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NATURAL HISTORY SURVEY

Erigenia FEB 1 8 1998 LIBRARY Number 15 November 1997

Journal of the Illinois Native Society Erigenia

Number 15, November 1997

The Illinois Native Plant Society Journal

The Illinois Native Plant Society is dedicated to the preservation, conservation, and study of the native and vegetation of Illinois.

Society Erigenia is named for Erigenia bulbosa (Michx.) Executive Committee of the (harbinger of spring), one of our earliest Nutt. President: Marty Vogt blooming woodland plants. The first issue was PreSIDENT-ELECT: Mark Basinger in August 1982. published Past President: Glen Kruse Treasurer: Jon Duerr ® 1997 The Illinois Native Plant Society Copyright Recording Secretary: Ray Boehmer (ISSN 8755-2000) Membership: Lorna Konsis Editor, Erigenia: Gerould Wilhelm Erigenia Staff 324 N. York Road, Elmhurst, IL 60126

EDITOR: Gerould S. Wilhelm Editor, The Harbinger: Ken Konsis Copy Editor-. Barbara A. Johnson 20301 E. 900 North Road, WestviUe, IL 61883 Typesetting and Design: Linda A. Masters E-MAIL: [email protected] Production: George D. Johnson MEMBERS-AT-LARGE: Todd Bittner, George Johnson, Mike Mason, Cover Illustration Pat Neighbors, Jack Shouba

Mark Mohlenbrock of five Original drawing by Chapter Presidents examples of our Illinois flora: Camassia scilloides

(Raf.) Cory, Carex pensylvanica Lam., Dodecatheon Central {Springfield): Mike Mason meadia L., Quercus macrocarpa Michx., and East-Central {Urbana): Ken Robertson Tradescantia virginiana L. Forest Glen {WestviUe): Reggie Romine Northeast {Naperville): George Johnson IS dedicated to Mark's father. Dr. Robert H. It Southern {Carbondale): Mark Basinger Mohlenbrock, who has devoted his career to the study of our state's flora.

Membership

Yearly membership includes a subscription to Erigenia and the quarterly newsletter. The Harbinger. For more information write to:

nUnois Native Plant Society Forest Glen Preserve 20301 E. 900 North Road WestviUe, Illinois 61883 Erigenia

Number 15, November 1997

Table of Contents

Floristic Quality Assessment for Vegetation in Illinois, A Method for Assessing Vegetation Integrity

John B. Taft, Gerould S. Wilhelm, Douglas M. Ladd, and Linda A. Masters 3

Abstract 3

Introduction 3 Background on Assessment Methods for Natural Areas 4

Principles of Plant Ecology Relevant to Floristic Quality Assessment 5

Methods 6 Terminology and Concepts 7 Coefficient of Conservatism 7 Ecological and Community Integrity 9 Floristic Quahty Index 9 Natural Area 10 Physiognomy 10

Apphcation of Floristic Quality Assessment 10 Floristic Quality Assessment Apphcation Computer Program 10

Sur\'ey Intensity and Spatial and Temporal Scales of Survey Units 11

Data Analysis 11

Results and Discussion 11 Examples of Floristic Quahty Assessment 13

Example 1: Four Herbaceous Communities 13

Example 2: Two Mesic Upland Forest Communities 15

Example 3: Two Southern Flatwoods Communities 15 Testable Paradigm 17

Conclusions 18

Glossan,' 19

Acknowledgments 20

About the Authors 20

Literature Cited 21

Appendix: Vegetation of lUinois Database 24

ERJGEXL4. Number 15, November 1997, pp. 3-95 © 1997, Illinois Native Plant SocieU'

Floristic Quality Assessment for Vegetation in Illinois A Method for Assessing Vegetation Integrity

John B. Taft\ Gerould S. Wilhelm', Douglas M. Ladd', and Linda A. Masters'

ABSTRACT: Floristic Qualirv' Assessment (FQA) is proposed as a metliod to assess floristic integrity in Illinois. For

the application of FQA, each taxon in the Illinois vascular flora was assigned an integer from to 10 termed a

coefficient of conservatism (C). Two basic ecological tenets that the coefficients represent are that plant species differ

in their tolerance to and disturbance rypes, and that plant species display varying degrees of fidelity to

integrit)-.

With these principles as a guide, the coefficient applied to each taxon represents a rank based on observed behavior

and panerns of occurrence in Illinois plant communities and our confidence that a taxon is remnant (natural area) dependent. Species given a C value of 0-1 are taxa adapted to severe disturbances, particularly anthropogenic disturbances, occurring so frequently that often only brief periods are available for growth and reproduaion. Species

ranked with a C value of 2-3 are associated with somewhat more stable, though degraded, environments. Those species with coefficients 4-6 include many dominant or matrLx species for several ; they have a high consistency of

occurrence within given community types. Species with C values 7-8 are taxa we associate mostly with natural areas, but that can be found persisting where the habitat has been degraded somewhat. Those species with coefficients 9-10

are considered to be restriaed to high-quality natural areas. A floristic qualirv- index (FQI) and a mean coefficient of conservatism (C) are two of the values derived from floristic inventory data. Other derived parameters include species richness, relative importance, percent of taxa that are native and adventive, number of rare species, and guild diversity (including wetness and conservatism ranks, and

physiognomic classes). We suggest that FQA is a promising tool that can be used to discriminate natural qualiry- of vegetation on the Illinois landscape and to make time-series comparisons in ecological studies. We suggest the use of

certain parametric and nonparametric statistical tests, such as analysis of variance, mean-separation techniques, and

goodness-of-flt tests, that can aid in distinguishing nonrandom differences in floristic quality.

Introduction ^^"^ design (size and shape) of areas often are influenced by quahtative differences in vegetation. This paper

Patterns of vegetation are rehable indicators of describes a method for discerning floristic integrity in

several biotic and abiotic factors. Biotic interactions Illinois.

among species and abiotic factors (including edaphic and Floristic Quahty Assessment (FQA) is a method chmatic characteristics) influence plant assemblages in that uses a floristic quahty mdex (FQI), mtroduced by many complex ways that lead to the expression of WUhehn (1977) and Swink and Wilhehn (1979, 1994),

differences at the species, community, and ecosystem and modified here for the Illin ois vascular flora. FQA

levels. Overlying these influences is disturbance history. integrates FQI with other vegetation parameters. These

Disturbances differ in frequency, intensity, and duration. include mean coefficient of conservatism, species Infrequent disturbances of low intensity and short richness, percent native and adventive species, guild duration can have relatively neghgible impacts on the diversity for various physiognomic and conservatism

integrity of a plant community. However, as frequency, classes, number of threatened and endangered species, intensitv, and/or duration increase, damage and and type of natural community and grades following the ukunately degradation can occur, resuhingm predictable classification and grading criteria estabhshed by the changes in plant communiry characteristics, panicularly Illinois Natural Areas Inventory (White 1978). FQA can

composition. Differentiating vegetation on the basis of be used to make spatial as well as time-series

level of degradation is an important step in attempting to comparisons, and in this way FQA can be effective in conserve . For example, preserve selection tracking vegetational changes in restoration.

Illinois Natural Histon" Survey, 607 E. Peabody Dnve, Champaign, IL 61820 - Conservation Research Institute, 324 N. York Road, Elmhurst, IL 60126

' The Nature Consen-ano-, 2800 S. Brenra-ood, St. Louis, MO 63144 FLORISTIC QUALITt- ASSESSMENT FOR VEGETATION IN ILLINOIS reconstruction, or control situations, and in evaluating Illinois Natural Areas Inventory (INAI) was an effon to parameters across environmental and disturbance conduct a comprehensive county-by-cotmty inventory of gradients. Species abundance measures also can be natural areas (White 1978). A method for assessing included in FQA evaluations. In this paper we discuss habitat quahties was developed for the INAI, to aid in key terminology, describe the method of FQA for the the identification of significant remnants of natural

Illinois vascular flora, offer suggested appUcations and communities. Several site characteristics were integrated statistical analyses, and urge experimental tests of in the natural community grading method, including hypotheses related to floristic quaUty. We caution that aspects of vegetation such as perceived successional stage, any vegetative assessment based on a single index is Ukely evidence of disturbance, and presence and relative- to be insufficient to account for all possible relevant abundance patterns for species characteristic of particular aspects. As an introduction, a shon history of habitat habitats and levels of disturbance. The INAI used a assessment methods, particularly those used in Illinois, is discontinuous, determinant grading scale, where habitat given. Selected issues in plant-community ecology are remnants received a grade of A, B, C, D, or E (defined included as background information. under Illinois Natural Areas Inventory Grades in the glossary) in accordance with increasing degrees of Background on Assessment Methods for Natural disturbance reflected in the community characteristics Areas (White 1978). Herein, reference to INAI natural areas will be made with capital letters (Natural Area). Methods for making qualitative assessments of Independent of the INAI was the development of a biological communities have had expanding roles in the method of natural area identification using a continuous, conservation of lands and habitats as development mdeterminate scale called a Natural Area Rating Index pressures increase. An Index of has (NARI) based on floristic composition (Wilhelm 1977, been developed based on characteristics of fish Swink and Wilhehn 1979, Wilhehn and Ladd 1988). The community composition (Karr et al. 1986) and for ant NARI was developed as an aid m discriminating natural populations (Majer and Beeston 1996). Migratory bird quahty of vegetation among open lands in the Chicago species have been ranked according to perceived region and is based on an index derived from the prioritization of habitat and species conservation goals composition of vascular plants at a site. Because (Hunter ei al. 1993). There is a recognized need for vegetation spans the entire disturbance gradient from an simple, sensitive, readily interpretable, and ecologically urban lot or cropland to relatively "pristine" habitats, a meaningful methods of classifying vegetation according continuous scale offers some refinement to quahtative to levels of ecological integrity (Keddy et al. 1993), distinctions of floristic characteristics. This characteristic particularly for use by the nonspeciahst (Grime 1974). In in particular made the Natural Area Rating Index a addition, a rapid method of assessment often is needed, valuable tool for identifying degraded remnants of native particularly when evaluating large portions of a vegetation having recovery potential, given appropriate landscape (e.g., proposed highway-construction corridors management. that cross numerous remnants of native vegetation and Principal criticisms of the method have included the natural community types). Ordination techniques can be following: 1) the coefficient range chosen, which began used effectively to examine relationships among with -3 for the most invasive adventive species and vegetation (and abiotic) sample data. However, these increased by intervals of 1 to a coefficient of 10 indirect measures are not particularly rapid and are value- (coefficients of 15 and 20 were used for very rare species), neutral, limiting their application for making quahtative 2) a lack of consideration for species abundance, and 3) assessments of biotic communities, particularly in the the subjective nature of coefficients assigned to each heterogeneous landscape. taxon and differences in interpretation of them. Two developments have been key in the Refinements of the method led to a revised scale of identification and protection of natural areas in Illinois. coefficients that ranged from to 10; all adventive species First, in 1963, the Illinois Nature Preserves Commission were assigned an asterisk with a numerical value of 0. For was formed to administer the development of a system of clarity the method was renamed Floristic Quahty nature preserves as representative examples of the natural Assessment (Swink and Wilhehn 1994). history of the state. Second, during the mid 1970s, the

ERJGENL4 November 1997 FLOFUsnc Qu.\LiT\' Assessment for Vegetation in Illinois

Abundance measures for species, as described later native biological diversity (original condition) is unhkely in this paper, are readily accommodated in FQA and under normal circumstances. Degraded lands have lost should be included in any assessment of vegetation when some aspects of ecosystem structure such as species possible. It is important to acknowledge that natural composition. Degraded lands are termed derelict when quahty assessments are subject to bias and require more land use becomes very hmited (Brown and Lugo 1994). or less subjective judgements at the current state of They can be further distinguished as those that can be community ecological science (Crovello 1970). The FQA restored to nearly original condition through some method, though subjective, permits dispassionate and management effort, rehabilitated to a condition repeatable appHcation because its value judgements are somewhat similar to the original but where predetermined. Assessment methods based on FQA have compositional differences remain (Lovejoy 1975) or, at

been developed in Ohio (Andreas and Lichvar 1995), best, reclaimed to a limited degree in severe cases such as

Michigan (Herman et al. 1996), Missouri (Ladd 1993), strip mining.

and southern Ontario (Oldham et al. 1995), and Many midwestern plant communities were formed elaborated on by Masters (1997). and historically maintained with landscape-scale

In addition to investigating the current composition processes that include disturbances such as periodic fire, and structure of the vegetation, any assessment of as well as grazing or browsing impacts by large vegetation quality should also give attention to herbivores (Anderson 1983, 1990). Additional

degradation factors at the landscape, ecosystem, and considerations in regard to disturbance regimes are

community levels, and the historic (presettlement) and addressed under Ecological Integrity in the methods contemporary natural disturbance regimes. section and in the discussion of succession below. Different survival strategies have evolved among Principles of Plant Community Ecology Relative organisms for coping with disturbances. Among the to Floristic Quality Assessment hypotheses of mechanisms to account for these strategies are MacArthur and Wilson's (1967) r- and K-selected Plants can be classified into groupings based on a species, Grubb's (1977) regeneration niche, and Grime's variety of species characteristics such as physiognomy, regenerative flexibihty for ecological amphtude (Grime phenology, and ecophysiology, and habitat 1974, Grime et al. 1988). For the latter, species survival characteristics such as soil type, Hght, moisture, and strategies are considered to be shaped by an equHibrium disturbance regimes. In heavily developed landscapes among the ecological forces of competition, stress, and such as Illinois, and similarly in Great Britain, disturbance. These forces serve as axes for ordinating contemporary' anthropogenic disturbances to vegetation species' responses in Grime's "triangles." These are often the predominant influences on composition ordinations yield three general life strategies referred to (Hodgson 1986), and thus are dominant among assembly as the C-S-R model: competitors, stress tolerators, and and response rules for communities {sensu Keddy 1992). ruderals. Species son selectively into this disturbance matrix; the Whittaker (1965) recognized that plant communities opponunistic species become more common as the could be described by three basic dominance-diversity landscape becomes more unstable. The coefficients of curves that differ in the cumulative proponion of conservatism used in FQA are an attempt to categorize importance of species. Species-poor communities are species according to their response to levels of habitat strongly dominated by a few taxa; in communities with degradation. high species richness, no species is strongly dominant. Three general topics in plant community Many communities are intermediate, composed of a few ecology—disturbance ecology, the maintenance of taxa with high relative abundance and many intermediate diversity, and successional theory—are particularly and rare species. Several studies suggest that intermediate relevant to the concept of floristic quahty because they levels of available resources (nutrients and physical provide a framework for understanding patterns and factors) support the greatest diversity (Tilman 1986, trends, particularly at the population and community Ashton 1989, Tiknan and Pacala 1993). Intermediate levels. Disturbance is a general term referring to any levels of disturbance also appear important in the penurbation. Plant communities can be damaged when maintenance of diversity in many communities (Connell severely disturbed and are degraded when recovery to its 1978, Tester 1989), although the maintenance of peak

ERIGEN14 November 1997 FLORJSTIC QU.^LITl- ASSESSMENT FOR \'EGETATION IN ILLINOIS

levels of plant in some panicularly fire- condition. Without knowledge of the immigration dependent systems appears to require frequent potential for replacement species, we have no way to perturbations (Walker and Peet 1983). predict accurately the composition or structure of The groupings described above are useful in that subsequent communities. Consequently, the assumptions they attempt to provide both order to species of directional trends in secondary succession leading assemblages and predictabihty regarding the rate and toward the original (presettlement) plant community direction of changes in response to such things as human- may have lost relevance where the landscape is highly influenced disruptions. In all of the models, spatial and fragmented. Using terminology from disturbance temporal heterogeneity within and among habitats is a ecology (e.g., degraded, dereUct) when describing the critical factor in the maintenance of species diversity at natural condition of a site may be clearer than the community level of organization or higher. speculations about successional phases (e.g., early

Succession is a frequently used term for the successional, late successional) of disturbed vegetation. description of vegetational change through time. Apparently, many degraded sites persist in states of

Clements (1936) argued that succession was an orderly perpetual botanical purgatory (Taft 1996). and predictable process leading to a "climax" community, depending on climate and other factors. Typically, Methods primar)- succession is initiated on exposed parent materials, while secondary succession involves changes in In Floristic Quahty Assessment (FQA), floristic vegetational characteristics following events such as inventory data are used to calculate several parameters of abandonment of cropland, clear-cutting of forests, or vegetation. These include the following measures, each

drainage of wetlands. However, climax is an ambiguous defined and described in greater detail in subsequent

term (Crawley 1986) and appears to have httle practical sections: 1) species richness, 2) floristic quality index

meaning if considered without regard to regional (FQI), 3) mean coefficient of conservatism (C), 4) guild disturbance regimes or historical antecedents. In diversity (frequency distribution among physiognomic

landscapes such as those in the Midwest, the and conservatism classes), 5) species relative imponance,

development of many native plant communities was 6) number and percent rare and adventive species, and 7) dependent on anthropogenic fires, the practice of which wetness characteristics. These data are presented in a dates back to the postglacial era. In such circumstances summary table. The FQI and C are derived from the cessation of fire could be regarded as a "disturbance." coefficients of conservatism assigned to each taxon in the

Indiscriminate use of the term succession may Illinois vascular flora. Important terms related to FQA obfuscate the fact that cenain plant communities require are defined in the glossary; key concepts and periodic penurbations such as fire for the maintenance of terminology tmderlying the general philosophy of FQA

structural characteristics and compositional diversity. If are discussed below. Recommendations for applying and unidirectional successional trends in these communities analyzing selected FQA results are included. We were among our conservation goals, we would not be undertake this effort with the knowledge that concerned with vegetational changes such as those from contending with the entire flora of Illinois overextends prairie communities to forest-hke assemblages or from our collective experience to some extent. The judgments biodiverse oak-dominated woodlands to maple- presented here are based primarily on our cumulative

dominated forest. Such changes, however, often result in total of over 60 years of botanical and ecological field

a loss of species richness (Wilhehn 1991, Taft et al. 1995), study throughout Illinois and the Midwest. particularly in our highly fragmented landscape, where Botanical nomenclature in the text and appendix species immigration, needed to compensate for local approximates Mohlenbrock (1986). Many hybrids and

extirpations of species, is seriously challenged. certain subspecific taxa such as forma, are not included; The term succession, when used for changes in siome varieties were omitted when we perceived them vegetation following severe anthropogenic disturbances, not to vary ecologically from the typical variety. may be misleading. Without detailed experimental Recently recorded species for Illinois are also included.

studies of various disturbance factors on different The hsting of species in Appendix I is not to be

vegetation types, we do not know how extensively interpreted as a definitive flora of Illinois; it is intended vegetation "succeeds" or recovers to a more stable

Eugenia November 1997 FLORISTIC QU-^LITi' ASSESSMENT FOR VEGETATION IN ILLINOIS solely to be reference database for applications of other end of the spectrum, species vinually restricted to

Floristic Quality Assessment. natural areas in Illinois received C values of 10. All 957

The list in Appendix I comprises 2,091 native taxa non-native species were assigned asterisks ("') and are and 955 non-native taxa, for a total of 3,046 taxa, treated as Os in the calculations for site indices (FQI and compared with Mohlenbrock's (1986) total of 3,203 taxa, C). These calculations are further discussed in comments which included 101 hybrids. It is beyond the scope of under Floristic Quahty Index below and in the glossarv-. this paper to hst currently accepted nomenclatural Species native to Illinois, but also occurring escaped from s}-nonomy for each taxon; such a list soon would be out cultivation (e.g., Pinns spp.), should be ranked as non- of date. Unfortunately, scientific names of plants in native species when found in such situations.

Nonh America are in a slate of flux, with often With these criteria for designating coefficients, our confhcting nomenclatural treatments (Little 1979, approach was somewhat different from past effons. For Kanesz and Kartesz 1980, Soil Conservation Service example, we are not intending to estimate the degree to

1982, Gunn et al. 1992, Morin 1993, and Kartesz 1994). which a species is restricted to a cenain habitat, or to

Only a single common name for each taxon is offered, gauge its modahty according to Curtis (1959). Many despite the fact that many taxa are known by a variety of relatively conservative taxa (e.g., Amorpha canescens, colloquial names. An attempt was made to use common Baptism leucophaea, Cypripedmm candidum, Drosera names with the widest appeal; they are taken mostly rotundifolia, Gaylussacia baccata, Osmunda cinnamomea, from Mohlenbrock (1986), Swink and Wilhehn (1994), Ceanothns americanus, and Viola pedata) occur regularly and Robenson (1994). in more than one plant community, as defined by White

Physiognomic designations are subject to and Madany (1978). In addition, we were not attempting interpretation. Terms such as annual, biennial, perennial, to estimate rarity, although some circularity of reasoning shrub, and tree sometimes imperfectly depict the habit of was unavoidable when evaluating very rare taxa known plants, but for the purposes of guild formation in FQA only from a few natural areas. analysis, such designations can be useful in describing Reasons for rarity m the Illinois flora are many (Taft structural differences or changes. 1995) and iaclude several recognized by Rabinowitz

(1981). Scale of inference influences what is considered a Terminolog}^' and Concepts rare species. Many species that are rare within the poUtical boundaries of Illinois are abundant elsewhere. Coefficient of Conservatism. For the application of Many conservative taxa are not at risk of extirpation FQA, each taxon in the Illinois vascular flora was from the state, but are regionally quite rare because of assigned an integer from to 10, termed a coefficient of habitat loss and degradation. Commonness and rarity of conservatism (C). The coefficients represent two basic plant species in England have been considered in terms of ecological tenets: plants differ in their tolerance to ecological, taxonomic, and evolutionary processes within disturbance type, frequency, and amplitude, and plants a landscape characterized by tremendous habitat loss and display var}-ing degrees of fidehty to habitat integrity. degradation (Hodgson 1986). Although common and With these principles as a guide, the C value appUed to rare species at local scales may be strongly correlated to each taxon represents a relative rank based on observed measurable traits, there is so much variability in behavior and patterns of occurrence in Illinois plant ecological, taxonomic, and evolutionary characteristics of communities and our confidence that a taxon is remnant species at the statewide scale (Schwanz 1993) that these (natural area) dependent. The authors reached consensus groupings do not address consistently our criteria for on these coefficients through committee effort and, in conservatism. Although rarity is not a criterion for some cases, with consultation from reviewers of the assignment of C values, it forms a pan of the matrix of manuscript. For certain taxa we supplemented our field parameters in FQA. experience by examining range maps (Mohlenbrock and The coefficients, in part, can be considered in terms Ladd 1978) and reviewing comments regarding habitats of Grime's (1974) survival strategies. Species given a C in several floras (Deam 1940, Gleason 1952, Steyermark value of 0-1 correspond to Grime's ruderal species and 1963, Sheviak 1974, Mohlenbrock 1986, Swink and those with a C value of 2-3 correspond to ruderal- Wilhelm 1994). The native species most successful in competitive species. This broad, combined species guild badly damaged habitats were given C values of 0. At the includes taxa adapted to frequent and severe disturbances.

ER1GESL4 November 1997 Floristic Qllvlit.- Assessment for Vegetation in Illinois including anthropogenic disturbances that often result in such as Asclepias tuberosa and Oxalis violacea. These only brief opportunities for reproduction. Under such a species are occasional to common in degraded habitats in disturbance regime, only species capable of maintaining far southern Illinois, but in central and nonhern Illinois populations under such conditions are present, including they are more restricted to remnant areas. In these those that rapidly grow, flower, and produce fruits (e.g., instances, we assigned an intermediate value such as 5.

Ambroiia trifida, Amaranthus rudis, Cassia fasciadata, The species having C values of 7-10 are less clearly

Conyza canadensis, Engeron annuus, Impatiens capensis, ahgned with Grime's model. Grime et al. (1988) defined Lactuca canadensis, Lepidmm virginicum, Oxalis stncta, the third guild, stress tolerators, to include species that

Panetaria pensylvanica, and Vulpia octoflora). Many are persist where plant productivity is continuously limited capable of persisting in seed banks, and some have wind- by the environment. A more specific definition of dispersed seeds—two strategies that allow species to son Grime's stress tolerators, offered in an editorial by into suitable, newly disturbed habitats. Some longer-lived Duffey (1986), includes "species that are slow-growing, species capable of persisting with frequent disturbances long-hved and often rather immobile plants of infertile such as siltation, flooding, and grazing are also included habitats or late-successional vegetation." Our criteria for in this group {e.2,., Acer saccharinum, Crataegus pruinosa, species ranked with coefficients 7-10 allow the inclusion

Gleditsia triacanthos, Populus deltoides, Ribes missouriense, of species that may tolerate stress, but through a variety

Rubus occidentalis, and Symphoncarposorbiculatus). These of mechanisms. More germane to quahtative floristic taxa constitute approximately 17% of our native flora. In assessments, these taxa do not tolerate much habitat conjunction with many of the adventive elements, these degradation. Consequently, this guild includes some species now dominate the contemporary Illinois annuals and biennials (e.g., Agalinis gattingeri, Draba landscape. cuneifolia, Hottonia inflata, Iresine rhizomatosa, Lechea

Species assigned coefficients 4-6 correspond roughly intermedia, Oenothera linifolia, Polygala incamata, and to Grime's competitors. These include many dominant Utricularia minor). However, like Grime's stress or matrix species for several habitats (e.g., Andropogon tolerators, most taxa in this guild are long-hved gerardii, Carex artitecta, C. pensylvanica, C. stricta, Carya perennials (e.g., Asclepias meadii, A. viridiflora, Carex ovata, Panicum virgatum, Quercus alba, Schizachyrium disperma, C. peduncidata, C. prasina, Clitoria manana, scoparmm, and Sorghastrum nutans) and species that are Cystopteris bulbifera, Gymnocarpium dryopteris, Lilium often expected, or have high consistency, in a given philadelphicum, Mentzelia oligosperma, Sedum telephioides, community type (e.g., Aesculus glabra, Ansaema S. tematum, and Talinum parviflorum, Woodsia ilvensis). triphyllum. Delphinium tricome. Phlox divancata, The species ranked with coefficients 7-8 include taxa we Silphmm integrifolium, Smilacina racemosa, Thalictrum associate mostly with natural areas but which can be dioicum, Trillium recurvatum, and Zizia aurea). Many found persisting where the habitat has been degraded can persist with hght to moderate disturbances for somewhat (e.g., Actaea pachypoda, Caulophyllum intermediate periods, but may decline with an increase in thalictroides, Ceanothus americanus, Lysimachia intensity, frequency, or duration of disturbance. Some quadriflora, Peltandra virginica. Phlox pilosa, Spigelia species that are range restricted, such as Boltonia manlandica, and Viburnum rufidulum). Like the matrix decurrens, which is hsted as a threatened species by the species (C values of 4-6), if the disturbance resulting in U.S. Fish and Wildlife Service (USFWS 1988) and the degradation increases in frequency, intensity, or

Illinois Endangered Species Protection Board (Herken duration, these taxa are expected to undergo reduction in

1991), and other species that are rare in Illinois such as population sizes and eventually be prone to local Scirpus paludosus, and Tradescantia hracteata, are included extirpation. Species with coefficients 9-10 are considered in the 4-6 category. In the contemporary Illinois to be restricted to relatively intact natural areas. landscape these species demonstrate considerable Though there is some commonahty between the tolerance to disturbance and even habitat degradation, C-S-R model (Grime et al. 1988) and the concept of but usually not to the extent characteristic of the ruderal- conservatism, we lack the experimental autecological competitor species guUd. evidence to ordinate species mto Grime's triangles.

On occasion, during the coefficient assessment phase Funher, species assigned C values of 7-10 do not fit of this project, we needed to evaluate taxa that consistently into Grime's C-S-R model, unless the stress- demonstrate regional behavioral differences in Illinois, tolerator guild is more broadly defined to include species

ERJGENL4 November 199' .

Florjstic Qu.^LiTi' Assessment for Vegetation in Illinois found primarily in semistable habitat remnants certain aboriginal practices, but also from the invasion of (sometimes referred to as "late-successional" adventive taxa. Adventive taxa in a system may sort into communities). disturbance or habitat niches, replace many native taxa Unfortunately, taxa included among each major over time, and interfere with rates of recovery processes cohon of coefficients (0-3, 4-6, 7-10) span a range that (Cohen et al. 1995). is too broad taxonomically, ecologically, and Measuring ecological integrity based on ecosystem physiognomically for any objective natural sorting to fimction alone may not provide the resolution needed to serv^e as a guide to species rankings that meet our guiding detect important changes. For example, biomass and principles for the coefficients of conservatism (see above) productivit}^ may not change dramatically in a palustrine For that reason, we based our judgments for the wetland impacted by siltation or altered flooding regimes assignments of the coefficients on the observed behavior where only a few tolerant taxa persist (e.g., Typha spp. of individual elements of the flora within the context of and Phalans arundinacea). However, the structural their Illinois ranges. Applying our judgments was integrity of a formerly diverse graminoid wetland is lost necessary since it is likely we will never have sufficient in this near monoculture, as when, for example, a experimental data to make predictions about floristic discharge wetland is convened to a surface runoff qualit}^ and ecological integrity for the diversity of wetland as a result of ambient watershed alterations. habitats, species, and disturbance regimes in Illinois using Integrity of both ecosystem structure and fimction is more ostensibly "objective" methods. Furthermore, rapid reduced in a heavily grazed (or browsed) woodland when and repeatable techniques for evaluating the integrity of soil compaction and intense herbivory result in losses in plant communities are needed now, particularly when moisture, nutrient availability, biomass, and diversity, as assessing complex patterns of vegetation in large sections well as changes in species composition. Floristic Quality of the landscape. Assessment addresses the structural aspects of ecosystem integrity. Ecological and Community Integrity. There are both functional and structural aspects of ecosystems. Floristic Quality Index. The FQI is a weighted index of

Ecosystem function involves the flow of energy and species richness (N), and is the arithmetic product of the matter, while structure is characterized by biotic average coefficient of conservatism (C) and the square- interactions, composition, and form. Ecological or root of species richness (V"N) of an inventory unit. The community integrity can be viewed as the degree to square-root transformation of N limits the variable which self-correcting properties are exhibited when an influence of area alone on species richness (Swink and ecosystem is exposed to disturbance (Regier 1993). Wilhekn 1979, 1994). In practice, it is possible for two

Natural disturbances are perturbations that occur sites with the same C to have different FQIs, and it is

routinely in a system and to which the component taxa possible for two sites with the same FQI to have

have tolerance or adaptations. They can occur at many different C values. Relatively degraded sites can have an

different scales. Tree falls and gopher mounds are FQI similar to or greater than high-quality natural areas

examples of small-scale perturbations. Fire is an example if they support a much greater native species richness.

of a large-scale natural disturbance in many Midwestern This can occur when there are substantial differences in

plant communities, and fire frequency and timing are size, levels of habitat heterogeneity, or inventor}^ effort

important determining factors for many community among compared sites. This and other relationships

characteristics. Fire absence can result in dramatic among the FQI, C, and N are illustrated in figure 1. changes in community structural characteristics (Taft Thus, rather than relying on a single index to describe

1997). Perturbations that exceed the intensity, frequency, floristic integrity, it is usually necessary to include more or duration of the natural disturbance regime can result than one parameter of the composition to estimate more

in loss of species that lack tolerance or adaptations to the precisely site floristic integrity.

new levels. When certain species, or assemblages of taxa, For the floristic parameters FQI, C, and N, we

are extirpated from a community, the system's capability recommend that calculations be made using all species

for restoration is diminished, and integrity is lowered. (native and adventive) as well as native species only. As Integrity can be lowered not only by the loss of noted previously, the estabUshment of exotic species in

species and the diminishment of abiotic processes and a natural community often can result in the replacement

ERIGESL-1 November 1997 Floristic Qu.\i.i-n- Assessment for Vegetation in Illinois

SiU B (N = x+E

FQI,

FQI, FQI3 FLORISnC QU.\LIPi- ASSESSMENT FOR VEGETATION IN ILLINOIS or systematic and may or may not be permanent) are spatially heterogenous habitats such as dune and swale used. This latter option is useful in tracking spatial and communities near Lake Michigan. temporal gradients of floristic integrity and wetness (see Wilhelm 1992), comparing data from large inventory- Data Analysis units, and conducting rapid ecological assessments When distinguishing the quahtative condition of (Heumann et al. 1993). habitat remnants using FQA, a typical goal is to

determine if the composition of two or more sites differs Survey Intensity and Spatial and Temporal Scales significantly from random expectation in the frequency of Survey Units distribution of the coefficients of conservatism. Three Measurements of an ecosystem or community propenies of the data influence the approach to be taken usually are at a smaller scale than the target system. Since to make this determination. If the sample data have an the FQI is a weighted index of species richness, larger acceptably normal distribution, have equal variances survey units and greater inventory efforts generally yield (homoscidastic), and are independent, then parametric higher indices of floristic quahty (figure 1), if increased statistics may be apphed (but see below). If, however, the size corresponds to increased richness of conservative data lack central-normal tendency or have unequal species. Determining the extent and configuration of the variances (heteroscidastic), a nonparametric or survey unit often is not a trivial question. Where the unit distribution-free method is suggested (independence of of floristic analysis is an isolated habitat fragment, the the data is assumed). Central-normal tendency usually sample area usually is readily apparent. In landscapes occurs with rank data when sample size (e.g., number of with more contiguous vegetation, however, determining species) is greater than about 50. the sample unit is less obvious and in many ways Methods used for examples in this text include dependent on the questions and interests of the parametric and nonparametric two-sample tests (e.g., investigation. Goals of the analysis may include a two-sample t-tests with unpooled variances, the Mann- complete species inventory, but it should be noted that Whitney U test, and the Kolmogorov-Smirnov [K-S] a complete inventory usually is not possible because of two-sample goodness-of-fit test). Comparisons of spatial and especially temporal variabihty in floristic multiple samples are tested with one-way analysis of composition. Thus, a single site visit will not variance (ANOVA), Tukey's Honestly Significant comprehensively account for all species in a community Difference (HSD) test, and the Kruskall-WaUis ANOVA. or site. With repeated visits over the growing season All statistical analyses were made using Systat version 7.0 most species that are actively growing at a site can be (Wilkinson 1997). identified, but this would not be adequate to evaluate the seed bank. Experience in midwestern vegetation types Results and Discussion has demonstrated that a single visit made between early

June and late August by a competent botanist can Coefficients of conservatism assigned to each taxon achieve a roughly 80% complete inventory. Subsampling, recognized here for the vascular flora of Illinois are spatially and temporally, is a practical option, presented in the appendix. The frequency distribution of panicularly where habitat integrity appears relatively coefficients of conservatism (0-10) for native species is uniform and the survey unit is too large to inventory left-skewed due to a strong peak at coefficient 10 (figure completely within the time available. Details of the 2). Distribution of species by physiognomic classes survey method and effort always should accompany any indicates that most species in the Illinois flora are reponing of results from FQA. Indiscriminate perennial dicot forbs, followed by adventive annual forbs comparisons of floristic quahty can be misleading if the (figure 3). Perennial sedges and grasses are notably more methods used for the evaluations are not similar. Where imponant m the native flora than in the adventive flora. area and heterogeneity of habitats or community The distribution of wetness coefficients for the native

remnants are considerably different, the mean coefficient and adventive flora of Uhnois (figure 4) shows that most of conservatism provides an area-independent variable taxa, including native and adventive, are (obhgate) upland

for comparisons of floristic quahty. Wilhehn (Swink and species; only about 91 adventive taxa are wetland species

Wilhelm 1979) provides insights for how to treat

ERIGE\i4 November 199' Floristic Qu.\Li-n- Assessment for \^egetation in Illinois FLORISTIC QUAUTi- ASSESSMENT FOR VEGETATION IN ILLINOIS tolerance to environmental stress (e.g., Opuntia humifusa, Examples of Floristic Quality Assessment QuercHS marilandica, and Vaccinium arhoreum). The The following three examples of Floristic Quality remaining flora—the species modal to relatively stable Assessment apphcation are not intended as proof or natural areas—may only loosely fit the stress-tolerator strenuous testing of the method, but rather as Despite a long history of habitat loss and guild. illustrations of cases where FQA and analytic methods there are remnant plant degradation in Illinois, are used in an anempt to differentiate vegetation quahty. communities in locahzed Httle-disturbed areas on both Example 1: Four Herbaceous Communities. Sites 1, 2, nutrient-poor and nutrient-rich sites. These remnants and 3 are prairie remnants. Site 1 is a high-quahty typically are rich in species and include many taxa that Natural Area; Sites 2 and 3 have been damaged by past lack ruderal characteristics, strong competitive abUities, disturbances but are dominated by native prairie species. or tolerance to high stress levels (e.g., Asdepias perenms, Site 4 is an old field with a histor)' of cultivation. All sites Caulophyllum thalictroides, Cypnpedium reginae, Dalea are similar in area (~2 to 4 ha) and were surveyed with Candida, Lilium philadelphicum, Trillium grandiflorum, similar inventory effons. Parameters of floristic quahty and Viburnum acerifolium). from all sites are compared m table 1. Comparisons of all assessment of ecosystem integrity based on a Any sites are shown for the cumulative proportion of species single index is likely to be insufficient to account for all by conservatism ranks (figure 6) and distribution pattern relevant aspects. For example, the FQI or when C of coefficients for each site using box plots (figure 7). reported alone can be misleading (figure 1). Also, species Data Analysis. Frequency of the coefficient of richness alone can be an insensitive indicator of habitat conservatism for each taxon present at each site are quahty, since it is possible for a degraded site to support normally distributed and meet the equal variance a similar or greater number of taxa than an undegraded assimiptions, although data from the old field (Site 4, n = site. Six measures of biological integrity for wetlands 51) are extremely skewed to the right (normahty test p = have been suggested by Keddy et al. (1993): species 0.084). Results are compared first using parametric diversity, indicator guilds, exotic species, rare species, techniques and then (as a precaution against possible plant biomass, and amphibian biomass. Diversity is nonnormal distributions and unequal group size) viewed as an essential indicator of integrity (Keddy et al. compared using results from nonparametric methods.

1993). However, instead of only measuring species For parametric tests, quahtative differences in richness, Keddy et al. (1993) also recommend assessing composition among all four sites were examined with guild diversity. FQA readily addresses the first four analysis of variance (ANOVA); multiple comparisons recommended measures, provides an index of wetness were examined with Tukey's HSD mean-separation indicates that a significant characteristics, and can be appHed to wetland and upland technique (table 2). ANOVA difference 0.000001) exists in floristic quahty among vegetation; moreover, it can be expanded to include other (p< the sites examined, as measured by the frequency community traits or particular interests such as INAI grades.

Table 1. Floristic integrity assessment summary data comparing four herbaceous commumties (Sites 1-4).

INAI Community Classsification INAI Grade Floristic Qu.\lit\' Assessment for Vegetation in Illinois

00 -

80 -

50 - FLORJSnC QU.\LIT\" ASSESSMENT FOR VEGETATION IN ILLINOIS

Table 3. Flonsac quality compansons among four Table 4. Floristic integrity assessment summary data comparing herbaceous communities. Probabilit\' levels shown two mesic upland forests. Woodland 1 has been grazed while compare results from two parametric tests and two Woodland 2, a smaller forest, apparendy has not. nonparametnc tests. See text for site descnpaons. The adjusted critical values for the two-sample tests are shown for these muluple compansons (e.g., p<0.0083).

Parametnc Tests Tukey HSD Test,

Site 1 Floristic Qu.\Li'n- Assessment for Vegetation in Illinois Florjstic Qu.\Li-n- assessment for Vegetation in Illinois

Dmax = 103 FLORISTIC QU.^LIT)- ASSESSMENT FOR VEGETATION IN ILLINOIS

duration of the disturbance are increased, species with diversity and composition also influence the FQI (e.g., regeneration intervals shorter than the disturbance habitat size, heterogeneity, disturbance history, and level frequencies (C guild 0-2[3]: Grime's ruderals) become of degradation). The mean coefficient of conservatism dominant, including many adventive species. (and quadrat-based sampling methods) provides an area-

The reverse of this paradigm is the recovery of a independent means of making qualitative comparisons degraded system. Restoration seeks to return damaged among sites. FQA can accommodate measures of species habitats or communities to their qualitative, abimdance and can accompany other measures of natural compositional, and structural states prior to degradation. community quaHty such as Illinois Natural Areas

We predict that both the FQI and C will increase at a site Inventory grades. We suggest testing the method by with the introduction of appropriate vegetation comparing floristic composition among sites and time management. In the Midwest, many studies have been intervals with known levels of disturbances and conducted, or are ongoing, that track the recovery of restoration activities using mean-separation techniques plant communities with the reintroduction of fire (Tester and analysis of guild diversity. Although similar results

1989; DeSekn and Clebsch 1991; Apfelbaum and Haney may be achieved with parametric statistics,

1991; Wilhekn ndMasters 1994; Taft, unpubhsheddata). nonparametric tests may be preferred for small sample

FQA offers a method to track changes in floristic sizes when all assumptions of parametric methods may composition that may be helpful in goal development not be met. and assessment (Masters 1997). Again, quantitative data provide the most accurate account of the relative abundance of species at a site. Species at low population levels sometimes are at greater risk of extinction (May

1973). If, by chance, most of the taxa with high C values are at low population levels, the species pool may be unstable and susceptible to rapid changes in the FQI and C. As always, the cost in time needed to collect and analyze quantitative data has to be contrasted with the ease, rapidity, and qualitatively thorough nature of floristic presence- absence data collection. Inventory goals will determine the approach to be taken.

Conclusions

We offer Floristic Quality Assessment (FQA) for the Illinois flora as a versatile, relatively rapid, dispassionate, and repeatable method for making qualitative assessments of plant communities and for assessing effectiveness of ecological restoration activities.

Using floristic inventory data, FQA summarizes several parameters of plant commimities, including a weighted measure of species richness (FQI), a mean coefficient of conservatism (C), guild diversity, proportion of adventive taxa, wetness characteristics, relative importance of native species, physiognomic characteristics, and rare species. The FQI is calculated from coefficients of conservatism (on a scale of 0-10) assigned to each taxon in the Illinois flora. The philosophy underlying the assignment of the coefficients

is a recognition that plant species are unequal contributors to habitat quahty. Factors that influence

ERJGENL4 November 1 997 FLORJSnC QU.VLITi' ASSESSMENT FOR VEGETATION IN ILLINOIS

Glossary propenies in an ecosystem or community exen themselves when that community is exposed to disturbance. Adventive - Not native to Illinois. Adventive is Natural Area - In a broad sense, a natural area is synonymous with the terms exotic and alien. Species that considered to be a natural community that is (presumably) have limited natural ranges in Illinois, but that are widely representative of the presettlement vegetation for the site. planted or escaped, such as Pinus strobus and Robinia This general definition includes all Natural Areas (INAI pseudoacacia, should be treated as adventive when sites graded A and B), but also areas that presently do not encountered outside their natural Illinois distributions, and meet the standards for the INAI but that, with assigned a C value of in the calculation of the floristic management and time, have potential for restoration to a qualitv index and mean coefficient of conservatism. community with floristic composition and diversity similar Coefficient of Conservatism (C) - An integer from to 10 to the presettlement condition. assigned to each taxon in the Illinois flora and used in Physiognomy - Broadly defined, physiognomy includes calculating the floristic quality index. Each value reflects an plant habit (architectural characteristics), life history, and estimate of a plant's tendency to be restricted to "natural certain taxonomic classes. Physiognomic classes assigned to areas" (see detailed description in methods section). The each taxon in the Illinois flora are Fern (including fern mean coefficient of conservatism (C) is calculated by allies), Annual Forb, Biennial Forb, Perennial Forb, summing all coefficients in an inventory unit and dividing Annual Grass, Perennial Grass, Annual Sedge, Perennial by number of species (N), or C = E C/N. Sedge, Herbaceous Vine, Woody Vine, Shrub, and Tree. Conservatism - The tendency of a taxon to be restricted to Tracking physiognomic classes can be an important natural areas. Similar to remnant dependency (Panzer et al. component of FQA, since it is theoretically possible for 1995). dramatic changes in community structure to occur without

Floristic Qualitv Index (FQI) - An index derived from changes in the FQI or C.

floristic inventor}' data and calculated by the following Rare Species - Plant species listed as threatened or formula from Swink and Wilhelm (1979, 1994): endangered by the Illinois Endangered Species Protection I = C (VN), in which: Board (Herkert 1991, 1994). C = coefficient of conservatism Species richness - Total number of native and adventive C = LC/N species. N = number of taxa. Wetness - Wetness classification is based on the National Guild Diversity - Guild diversity is measured from Wetland Categor}- for Region 3 of the Fish frequency distributions for species among traits such as and Wildlife Service (Reed 1988). Plants are designated as physiognomic classes, wetness ranks (see Wetness), or Obligate Wetland, Facultative Wetland, Facultative, conservatism ranks. These frequency data allow for Facultative Upland, and Upland. These classes are further graphical depictions of these guilds for comparison among ranked by " + " and "-" values for the three facultative sites and time periods (see Data Analysis in results section). classes, thereby providing greater resolution. These Illinois Natural Areas Inventory Grades - Definitions taken nominal classes have been sorted into ordinate values: from White 31): (1978, p. -5 = Obligate Wetland (OBL) Grade = Relatively stable or undisturbed communities. A -4 = Facultative Wetland + (FACW+) Example: old growth, ungrazed forest. -3 = Facuhative Wetland (FACW) Grade = Late successional or lightly disturbed B -2 = Facultative Wetland - (FACW-) communities. Example: old growth forest that was -1 = Facultative + (FAC+) selectively logged 5 years ago. = Facultative (FAC) Grade C = Mid-successional or moderately to heavily + 1 = Facultative- (FAC-) disturbed communities. Example: young to mature second- + 2 = Facultative Upland + (FACU+) gro-wth forest. + 3 = Facultative Upland (FACU) Grade = Early successional or severely disturbed D + 4 = Facultative Upland - (FACU-) communities. Example: severely grazed forest of any age. + 5 = Upland (UPL). Grade E = Very early successional or very severely Mean wetness is an average derived from all wetness disturbed communities. Example: cropland. (ordinate) values in a floristic inventory unit; it provides an Integrity, Ecological and Community - Integrity implies an index that characterizes the plant community in terms of unimpaired, complete condition. Ecological or community hydrological characteristics.

integrity refers to the degree to which self-correcting

EK1GESL4 November 1997 Floristic Qu.^LiTi' Assessment for \"egetation in Illinois

Acknowledgments

The authors gratefully acknowledge several reviewers who improved this manuscript with written comments

and discussion. These include Mark Schwartz (University of California at Davis), Geoff Levin (Illinois Natural History Survey [INHS]), Ken Robenson (INHS), John White (Ecological Services), John Ebinger (Eastern Illinois University), and Mary Kay Solecki (Illinois Nature Preserves Commission). Marlin Bowles

(The Monon Arboretum) reviewed the manuscript and provided alternative viewpoints. Jeff Brawn (INHS)

and Susan Aref (University of Illinois at Champaign-Urbana) provided statistical advice during preliminary phases of this paper. Louis Iverson (U.S. Forest Service), Ken Robenson, and Mark Schwartz offered

encouragement during early stages of the project that in many ways inspired the effort. We thank the Illinois Depanment of Transponation, Bureau of Design and Environment, for suppon and encouragement for the

development of this paper. Finally, thanks to the Illinois Native Plant Society for providing a special issue for the publication of this paper.

About the Authors

John Taft is a research scientist for the Illinois Natural History Survey, where he performs botanical and

ecological evaluations on Illinois Department of Transportation project areas throughout the state. He also conducts independent research in plant community ecology. John earned an M.S. in botany from Southern

Illinois University and is a Ph.D. candidate at the University of Illinois in the field of natural resources and environmental sciences.

Gerould Wilhelm has a Ph.D. in botany from Southern Illinois University where he worked under Dr. Robert Mohlenbrock. Much of his botanical research has been centered in the Chicago region, where he coauthored with Floyd Swink the 3rd and 4th editions of Plants ofthe Chicago Region; he has also compiled a lichen flora for the region. Gerry works as the principal environmental scientist for Conservation Design

Forum, Inc. and as a research scientist for Conservation Research Institute.

Douglas Ladd is the director of science and stewardship for the Missouri Chapter of the Nature Conservancy.

He received his M.S. in botany from Southern Illinois University, where he later worked with Dr. Robert Mohlenbrock on the Distribution ofIllinois Vascular Plants. His recent publications mclude

Wildflowers and Checklist and Bibliography ofMissouri Lichens. One of his special interests is .

Linda Masters is the director of Conservation Research Institute. She works as a restoration ecologist

professionally and devotes volunteer efforts to natural land preservation and stewardship as well. She has

developed the application computer programs for floristic quality assessment in the Chicago region and is

continuing to adapt the methodology and programming for use in other geographical regions. Linda wrote the chapter "Vegetation Monitoring" in the 1997 publication The Tallgrass Restoration Handbook and will

soon be receiving an M.S. in ecology from the University of Illinois at Chicago.

ERIGENL4 November 1997 Floristic Qu.\Li'n' Assessment for Vegetation in Illinois

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Erigenia November 1997 Floristic Qu.\LnT Assessment for Vegetation in Illinois

APPENDIX: Vegetation of Illinois Database

The following is a listing of selected acronyms serve as easUy rendered extraction tags for the taxa, sorted alphabetically by and then by species, plants in the data base. for use in the apphcation of Floristic Quahty Following the acronym is the assigned C value

Assessment in Illinois. Native species are rendered in a (coefficient of conservatism) for native species or by an standard type face, while introduced or adventive species asterisk for non-native species. After the colloquial name

are shown in ALL CAPS; each is followed by a single is the coefficient of wetness, followed by its

colloquial name. This listing is not to be construed as an corresponding National Wetland Category. The authoritative treatise on the flora of Illinois, nor was categories were assigned based on observations of their there any attempt to justify the Latin name as behavior throughout "Region 3" of the U.S. Fish & nomenclaturally legitimate. Indeed, for taxonomic Wildhfe Service. Obligate wetland species (OBL, - 5) concept and nomenclature, we have approximated have 99% probabihty of occurring in wetlands,

Mohlenbrock (1986), wherein authorities for most of the facultative/wet species (FACW, -3) a 67%-99% names may be found. probability, facultative species (FAC, 0) a 34%-66%

Each species is preceded by a six-letter acronym, based probabihty, facultative/upland species (FACU, 3) a

upon the first three letters of the genus followed by the l%-33% probabihty, and upland species (UPL, 5) have

first three letters of the species, or by the first two letters less than a 1% probabihty of occurring in wetlands.

of the species and the first letter of a subspecific taxon Each species has been designated with a physiognomic (e.g./l^«tilonJ/7eophrasti = ABUTHE;/lcerrMbrumvar. characteristic, using commonly apphed terms such as iirummondii (ACERUD). Where ambiguity results, tree, shrub, forb, vine, grass, sedge, and cryptogam. The

such as in Polygonum frydiopiper (POLHYR) and forbs, grasses, and sedges are preceded by modifiers such Polygon-am. /7Wropiperoides (POLHYS) a nonintuitive as A (annual), B (biennial), and P (perennial). These are acronym has been created. Use of such acronyms makes followed by a family name, following the delineation in

field notes go much faster, and the Mohlenbrock (1986).

ERIGENL4 November 1997 1

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ERIGENLA. is a peer-reviewed journal published by the and an indication of the top of the figure should be Illinois Native Plant Society. The journal publishes lightly penciled on the back of every figure. Figure research papers, reviews, essays, and technical reports legends should be typed double-spaced on a separate pertaining to the biota of Illinois and adjacent states. sheet.

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All manuscripts and communications should be sent to Headings and hterature citations in the text and in the the editor, Gerould S. Wilhelm, Conservation Literature Cited section should be in the formal Research Institute, 324 N. York Road, Elmhurst, IL outlined below: 6C126. Three copies of the manuscript should be MAIN HEADING submitted. Authors should retain a copy of all Subheading on Own Line material. The title page should state the affiliation and Sub-subheading at beginning of line. The text continues the complete addresses of all the authors; telephone Sub-sub-subheading indented and at beginning of

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Common names, if used, should be referenced to a habitats on the Yucatan Peninsula. Pages 273-286 scientific name. Thereafter, scientific names are in J. M. Hagan III and D. W. Johnson, eds. recommended, but either may be used if done so Ecology and conservation of neotropical migrant consistently. landbirds. Smithsonian Institution Press, Washington, D.C. Tables and Illustrations

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Erigenia 15 November 1997

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