHYTOLACCAECEAE (Pokeweed Family) *Marrubium vulgare L. R 8 9 Phytolacca americana L. E 8 10 Monarda fistulosa L. var. fistulosa E 2 3 4 5 6 8 9 10 PLANTAGINACEAE (Plantain Family) *Nepeta cataria L. R 2345678 10 Plantago aristata Michx. R 5 6 7 8 *Prune/la vulgaris L. R 2 5 6 8 10 *P. lanceolata L. R 4 6 7 8 10 Pycnanthemum pilosum N ucc. EW 8 *P. major L. R 3 8 P. tenuifolium Schrad. EW 8 P. patagonica Jacq. P R 2 3 4 678910 Salvia reflexa Hornem. p R 2 3 4 678910 var. patagonica (P. purshii R. & S.) (S. lanceolata Willd.) P. rugelii Decne. R 2 3 4 7 8 10 Scutel/aria lateriflora L. A 2 3 4 8 9 P. virginica L. R 7 8 9 10 Teucrium canademe L. var. E 2 3 4 678910 PLATANACEAE (Sycamore Family) occidentale (Gray) McCI. & Epl. Platanus occidentalis L. W 6 7 8 9 10 (incl. T. occidentale Gray) POLEMONIACEAE (Phlox Family) T. canademe L. E 3 5 6 8 Phlox divaricata L. w 2 3 4 678910 var. virginicum (L.) Eat. ssp. laphamii (Wood) Wherry LINACEAE (Flax Family) P. pilosa L. ssp. fulgida p 2 3 4 678910 Linum rigidum Pursh var. rigidum P 1 2 3 4 6 7 8 (Wherry) Wherry L. sulcatum Riddell P 1 2 3 4 678910 POLYGALACEAE (Milkworc Family) LOASACEAE (Sand Lily Family) Polygala verticil/ata L. P 2 3 4 5 6 8 9 10 Mentzelia decapetala (Pursh) P 2 3 6 POLYGONACEAE (Buckwheat Family) Urban & Gilg. *F agopyrum es cu lentum Moench R 6 MALVACEAE (Mallow Family) Polygonum achoreum Blake R 2 345678 10 *Abutilon theophrasti Medic. R 1 2 ~ 4 678910 (incl. P. erectum L.) Cal/irhoe involucrata P 6 7 8 10 P. arenastrum Jord. ex Bar. R 2 3 4 6 8 10 (T. & G.) Gray (P. aviculare L.) *Hibiscus trionum L. R 3 8 9 10 P. coccineum Muhl. A 2 3 4 678910 *Malva neglecta Wallr. R 2 3 9 *P. convolvulus L. R 2 3 4 6 7 8 *M. rotundifolia L. R 4 8 *P. hydropiper L. A 4 *Sida spinosa L. R 8 10 P. lapathifolium L. A 3 4 5 6 7 8 10 Sphaeralcea coccinea (Pursh) Rydb. P 2 6 P. pemylvanicum L. A 2 3 4 5 6 7 8 9 10 MENISPERMACEAE (Moonseed Family) *P. persicaria L. A 2 3 4 5 6 7 8 10 Menispermum canademe L. E W 1 2 3 4 5 678910 P. punctatum Ell. A 4 6 8 9 MONOTROPACEAE (Indian Pipe Family) P. ramosissimum Michx. R 3 5 6 8 10 Monotropa uniflora L. W 8 10 P. scandens L. R 5 7 8 9 10 MORACEAE (Mulberry Family) P. virginianum L. w 4 8 9 10 *Maclurapomifera(Raf.)Schneid. R 1 8 9 10 *Rumex acetosel/a L. R 5 6 7 8 *Marus alba L. R 1 2 3 4 678910 R. altissimus Wood AR 2 45678910 M. rubra L. W 4 678910 *R. crispus L. R 2 3 4 5 6 7 8 9 10 NYCTAGINACEAE (Four O'Clock Family) R. mexicanus Meisn. AR 6 10 https://scholarworks.uni.edu/pias/vol92/iss5/12 10 Novacek et al.: The Vegetation of the Loess Hills Landform Along the Missouri Riv

VEGETATION OF LOESS HlllS 209

PORTULACACEAE (Purslane Family) G. obtusum Bigel. W Claytonia virginica L. W 4 6 8 9 10 10 G. trif/orum Michx. W *Portulaca oleracea L. R 3 1234 6 8910 5 6 7 Hedyotis nigricans (Lam.) Fosb. p 4 5 6 7 8 9 10 PRIMULACEAE (Primrose Family) (Houstonia nigricans {Lam.) Fern.) Androsaa occidentalis Pursh P 1 Lysimachia ciliata L. A E 1 2 4 RUTACEAE (Rue Family) 6 8 10 Zanthoxylum americanum Mill. E 1 2 3 4 RANUNCULACEAE (Buttercup Family) 5 6 7 8 9 10 Actaea alba (L.) Mill. W SALICACEAE (Willow Family) 9 10 *Popu/us alba L. (A. pacbypods Ell.) R 1 2 3 4 5 8 P. de/toides Marsh. A. rubra (Ait.) Willd. AE 1 2 3 4 5 6 7 8 9 10 w 3 4 6 8 9 10 P. tmnuloides Michx. E Anemone canadensis L. AE 2 3 4 5 6 7 8 7 9 10 *Salix alba L. R 4 5 A. cylindrica Gray p 2 3 4 5 6 7 8 9 10 S. amygdaloides Anderss. A 2 3 A. patens L. p 2 3 4 4 5 6 7 8 9 10 5 S. discolor Muhl. A 2 3 4 A. virginiana L. w 2 3 4 6 S. exigua Nutt. 8 9 10 A 2 3 4 5 6 7 8 9 10 Aquilegia canadensis L. w 2 3 4 5 6 8 9 10 ssp. interior (Rowlee) Cronq. Clematis pitcheri T. & G. E 6 8 10 (incl. S. interior Rowlee) C. virginiana L. E 2 3 4 5 6 10 *S. fragi/is L. R 2 3 *Delphinium ajacis L. R 4 S. humilis Marsh. 10 p 23456 7 8 9 10 D. tricorne Michx. E 4 6 10 S. nigra Marsh. A 5 6 7 8 9 10 D. virescens Nutt. P 2 3 4 5 6 7 8 9 10 S. rigida Muhl. A 123456 8 9 10 Isopyrumbiternatum(RM.)T. &G. W 10 SANTALACEAE (Sandalwood Family) Riln1111CUlus abortivus L. E 2 3 4 5 6 7 8 9 10 Comandra umbtllata (L.) Nutt. E P 1 2 3 4 5 6 7 8 9 10 R. fascicularis Muhl. w 6 ssp. umbellata R. 1?£11f'Vatus Poir. w 8 10 (incl. C. richardsiana Fern.) R. sceleratus L. A 2 6 SAXIFRAGACEAE (Saxifrage Family) Thalictrum dasycarpum w 2 3 4 5 6 8 9 10 Heuchera richardsonii R. Br. E P 4 5 7 8 Fisch. & Lall. (incl. H. hirsuticaulis (Wheeler) Rydb.) RESEDACEAE (Reseda Family) Ribes americanum Mill E P 2 3 8 *Reseda lutea L. R 5 R. missouriense Nutt. E 2 3 4 5 6 7 8 9 10 RHAMNACEAE (Buckthorn Family) R. odoratum Wendi. f. E 1 3 4 7 8 Ceanothus americanus L. E 5 6 7 8 SCROPHULARIACEAE (Figwott Family) var. pitcheri T. & G. Agalinis aspera (Bench.) Britt. P 1 2 3 4 5 6 7 8 9 10 C. herbaceus RM. E P 1 2 3 4 5 6 7 8 9 10 (Gerardia aspera Dougl.) var. pubescens (T. & G.) Shinners Castilleja sessiliflora Pursh p 2 3 4 5 6 7 8 9 JO (C. ()l)afus Desf.) Mimulus ringens L. A 4 5 8 10 *Rhamnus catharticus L. E R 3 Penstemon albidus Nutt. p 2 3 R. lanceolatus Pursh E 3 4 5 6 7 8 9 P. cobaea Nutt. p 8 var. glabratus GI. P. digitalis Nute. p 8 9 10 ROSACEAE (Rose Family) P. gracilis Nutt. p 5 Agrimonia pubescens Walle. E 2 3 4 9 10 P. grandiflorus Nutt. p 2 3 4 5 678910 A. striata Michx. E 4 6 Scrophularia lanceolata Pursh E 2 3 4 6 Amelanchier arborea w 678910 S. marilandica L. E 2 3 4 8 9 10 (Michx.) Fern. *Verbascum blattaria L. R 5 6 Crategus ,,,,,I/is (T. & G.) Scheele E 2 3 4 5 6 7 8 9 10 *V. thapsus L. R 2 3 4 5 6 7 8 9 JO Fragaria mca L. E W 2 3 4 8 Veronica peregrina L. R 3 4 5 6 7 8 9 10 var. americana Potter var. peregrina F. virginiana Duchne. E P 2 3 4 5 6 7 8 9 10 SOLANACEAE (Nightshade Family) var. illinoensis (Prince) Gray Datura stramonium L. R 4 5 6 7 8 9 JO Geum canadense Jacq. E 2 3 4 5 6 7 8 9 10 *Lycium halimifolium Mill. R 3 5 6 8 9 Potentilla arguta Pursh P 2 3 4 5 8 Pbysalis heteropbylla Nees. PR 3 5 78910 P. 11Qr11Bgica L. A 2 3 4 5 7 8 P. virginiana Mill. ER 2345678910 *P. rerta L. R 2 4 10 (incl. P. longifolia Nutt. and Prunus americana Marsh. E 2 3 4 5 6 7 8 9 P. subglabratum Mack. & Bush) P. mexicana Wars. E 9 *Solanum americanum Mill. E R 2 3 4 6 7 8 10 P. serotina Ehrh. E 3 4 5 6 7 8 9 10 (S. nigrum L.) P. virginiana L. E 2 3 4 5 6 7 8 9 10 *S. carolinense L. R 5 6 7 8 9 10 Pyrus ioensis (Wood) Bailey E 3 4 5 6 7 8 9 10 *S. rostratum L. R 2 3 4 5 6 7 8 9 10 Rosa arkansana Porter P 2 3 4 5 6 7 8 9 10 STAPHYLEACEAE (Bladdernut Family) (incl. R. su/fulta Greene) Stapbylea trifolia L. w 3 4 678910 R. blanda Ait. E 2 3 4 5 6 THYMELAEACEAE (Leatherwood Family) R. setigera Michx. P 9 10 *P asserina annua Wikstr. R 2 3 4 6 R. woodsii Lindi. E 2 3 4 5 8 (Thymelaea passerina (L.) Coss. & Germ.) (incl. R. fendleri Crepin) TILIACEAE (Basswood Family) Rubus allegheniensis Potter E 6 7 8 9 Ttlia americana L. w 1 2 3 4 5 6 7 8 9 10 R. f/agellaris Willd. E 4 ULMACEAE (Elm Family) R. idaeus L. E 5 6 7 8 9 10 Ce/tis occidentalis L. w 1 2 3 4 5 6 7 8 9 10 ssp. sachalinensis {Levi.) Focke UImus americana L. w 2 3 4 5 6 7 8 9 10 R. ocridentalis L. E 1 2 3 4 5 6 7 8 9 10 *U. pumila L. R 2 3 4 5 6 7 8 RUBIACEAE (Madder Family) U. rubra Muhl. w 2 3 4 5 6 7 8 9 10 Galium aparine L. E 1 2 3 4 5678 10 u. thomasi Sarg. w 3 5 6 7 9 G. circaezam Michx. w 3 4 6 8 9 10 URTICACEAE (Nettle Family) G. concinnum T. & G. w 8 9 10 Laportea canadensis (L.) Wedd. A E 1 2 3 4 5 6 7 8 9 10

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C. tetanica Schkuhr A 9 Parietaria pensylvanica Muhl. AE 2 3 4 5 6 8 10 Cyperus erythrorhizar Muhl. A 3 5 8 9 10 Pi/ea pumi/a (L.) Gray A 2 3 4 10 C. escu/entus L. AR 345678 10 Urtica dioica L. ssp. AER 2 3 4 5 6 7 8 9 10 C. ferruginescem Boeck!. A 2 3 4 5 8 10 graci/i.r (Ait.) Seland. VERBENACEAE (Vervain Family) (C. odoratus L.) R 4 6 9 10 Verkna bracteata Lag. & Roclr. R 1 2 3 4 5 6 8 9 10 C. strigosus L. Scirpus atrwirens Willd. A 1 2 3 4 *V. canadensis (L.) Britt. R 6 10 S. va/idus Yahl. A 2 3 V. hastata L. A 4 DIOSCOREACEAE (Yam Family) V. stricta Vent. PR 1 2 3 4 5 6 7 8 9 10 Dioscorea ~i//osa L. w 8 10 V. urticifo/ia L. AE 1 2 3 4 5 6 7 8 9 10 VIOLACEAE (Violet Family) IRIDACEAE (Iris Family) Sisyrinchium campestre Biclcn. P 1 2 3 4 5 6 7 8 9 10 Viola canadensi.r L. w 1 2 3 4 JUNCACEAE (Rush Family) (incl. V. rugu/osa Greene) A 3 4 V. mi.rsouriensis Greene w 8 10 ]uncus dudleyi Wieg. 6 8 10 V. pe,datifida G. Don p 2 3 4 5 6 8 9 ]. tenui.r Willd. R 3 LEMNACEAE (Duckweed Family) V. pratinco/a Greene RW 4 5 7 9 7 8 V. pubescem Ait. w 2 3 4 6 8 9 10 Lemna minor L. A 4 8 (incl. V. pensy/vanica Michx.) Spirodt/a polyrhiza (L.) Schie id A V. sororia Willd. E 2 3 4 5 6 9 10 LILIACEAE (Lily Family) 4 5 6 7 8 10 V. viarum Pollard p 4 5 7 8 9 10 A//ium canadense L. var. canadense P 1 VITACEAE (Grape Family) *Asparagus officinalis L. R 1 2 3 4 5 7 8 10 Ampelopsi.r cordata Michx. w 8 10 *Conval/aria majalis L. R 3 Parthenoci.rsus quinquefo/ia (L.) E 2 3 4 5 6 7 8 9 10 Erythronium albidum Nutt. W P 2 3 4 6 8 9 10 Planch. (incl. var. mesochoreum Knerr) P. vitaaa (Knerr) Hitchc. E 2 3 4 6 9 10 Li/ium canadense L. EW 3 10 Vitis cinerea Engelm. E 8 9 10 ssp. michiganense (Farw.) Boivin V. riparia Michx. E 2 3 4 5 6 7 8 9 10 & Cody (L. michiganense Farw.) V. vu/pina L. E 6 9 10 Polygonatum biflorum (Walt.) Ell. E 123456 8 9 10 Z¥GOPHYLLACEAE (Caltrop Family) (P. cana/iculatum (Muhl.) Pursh) *Tribu/us terrestris L. R 1 R o 1ll Smilacina racemosa (L.) Desf. W 3 4 6 MAGNOLIOPHYTA:LILIOPSIDA (Monocots) S. stel/ata (L.) Desf. W 2 3 4 6 10 ALISMATACEAE (Water Plantain Family) Smilax ecirrhata (Engelm. E W 2 3 4 6 7 10 A/i.rma plantago-aquatica L. A 4 ex Kunth) Wats. var. americana R. & S. S. herbacea L. E 3 4 8 10 Sagittaria latifo/ia Willd. A 3 var. lasioneuron (Small) Rydb. ARACEAE (Arum Family) S. hi.rpida Muhl. E 2 3 4 5 6 7 8 9 10 Arisaema dracontium Schott W 6 8 9 10 Uvu/aria grandiflora Sm. W 3 10 A. triphy//um (L.) Schott W 1 2 3 4 5 6 7 8 9 10 Yuait glauca Nutt. P 2 3 4 5 6 7 8 9 10 COMMELINACEAE (Spiderwort Family) ORCHIDACEAE (Orchid Family) *Comme/ina communi.r L. R 6 10 Cora//orhiza odontorhiza w 4 8 Tradescantia bracteata Small P R 1 2 3 4 5 6 7 8 9 10 (Willd.) Nutt. CYPERACEAE (Sedge Family) Cypripedium calceolus L. w 10 Came amphibo/a Steucl. W 1 3 4 8 9 var. pubescem (Willd.) Correll var. turgida Fern. (C. grisea Wahl.) Orchi.r s{Jectabili.r L. w 6 8 9 10 C. anmctens Biclcn. A 3 10 POACEAE (Grass Family) C. artitecta Mackenz. A 8 9 *Agropyron pectiniforme R 2 3 4 C. atherodes Spreng. A 3 6 Roem. & Schult. C. bickne//ii Britt. P 2 *A. repens (L.) Beauv. R 2 3 4 8 C. b/anda Dewey W 2 3 4 5 7 8 9 10 A. smithii Rydb. PR 2 3 4 5 6 8 C. brevior (Dew.) Mack. E P 2 3 4 5 6 7 8 9 10 Agrarti.r hyemali.r (Walt.) p 4 C. cepha/ophora Muhl. E 4 6 8 B.S.P. var. hyemalis C. davisii Schwein & Tort. A 8 9 10 *A. stolonifera L. R 2 3 4 6 8 10 C. eburnea Boott. W 3 4 8 (incl. A. alba L.) C. gravida Bailey var. gravida A 2 3 4 6 7 8 9 Andropog~n gerardi Vitman p 2 3 4 5 6 7 8 9 10 C. htliophi/a Mackenz. P 2 3 4 5 8 A. ha/Iii Hack. p 3 (C. pensylvanica Lam. var. A. scoparius Michx. p 2 3 4 678910 digyna Bock!.) Aristida longi.reta Steud. p 2 3 C. hitchcockiana Dew. W 4 7 8 A. oligantha Michx. p R 2 3 4 5 6 7 8 9 10 C. hystericina Muhl. ex. Willd. A 2 3 4 8 Bouteloua curtipendu/a p 2 3 4 5 6 7 8 9 10 C. /anuginara Michx. A 4 5 6 9 10 (Michx.) Torr. (C. lasiocarpa Ehrh.) B. gracili.r (H.B.K.) Griffiths p 2 4 5 6 8 9 10 C. meadii Dewey P 2 5 9 B. hirsuta Lag. p 2 4 7 8 9 10 C. moles ta Mack. P 4 5 6 8 10 Brachyelytrum erectum w 3 4 6 7 10 C. norma/i.r Mackenz. W 6 8 (Schreb.) Beauv. C. o/igocarpa Schkuhr W 3 4 7 8 *Bromus inermis Leyss. R 2 3 4 5 6 7 8 9 10 C. pensy/vanica Lam. W 3 5 8 *B. japonicus Thunb. R 2345678 C. rarea Schkuhr ex Willd. W 2 3 4 6 8 9 10 B. latiglumi.r (Shear) Hitchc. E 3 5 var. pusi//a Peck ex Howe B. pubescem Willd. R 3 4 6 8 10 (incl. C. contJO!uta Mackenz.) *B. racemosus L. R 6 8 C. saximontana Mack. W 2 3 (B. commutatus Schrad.) C. sparganioitks Muhl. A 3 4 8 9 10 *B. secalinus L. R 3 5 C . .rpmzgelii Dewey W 2 3 4 8 *B. tectorum L. R 12345678 10

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B11Chloe dactyloides P 3 9 10 Phalaris arundinacea L. A (Nutt.) Engelm. 2 3 4 6 8 9 10 *Phleum pratense L. R Calamovilfa longifolia P 2 3 4 6 8 9 10 2 4 6 8 PhragmiteJ austral is (Cav.) Trin. A (Hook.) Fern. 8 (P. communis Trin.) Cenchrus longispinus (Hack.) Fern. R 2 3 4 5 6 7 8 10 *Poa compmsa L. P R (C. pa11Ciflorus Bench.) 2 3 6 8 9 *P. pratensis L. E P R 2 3 4 Cinna arundinacea L. W 6 7 8 9 10 4 9 P. sy!VeJtris Gray W *Dactylis glomerata L. R 9 10 2 3 4 6 7 8 9 10 Schedonnardus paniculatus P 2 3 Diarrhena americana Beauv. W 3 4 7 9 (Nutt.) Trel. var. obovata Gl. *S ecale cereale L. R 2 3 4 5 6 7 8 10 Dicanthelium acuminatum (Sw.) P 9 *Setaria faberii Herrm. R 5 6 7 8 9 10 Gould & Clark var. villosum (Gray) *S. glauca (L.) Beauv. R 2 3 4 5 6 7 8 9 10 Gould & Clark (Panicum praecocius (S. luteJcens (Weigel) Hubb.) Hitchc. & Chase) *S. italica (L.) Beauv. R 2 9 D. latifolium (L.) Gould & Clark W 10 *S. verticillata (L.) Beauv. R 2 3 4 6 8 9 (Panicum latifolium L.) *S. viridis (L.) Beauv. R 2 3 4 6 7 8 9 10 D. leibergii (Vasey) Freckm. P 2 6 9 Sorghastrum avenaceum P 2 3 4 6 7 8 9 10 (Panicum leibergii (Vasey) Scribn.) (Michx.) Nash (S.nutans (L.) Nash) D. oligosanthes (Schultes) Gould P 2 4 5 6 7 8 9 *Sorghum halepense (L.) Pers. R 4 6 8 10 var. scribnerianum (Nash) Gould Spartina pectinata Link. A 2 4 6 7 8 9 10 (Panicum scribnerianum Nash) Sphenopholis obtusata E 4 6 8 10 D. oligosanthes (Schultes) Gould P 2 4 6 (Michx.) Scribn. var. wilcoxianum (Vasey) Gould Sporobolus asper (Michx.) Kunth P 2 3 4 6 7 8 9 10 & Clark (Panicum wilcoxianum Vasey) Vat. asper *Digitaria ischaemum R 2 3 4 8 9 10 S. cryptandrus (Torr.) Gray P 2 3 4 6 7 8 9 10 (Schreb.) Muhl. S. vaginiflorus (Torr.) Wood P R 2 3 4 6 7 8 9 *D. sanguinalis (L.) Scop. R 3 4 5 6 7 8 10 var. neglectus (Nash) Scribn. *Echinochloa crusgalli (L.) Beauv. R 2 3 4 5 6 7 8 9 10 (incl. S. neglectus Nash) E. muricata (Beauv.) Fern. R 2 3 5 6 7 8 10 S. vaginiflorus (Torr.) Wood P R 3 var. microstachya Wieg. var. vaginiflorus (E. pungens (Pair.) Rydb.) Stipa coma/a Trin. & Rupr. P 3 *Eleusine indica (L.) Gaerrn. R 3 6 7 8 10 S. spartea Trin. P 2 3 4 6 Elymus canadensis L. P 2 3 4 5 6 7 8 9 10 S. viridula Trin. P 2 3 E. villosus Muhl. W 2 3 4 5 6 7 8 9 10 Tri dens flavus (L.) Hi tchc. A 8 9 10 (E. striatus Willd.) (Triodia flava (L.) Smyth) E. virginicus L. E 2 3 4 6 7 8 10 POTAMOGETONACEAE (Pondweed Family) E. wiegandii Fern. E W 3 4 Potamogeton foliosus Raf. A 4 8 *Eragrostis cilianensis (All.) R 2 3 4 6 7 8 9 10 P. illinoensis Morang A 8 E. Mosher P. nodosus Pair. A 7 8 E. pectinacea (Michx.) Nees. P R 2 3 4 6 7 8 9 10 P. pectinatus L. A 7 8 E. spectabilis (Pursh) Steud. P R 2 3 4 6 7 8 9 10 TYPHACEAE (Cattail Family) E. trichodes (Nutt.) Wood R 2 3 Typha angustifolia L. A 2 3 4 8 FeJtuca obtusa Biehler W 2 3 4 5 6 8 9 10 T. latifolia L. A 4 F. octoflora Walt. P R 2 3 7 8 *F. pratensis Huds. (F. eliator L.) R 2 3 4 6 8 9 Glyceria striata (Lam.) Hitchc. A 2 3 4 6 8 *Hordeum jubatum L. R 2 3 4 6 7 8 10 REFERENCES H. pusillum Nutt. R 4 6 7 8 10 Hystrix patula Moench W 2 4 6 10 ALBERTSON, F. W., and J. E. WEAVER. 1945. Injury and death or Koeleria pyramidata (Lam.) Beauv. P 2 4 6 7 8 9 recovery of trees in a prairie climate. Ecol. Monog. 15:393-433. (K. cristata (L.) Pers.) AIKMAN, J. M. 1926. Distribution and structure of the forests of eastern Leersia oryzoides (L.) Sw. A 2 Nebraska. Univ. of Nebraska Studies 26: 1-75. L. virginica Willd. W 2 3 4 6 8 9 10 AIKMAN, J. M. , AND C. L. GILLY. 1948. A comparison of the forest floras *Lolium perenne L. R 3 8 along the Des Moines and Missouri Rivers. Proc. Iowa Acad. Sci. 55:63- *Miscanthus sacchariflorus R 3 73. (Maxim.) Hack. BARKLEY, T. M., ed. 1977. Atlas of the flora of theGreat Plains. Iowa State Muhlenbergia cuspidata P 2 3 4 5 6 7 8 9 10 University Press, Ames. (Torr.) Rydb. BENNETT, H. D. 1948. Notes on the Cyperaceae oflowa. Proc. Iowa Acad. M. /rondos a (Pair.) Fern. 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