Format of Page

In general, each map shows county records, with family, genus and species name to upper left (plus selected synonyms or alternative names informally indicated in parentheses or brackets). Below each map, the full binomial name is displayed with author, and the right there are codes for alien status (ALI), habitat (HAB) and abundance (ABU). Then in the lower block of text are brief notes on the species, with special attention to problems with taxonomic recognition or identification.

Format for family, genus and species names at upper left

If a species name lacks a variety designation, this means that only the typical variety is recognized in the state; or in a few cases, as explained in footnotes, varieties may be recognizable but are combined in these maps.

<  >      Indicates divisions of families or genera in general use: subfamilies, tribes, subgenera, sections and series. These are only informal listings.

(   )       Indicates synonym for epithet; hybrid name; or informal name (“ined.”).

{  }      Indicates brief note on name or suggested name; the + symbol indicates inclusion of a variety that may be segregated with further work.

"  "       Indicates wrong or uncertain application of an earlier name, not a proven synonym.

x       Indicates hybrid between the two species either side of this symbol.

X       Precedes the formal name for the hybrid, if this epithet is available.

The following symbols are also applied.

*          Indicates that this different name was used by Jones (2005).

**        Indicates that a different name was used by Jones (2005), but the unsettled situation in these old “Liliaceae” is too complex to show details.

[  ]       Indicates a broader family or genus concept, generally derived from old fashioned taxonomy.

Taxa excluded from mapping

In some species without consistently definitive records, their county records are not displayed and these species are indicated with the following categories of exclusion.

C	Well-documented but records may just come from cultivated plants that may persist for some time into wilder contexts, but perhaps not indefinitely; also includes a few weeds largely restricted to intensively cultivated sites (e.g. flower beds and greenhouses).
R	Reported and may occur in the wild, but records are uncertain or unverified, collections are lost or incomplete, or there are other problems with the data; explanations are provided in the footnotes.
T	Reported and may occur in the wild, but taxonomic distinction from close relatives remains unclear, and in some cases only intermediate plants may occur; in some cases, data for these taxa are mapped with their relatives, as explained in footnotes.
W	Well-documented but records may just come from rare waifs, defined as briefly established plants that have dispersed into the state through some long-distance mechanism (including occasional escapes from cultivation) but not truly perpetuating themselves in the wild.
These are not mutually exclusive categories, especially C versus W, and R versus T, but what appears to be the most applicable code is provided in each case. Note also that hybrids are generally not mapped or listed, unless established in populations independently of parent species, but their occurrence is indicated in footnotes of the Index.

Notes on distribution, ecology and taxonomy

These notes deal with details of some individual records, taxonomic problems, distinction of varieties or other segregates, occurence of hybrids, characters for identification, uncertainty in alien versus native status, priorities for further research, comments on biogeography or ecology, and other miscellaneous information of interest. Individual counties are referred to by four-letter abbreviations (Table 3, Figure 4).

The following abbreviations are used for some of the most frequently used bibliographic references.

APG    Angiosperm Phylogeny Group (2003); Stevens (2008) and his cited literature.

B         Braun (1943); including her references to other sources; note that her collections are mostly at US, but some are at GH, NY and perhaps elsewhere.

BA      Browne & Athey (1992)

BT       Beal & Thieret (1986)

Ch       Chester et al. (1993, 1997, 2009); plus updates at tenn.bio.utk.edu

Cr        Gleason & Cronquist (1991); plus Cronquist (1980) for Asteraceae

Co       Cooperrider (1995; et al. 2001; in prep.); including their major references

CW      Clark & Weckman (2008)

D         Deam (1940)

F          Fernald (1950)

FNA    Flora of North America (1993-2007; see vols. listed in bibliography)

Gm      Garman (1900, 1902, 1913, 1914)

Gl        Gleason (1952)

HFG    Harmon, Ford-Werntz & Grafton (2006); see also SC

HW+   Harvill et al. (1986); with updates of T. Wieboldt et al. (www.biol.vt.edu)

J           Jones (2005)

K         Kartesz (1999, 2011).

M         Medley (1993)

ML      Mohlenbrock & Ladd (1978) and Mohlenbrock (1988)

NP       Kentucky State Nature Commission Database (2001)

NS       NatureServe Explorer (2010); continuing, at www.natureserve.org

PL       PLANTS website of U.S.D.A. (2010); continuing, at www.plants.usda.gov

Pr         Price (1893); plus handwritten notes in copy at Missouri Botanical Garden

RAB    Radford, Ahles & Bell (1964)

SE       Southeast Exotic Pest Plants Council (2011); continuing, at www.eddmaps.org

Sm       Small (1933)

St         Steyermark (1963); see also Y

SC       Strausbaugh & Core (1978)

W        Weakley (2012); see also Atlas at www.herbarium.unc.edu

Y         Yatskievych (1999, 2006; continuing)

The following abbreviations are used for references to herbaria (Holmgen et al. 1990). These references are generally placed in parentheses after the abbreviations for counties. Note that, although locations of specimens are often reported in the literature, several collections have not been relocated, or may be temporarily mislaid, or lost. In the text below "check" implies that further search is warranted.

APSU Austin Peay State University (Clarksville, Tennessee)

BEREA Berea College (Berea, Kentucky)

CINC  University of Cincinnati (Cincinnati, Ohio)

DHL    Davies Herbarium, University of Louisville (Kentucky)

      Note: this herbarium was transferred to WKY in 2008.

DOV   Delaware State University (Dover, Delaware)

EKU    Eastern Kentucky University (Richmond, Kentucky)

F[I]      Field Museum (Chicago, Illinois)

GH      Gray Herbarium, Harvard University (Cambridge, Massachusetts)

KNK   Northern Kentucky University (Alexandria, Kentucky); now renamed “The John W. Thieret Herbarium”

KY      University of Kentucky (Lexington, Kentucky) [Note that important historical collections and county records that have been housed separately in the School of Agriculture are transferred to the main herbarium, or should be.]

MDKY Morehead State University (Morehead, Kentucky)

MEM   University of Memphis (Memphis, Tennessee); now renamed “The Raymond Athey Herbarium”

MICH University of Michigan (Ann Arbor, Michigan)

MO      Missouri Botanical Garden (St. Louis, Missouri)

MUR   Murray State University (Murray, Kentucky)

NCU   University of North Carolina (Chapel Hill, North Carolina)

NY      New York Botanical Garden (Bronx, New York)

PH       Philadelphia Academy of Science (Philadelphia, Pennsylvania)

PUR    Purdue University (Purdue, Indiana)

SIU     Southern Illinois University (Carbondale, Illinois)

TENN University of Tennessee (Knoxville, Tennessee)

UC      Jepson Herbarium, University of California (Berkeley, California)

US       U.S. National Herbarium, Smithsonian Institution (Washington, DC)

WKY  Western Kentucky University (Bowling Green, Kentucky)

 

Other miscellaneous abbreviations are as follows.

2n        chromosome number of sporophyte

(Goldblatt & Johnson 2006, 2008)

aff.      affine (Latin for ‘with affinity to’)

auct.    auctores (Latin indicating ‘in the sense of some authors but not the original one’)

c.         central

ca.        circa (= about)

cf.        confere (Latin for ‘to be compared with’)

Co.      County

coll(s). collection(s)

comb.  combination

dbh      diameter at breast height (1.45 m)

det.      determined

e.         eastern

f.          filius (Latin for ‘son’)

flw       flower

l/w       length/width ratio

Mt.      Mountain

n.         northern

p.p.      pro parte (Latin for ‘in part’)

pro sp. pro species (Latin implying ‘an incipient species from well-established hybrids’)

Rv.      River

s.          southern

ssp.      subspecies

var.      variety

w.        western

´          indicates hybrid between the two species either side of this symbol

X         precedes the formal name for the hybrid, if this epithet is available

Abbreviations for the states of U.S.A and the provinces of Canada follow Fernald (1950) and FNA (1993-2007). Abbreviations for months are Jan, Feb, Mar, etc.

Codes for alien origin (ALI), typical habitat (HAB), abundance and change (ABU).  Only mapped species with certain presence in the state are provided with a full set of codes.

ALI     Alien Origin. Species are certainly native to Kentucky unless coded as follows; codes in lower case apply to species with somewhat uncertain status.

AF       Native mostly in Africa, south of the Mediterranean region.

AS       Native mostly in east Asia, from India to Japan.

AU      Native mostly in Australasia.

e          Native mostly to the east in North America; adventive or mixed in Kentucky.

E          Native to the east in North America.

EU       Native mostly in Eurasia—Europe, western Asia and Mediterranean region.

m         Native and alien plants may be mixed; range generally including Eurasia.

n          Native mostly to the north in North America; adventive or mixed in Kentucky.

N         Native to the north in North America; circumboreal in some cases.

s           Native mostly to the south in North America; adventive or mixed in Kentucky.

S          Native to the south in North and Central America; cosmopolitan in some cases.

SA       Native mostly in South America, south of Mexico and Central America.

w         Native mostly to the west in North America; a          dventive or mixed in Kentucky

W        Native to the west in North America, and in some cases elsewhere.

 

HAB   Typical Habitat. These codes are in four sequential groups: (i) 1-12 for broad overall habitat classes; (ii) miscellaneous microhabitats; (iii) A-E for acid-versus-basic soil; (iv) 1-6 for sun-versus-shade. This classification is an attempt to bridge gaps between language that has been traditionally used by botanists (e.g. Braun 1950), ecologists (e.g. Ky. State Nature Preserves website) or foresters (e.g. Eyre 1980), and the more complex phytosociological system that is being developed by NatureServe (see their website).

 

(i) Habitat classes. Numbers 1-12 indicate the classes of natural habitat (or native vegetation) where each species generally occurs, with the most typical class listed first. These broad intergrading classes have been developed from a general model of habitat gradients in the state (e.g. Campbell 1987, Campbell & Grubbs 1992), as diagrammed in Figure 1. There is an important third dimension to habitat that is not shown in Figure 1; this is the pH-related gradient that is indicated by the coding (A-E) of the fourth column, and diagrammed for upland forest types in Figure 2.

  

   For species that are frequently associated with more alien vegetation in artificially modified habitats, codes for natural habitats (1-12) are preceded by the following lettered codes. Only the most common type of artificial habitat is usually indicated, pending refinement of this system in future editions. Upper case codes indicate species that usually prosper in vegetation dominated by alien species. Lower case codes indicate that aliens are less prevalent but estimated to comprise at least 10% of typical associates.

F          Fields, farmland or suburban land in general, including fencerows and older thickets; this is the default code if G, H, R or S are not prevalent, and it generally indicates artificial modifications of habitat classes 7, 8, 9, 10 (especially) or 12.

G         Grazed/hayed areas, powerlines or other sites treated no more than once per year.

H         Horticultural contexts, cropland and similar sites with annually exposed soil.

R         Roadsides, lawns and similar sites cut or sprayed more

            than once per year.

S          Short trampled or mowed vegetation; paths, sidewalks, ball-fields, parking-lots, etc.

Especially on roadsides (R), deeper research may reveal further local selection from seasonal uses of herbicide, both intentional and from agricultural drift. In regions where small patches of native woodland or grassland are largely surrounded by cropped fields, there can be remarkable absences of even common native perennials. For example, recent drives through lowlands of the Shawnee Hills suggest general absences of early flowering species along roads (Hieracium, Houstonia, Krigia, Monarda, Phlox). Instead abundant weedy annuals such as Erigeron annuus and Packera glabella tend to spread from fields into roadsides, fencerows and woodland edges. 

The 12 classes of natural habitats are outlined below in terms of hydrology and overall development of vegetation, from open (stressed or disturbed) to closed (and more or less mesic) conditions. Ecologists have employed several descriptive adjectives for habitats based on Greek roots.  These words do not have broad consistency in scientific usage, but for Kentucky they can be defined in simple terms as follows.

Aquatic            With soil submerged for most of the year; usually open except with cypress

Dystrophic      With soil poor in nutrients, especially nitrogen and phosphorous

Eutrophic        With soil rich in nutrients, especially nitrogen and phosphorous

Hydric             With soil wet enough to be anaerobic for much of the year

Hydroxeric      With soil usually wet in winter, dry in summer; opposite of mesic

Mesic               With neither hydric nor xeric tendencies; oak usually local to absent

Pyric                With frequent fires greatly influencing the vegetation

Rheic               With forceful floods greatly influencing the vegetation

Ruderal           Associated with farmland on more or less mesic eutrophic sites

Seral                Changing to more mature or mesic forest unless redisturbed

Subhydric        Intermediate between hydric and mesic; often grading into hydric oak

Submesic         Somewhat mesic but modified in past/present by stresses/disturbances

Subxeric          Intermediate between xeric and mesic; oak potentially abundant

Xeric                With soil dry enough to maintain much red cedar or pine or rocky glades

Xerohydric      With soil usually dry in summer, wet in winter; grades into hydroxeric.

More popular terms closer to traditional usage in floristic manuals are provided in quotation marks below.

As outlined below, Classes 2 and 3 contain hydrophytic vegetation mostly composed of ‘obligate’ wetland species (Reed 1988, Jones 2005). Classes 1 and 9 contain mixtures or zonations of species from ‘obligate’ to ‘facultative wetland’ to ‘facultative’ species. Classes 4 and 6 contain mostly ‘facultative wetland’ or ‘facultative’ species. Classes 7, 8 and 10 are generally not at all hydric, but can include swales or damp flats with patches of ‘facultative wetland’ or ‘facultative’ species.

Class 12 contains the most xerophytic vegetation. Classes 11 and 10 usually contain moderately xerophytic vegetation. Classes 1, 7, 8 and 9 contain various mixtures or zonations that can include some moderately xerophytic vegetation. Classes 2, 3, 4, 5 and 6 contain virtually no xerophytic vegetation, except for included microhabitats such as seasonally exposed shorelines, occasional rocky ledges, and epiphytic situations.

1          Rivers and stream corridors, including shrubby/grassy streambanks. This class of habitat is defined to include the whole non-forested zone along flowing watercourses. Aquatic plants (Podostemon, Potamogeton, Vallisneria) can occur within relatively unpolluted free-flowing streams, and water-willow (Justicia americana) is often abundant in riffles. The banks are temporarily flooded, but often violently so, which tends to prevent complete tree cover. Also, seasonal drought can be severe in sections with coarse sandy, gravelly or rocky substrate, which hinders forest further. Various species of willows (Salix) are typically abundant at lower levels, often with much silky dogwood (Cornus obliqua). Other examples of characteristic shrubs, locally, include alder (Alnus serrulata) in more swampy transitions, white azalea (Rhododendron arborescens) on Appalachian sandstone boulders, and indigo-bush (Amorpha fruticosa) along larger rivers. Some streams have complex zonations, from shoals with partly submerged plants, to low lagoons with sedges (e.g., Carex emoryi along larger rivers), to exposed substrates with abundant annuals (e.g., Bidens, Eragrostis, Persicaria, Xanthium), to more stable shrubby levees, to higher rocky banks and ledges with perennial grasses (especially Panicum virgatum or Andropogon gerardii) and characteristic forbs (e.g., Baptisia australis, Physostegia virginiana, Solidago spp., Zizia aurea). Also, wood oats (Chasmanthium latifolium), wild ryes (Elymus) and other species are characteristic of more shady transitions to adjacent riparian forest. Many rare, uncommon or disjunct species occur in some variants of this class. In general, the vegetation along banks of larger streams is moderately secure from human interferences, but some variants have been much reduced by impoundment. Also, aquatic vegetation is probably much reduced in most larger streams and rivers due to pollution and impoundment.

2          Lakes and ponds, with associated shrubby/grassy marshes; including artificial impoundments. This class is defined to include the whole non-forested zone around relatively stagnant water-bodies. The banks are seasonally or semipermanently flooded, and rarely rocky (except for some artificial impoundments). There are often complex zonations with increasing water depth or flooding frequency. Buttonbush (Cephalanthus occidentalis) is often dominant in the woody fringe around these sites; other shrubby species include alder (Alnus serrulata), black willow (Salix nigra), swamp dogwood (Cornus foemina), swamp privet (Forestiera acuminata), and water elm (Planera aquatica). Wetter zones can include locally abundant cat-tails (Typha), grasses (especially Phragmites australis, Phalaris arundinacea, Leersia oryzoides), sedges and their allies (Carex, Cyperus, Scirpus), knotweeds (especially Persicaria hydropiperoides), swamp-loosestrife (Decodon verticillatus), and others, grading into duckweeds (Lemnaceae), pondweeds (Naijadaceae, Potamogetonaceae), water-lilies (Nymphaeaceae) and other floating or submerged aquatic vegetation in open water. Mudflats and other exposed shorelines during the drier seasons can become colonized by many annual species (e.g., Bidens, Cyperus, Fimbristylis, Ipomaea, Iva, Persicaria, Xanthium). Although there are many artificial or semi-natural areas with this kind of vegetation, there are few good natural examples, except in the lowlands of western regions. Many variants probably existed before settlement, and several may have virtually disappeared in some regions, e.g., smaller oxbows, beaver-ponds, seasonally ponded swales, and associated boggy openings.

3          Cypress/tupelo or other deep swamp forest, with associated open woodland. These seasonally or semi-permanently flooded sites are concentrated in southwestern regions. Bald-cypress (Taxodium distichum) and water tupelo (Nyssa aquatica) are locally dominant; other characteristic trees may include swamp red maple (Acer rubrum var. trilobum), green ash (Fraxinus pennsylvanica), pumpkin ash (F. tomentosa), sweetgum (Liquidambar styraciflua), water elm (Planera aquatica), swamp privet (Forestiera acuminata), water locust (Gleditsia aquatica), swamp cottonwood (Populus heterophylla), water hickory (Carya aquatica) and overcup oak (Quercus lyrata). Further north and east suitable habitat does exist, but the typical southern forest types are mostly replaced by open woods with trees less tolerant of flooding (transitional between Classes 6 and 2), or by more open shrubby/grassy vegetation (Class 2). More disturbed or peripheral sites have often become locally dominated by clonally spreading species like the common reed (Phragmitis australis), cat-tails (Typha), or swamp-loosestrife (Decodon verticillatus). Although there are still some extensive areas in fair condition, several of the characteristic southern species are rare in the state. More depauperate outlying variants are highly threatened.

4          Riparian (“streamside”) forest. These sites have temporary flooding, scouring and siltation that significantly influence the canopy composition, but with rapid recovery after disturbance. Typical trees are able to resucker rapidly after damage, and have root systems that can form highly branched ‘wads’ on unstable banks (E. Hartowicz, pers. comm., especially asterisked species*). These species are mostly boxelder (Acer negundo*), silver maple (A. saccharinum*), river birch (Betula nigra*), sycamore (Platanus occidentalis*) and cottonwood (Populus deltoides). Other common trees, including swampy transitions, are sweetgum (Liquidambar styraciflua), green ash (Fraxinus pennsylvanica), black willow (Salix nigra) and white elm (Ulmus americana). These forests are still widespread in the state, though often confined to narrow strips.

5          Mesic forest (“moist woods”). This class can occur on well-drained, temporarily flooded alluvial terraces, on various types of toe slope and upland swale, on some relatively undisturbed uplands with residual soils, and on steeper colluvial slopes, especially slopes with NE-facing aspects. Mesic sites have little or no hydric, xeric or pyric influence. The forest is fairly continuous, except for streamsides and temporary gaps after trees die. Among the most typical trees are sugar maples (Acer saccharum and its relatives), basswoods (Tilia), buckeyes (Aesculus), bitternut hickory (Carya cordiformis), beech (Fagus grandifolia), and tulip poplar (Liriodendron tulipifera). More locally, on acid soils, there are characteristic magnolias (Magnolia), birches (especially Betula lenta) and hemlock (Tsuga canadensis). More stressed or disturbed transitions can also have locally abundant white ash (Fraxinus americana), slippery elm (Ulmus rubra), northern red oak (Quercus rubra), and white pine (Pinus strobus). Also, chestnut (Castanea dentata) was formerly frequent in subxeric transitions on sandy soils. In the modern landscape, mesic forests has become increasing restricted to steeper slopes. It is still widespread in hilly regions, but variants on fertile floodplains and on calcareous uplands have become very rare or virtually eradicated.

6          Subhydric forest (“damp” or “wet woods”) and associated open woodland. This class occurs on lower floodplain terraces or swampy upland swales and seeps, with temporary to seasonal flooding or saturation, and high water tables. Although soils are generally damp or wet, there can be occasional droughts and even fires. There can be scattered small openings, especially along streams. Common trees include swamp red maple (Acer rubrum var. trilobum), blackgum (Nyssa sylvatica), sweetgum (Liquidambar styraciflua), sugarberry (Celtis laevigata), green ash (Fraxinus pennsylvanica), white elm (Ulmus americana); and locally hickories—notably shellbark (Carya laciniosa), and pecan (C. illinoinensis), and oaks—notably swamp chestnut oak (Quercus michauxii), bur oak (Q. macrocarpa), cherrybark oak (Q. pagoda) and shumard oak (Q. shumardii). There is much intergradation with the more oak-dominated forest of Class 9. Sedges, grasses, cane and other shrubs are often abundant in the understory, especially in more open woods close to streams and seeps. These forests are widespread in the state, but good examples are mostly small and threatened by human disturbances. Variants on more fertile soils have been largely cleared or intensively grazed. Variants on less fertile soils, especially small acid seeps, have often been degraded by pollution of surface or ground water.

 

7          Submesic forest (average “upland woods”) and associated open woodland. This is a widespread, somewhat miscellaneous class that is often a broad transition between mesic forest (Class 5) and more frequently disturbed thickets (Class 8) or open woodland (Class 10). It can occur on a wide range of topographic sites, but is most extensive on gentle slopes and flats with residual or alluvial soils that have some dry or damp season. These sites include moister zones in rolling plains, seasonally damp swales on uplands, and terraces with temporary flooding. Though somewhat mesic, the forests experience stresses from occasional dry or wet conditions, and they have often been disturbed by burning, grazing or farming. The highly varied tree composition can include locally abundant red maple (Acer rubrum), blackgum (Nyssa sylvatica), holly (Ilex opaca), tulip poplar (Liriodendron tuplipifera), hackberry (Celtis occidentalis), black walnut (Juglans nigra), buckeyes (especially Aesculus glabra), mulberry (Morus rubra), coffee tree (Gymnocladus dioicus), and various ashes, elms, hickories and oaks. Sugar maple (Acer saccharum) and beech (Fagus grandifolia) are locally abundant in transitions to more mesic forest. On more fertile soils, the forest can be especially distinctive, with locally dominant walnut, hackberry or buckeye; with thickets of cane, pawpaw, coralberry, other shrubby species and common woody vines; and with dense ground vegetation dominated by tall herbs (e.g. Ageratina altissima) and cool-season grasses (e.g. Elymus spp.),  Much disturbed forest belongs in this class, but probably few areas are similar to presettlement conditions, because of general conversion to farmland, frequent wood-cutting, and overgrazing in the remnants.

8          Deciduous seral thickets, maintained by repeated disturbance. Ecological distinction of this varied habitat class, typified by several thorny or root-sprouting species, is often bypassed by biologists. It occurs on a wide range of soils, but usually not hydric or xeric enough for permanent openings without disturbance. Much is in forest-edges, brushy grasslands, and old fields with frequent fire or grazing. While many woody species can locally prosper, some of the most characteristic larger trees include persimmon (Diospyros virginiana), sassafras (Sassafras albidum), honey locust (Gleditsia triacanthos), black locust (Robinia pseudoacacia), black cherry (Prunus serotina) and osage orange (Maclura pomifera). Characteristic smaller trees and shrubs include devil’s walking-stick (Aralia spinosa), cane (Arundinaria gigantea), pawpaw (Asimina triloba), redbud (Cercis canadensis), dogwoods (Cornus), black haws (Viburnum), hawthorns (Crataegus), crabapples (Malus), plums (Prunus), sumacs (Rhus), roses (Rosa), dwarf willows (Salix), coralberry (Symphoricarpos orbiculatus) and prickly-ash (Zanthoxylum americanum). As a class, such vegetation is extensive in today's disturbed landscape, often beginning in old-field succession with abundant briars (Rubus). However, it is probably much changed from presettlement conditions, and it is likely that some variants have now virtually disappeared. For example, canebrakes and plum thickets were probably widespread at grassland margins, along animal trails, and around regular human (Native American) encampments, but have now become highly reduced in the modern landscape. Also, pines and cedars have tended to displace many of these species in succession to submesic forest (Class 7) or other classes, especially on drier sites where regular burning and browsing has declined.

9          Hydric-tending flats with oak woodland or grassland (“swampy woods and openings”); including artificial variants along shores and other seasonally wet zones around impoundments.  This class occurs on seasonally to temporarily flooded or saturated soils, but in some cases there is also frequent drought stress and even occasional fire in late summer or fall. Ungulates probably had local influence before settlement, especially near seeps, licks and wallows. Oaks are common in wooded remnants—including characteristic swamp white (Q. bicolor), overcup (Q. lyrata), pin (Q. palustris), and willow oak (Q. phellos), but there is also much red maple (Acer rubrum var. trilobum), blackgum (Nyssa sylvatica), sweetgum (Liquidambar styraciflua), and green ash (Fraxinus pennsylvanica), especially in transitions to subhydric forest (Class 6). Openings with native vegetation include rushes (Juncus), sedges (Carex), beak-sedges (Rhynchospora), bulrushes (Scirpus), umbrella-sedges (Cyperus), spike-rushes (Eleocharis), and many grasses. Wetter openings often grade into marshy vegetation transitional to Class 2 or 3. Ephemeral ponds of various types can be included within the definition of this class. >Although small remnants of such vegetation are widely scattered in much of the state, the few good extensive examples are concentrated in western regions. Most land has been converted to farmland, especially with artificial drainage after 1950. More open, grassy variants with a history of burning—and perhaps licking, stamping and wallowing by megafauna—have virtually all disappeared.

10        Xeric-tending plains with oak woodland or grassland (“open oak woods and barrens”); now mostly converted to farmland.Native vegetation of this class was formerly widespread on gentle slopes, rolling plains and upland flats with a history of much burning and grazing. Although soils are deep and often somewhat mesic, they are subject to occasional intense droughts. The land is not typically flooded or saturated, but some soils are relatively impermeable or poorly drained in winter or spring.  Characteristic trees in wooded areas include oaks—especially blackjack (Q. marilandica), post (Q. stellata), southern red (Q. falcata), shingle (Q. imbricaria), shumard (Q. shumardii), and bur oak (Q. macrocarpa), hickories (especially Carya laciniosa, C. ovata and C. tomentosa), locally ashes (especially Fraxinus americana) and elms (especially Ulmus alata and U. americana). Pines (especially Pinus echinata in more natural areas of Appalachian regions) and red cedars (Juniperus virginiana) can be locally abundant, depending on the history of disturbance and succession, and then distinction from Class 12 can be more arbitrary. Many native grasses and wildflowers are typical of the more open variants. This vegetation has been virtually all converted to farmland, and the woodland remnants, without burning or browsing, tend to become denser and shadier in various ways. Without disturbance, there can be increases of pine or cedar (seral extensions of Class 12), development of shrubby thickets (Class 8), gradual invasion of more mesic trees such as maples (Class 7), or various mixtures of these compoments. Much farmland can be considered artificially analogous to this class, though with much more soil disturbance and with a more weedy, ruderal component in the vegetation. The clearance of trees, erosion of topsoil, soil-compaction, and more flashy hydrology in farmland has allowed expansion of this broadly defined habitat class onto formerly more mesic sites. It may be reasonable to segregate a distinct class for the more ruderal or pastoral vegetation that is typical of farmland.

11        Subxeric oak forests (“dry or rocky woods”).  This class is on moderate to steep slopes and ridges, with well-drained soils that are generally rocky with outcrops or colluvium. Disturbance from burning or browsing may have occurred before settlement but less intensively than in Class 10, and not enough to thin out the tree canopy and maintain grassy openings. Typical trees included chestnut (Castanea dentata, before the blight), chestnut oak (Quercus montana), scarlet oak (Q. coccinea), white oak (Q. alba), black oak (Q. velutina), southern red oak (Q. falcata), chinquapin oak (Q. muhlenbergii), shumard oak (Q. shumardii), pignut hickory (Carya glabra), mockernut hickory (C. tomentosa), shagbark hickory (C. ovata), white ash (Fraxinus americana), and blue ash (F. quadrangulata). Without disturbance, the understory often shifts to include more mesic or submesic species such as red maple (Acer rubrum), sugar maple (A. saccharum), blackgum (Nyssa sylvatica), red elm (Ulmus rubra), and white pine (Pinus strobus). This vegetation is still extensive, because much of the land is not suitable for farming, but there are very few examples of old-growth.

12        Xeric coniferous forest/glades and transitions to oak (“dry pine- or red cedar- oak woods and glades”); including artificial variants in roadcuts, quarries, mines, etc.These habitats typically occur on thin rocky or eroded soils where droughts maintain relatively open conditions and reduce the rate of succession to oaks or other forest trees. Such vegetation was formerly concentrated on or near rocky slopes (especially with SW-facing aspect), narrow ridges, clifftops, flatrocks, other outcrops, and eroded areas. Included also are various kinds of transition to oak or successional forests on deeper soils, often influenced by disturbance history as well as dry soils. There has usually been exposure to fires but fuels are often thin and interrupted by outcrops. Ungulate use was presumably varied before settlement, ranging from insignificant along clifftops to highly intense in eroded variants around mineral licks.With changes in disturbance-regime, red cedar (Juniperus virginiana) and scrub pine (Pinus virginiana) have now become much more extensive, especially in less xeric sites with young woods and thickets transitional to Classes 10, 8 and 7, and then distinction from those classes is often arbitrary. Other locally abundant trees include pitch pine (P. rigida), short-leaf pine (P. echinata), and various oaks, hickories, ashes and elms. More xeric variants have stable grassy openings, and these have generally survived fairly well since settlement. However, most subxeric transitions have become highly modified or reduced in extent, due to clearance for farmland, reduction in fire-frequency, and other environmental changes.

(ii) Microhabitat: association with rock, water, bare ground, or unusual substrates.

+          Loosely associated with rock outcrops or boulders

--          Partially associated with flat rock outcrops

==        Strongly associated with flat rock outcrops

\           Partially associated with cliff tops or ledges

\\          Strongly associated with cliff tops or ledges

           

|           Partially associated with vertical rock faces

||           Strongly associated with vertical rock faces

/           Partially associated with cliff bases or rockhouses

//          Strongly associated with cliff bases or rockhouses

~|         Associated with seeping, dripping or sprayed rock faces

~          Partially associated with water in growing season; often on receding shoreline

~~        Strongly associated with water in growing season; floating or submerged

::          Partially associated with bared ground: often due to establishment from seed

::::        Strongly associated with bared ground: eroded, trampled, burned, plowed, flooded

^          Facultative epiphyte on trees

^^        Obligate epiphyte on trees

 

(iii) Association with acidic infertile soils versus basic fertile soils. Typical position along this gradient is assigned from general knowledge in Kentucky and interpretation of regional floras. Associations of species with ecological regions of the state, which can be defined largely using the U.S. Dept. of Agriculture’s mapping of soils (Figure 3), form important provisional clues for this classification. The gradient in forests of east-central North America has been revealed by ordination studies (e.g. Campbell 1987; see Figure 2), but much more detailed work is needed to examine relationships of nitrogen, phosphorous, aluminum, other minerals, and associated factors. The parallel gradient in Europe has been well documented by Ellenberg (1988; and previous German editions), leading to many applications of botanical indices for pH and nutrient levels (e.g., Wittig & Neite 1986, Meerts 1997, Ertsen et al. 1998, Wamelink et al. 2002, Chytrý et al. 2003, Ewald 2003, Thimonier et al. 2006). However, applications of this type in North America are just beginning (e.g., Peet et al. 2003).

While there is much general knowledge of the processes that can correlate vegetation types with soil types in North America (e.g. Fisher & Binkley 2000), there have been few studies in which the relevant processes of soil chemistry and ecophysiology have been investigated. It is generally accepted that soils with pH 6-7 tend to have high natural levels of available nitrogen and phosphorous, as in the Bluegrass region, but raw data have not been analyzed, and this may only be a loose correlation. The highest fertility, in terms of phosphorous and nitrogen, may often occur on slightly acid soils, with pH of about 6; at pH 7 and above, phosphates become much less available to plants. The form of nitrogen—ammonium at lower pH and nitrate at higher pH—may also be a significant factor. And at low pH, aluminum and manganese can become more soluble and toxic to some plants. As in Europe, it will be important eventually to develop indices for nitrogen and phosphorous availability that are independant of indices for overall base status or pH.

 

Guidelines for this initial coding are as follows.

 

A         Association with strongly acid soils (ca. pH 4-5) and low overall fertility; typical upland oaks include Quercus montana and Q. coccinea; Castanea dentata was formerly frequent; Ericaceae are common. This is indicated by concentration on relatively acid shales and sandstones in the Knobs and Appalachian regions, coupled with virtual absence in the Bluegrass region or other calcareous regions.

B         Transitional or uncertain assignment between A and B.

C         Association with medium acid soils (ca. pH 5-6) and medium overall fertility; typical upland oaks include Q. alba, Q. velutina, Q. stellata and Q. falcata.

This is indicated by widely scattered distribution over the state, including parts of the Bluegrass, or other calcareous regions, as well as the Knobs and Appalachian hills.

D         Transitional or uncertain assignment between C and E; widespread species that are common on farmland or alluvial soils with relatively high fertility (especially in N and P) are generally assigned here.

E          Association with weakly acid to neutral soils (ca. pH 6-7) and medium to high overall fertility, especially in bases (Ca, Mg, K); typical upland oaks include Q. muhlenbergii, Q. shumardii, Q. macrocarpa and locally Q. imbricaria. This is indicated by higher frequency in the Bluegrass or other calcareous regions, compared to the Knobs and Appalachian hills (excluding more fertile valleys and other unusual sites).

(iv) Association with sun versus shade. Typical position along this gradient is based on general knowledge, including much vegetation survey and discussion among plant ecologists.

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1          Persistent in deep shade, and generally out-competed in more sunny areas.

2          Moderately persistent in shady forest, but more vigorous in sunny areas.

3          Mostly in partial shade, thin woods or edges, but declining in shady forest.

4          Often in full sun, but most abundant in thin woods and         brushy transitions.

5          Most abundant in full sun with relatively intact grassland or similar vegetation.

6          Most abundant in full sun with much bare ground, rock or water.

ABU   Abundance versus Rarity; and likely historical change. Only native species are provided with global and state abundance ranks. These ranks were assessed by applying the guidelines outlined below to a general review of distributional data across North America and Kentucky. Ranks for abundance were initially developed by The Nature Conservancy and the States’ Natural Heritage Programs during the 1970s and 1980s. Although these were partly updated during the 1990s in preparation for the Conservancy’s ecoregional planning, there has been little systematic effort to incorporate new taxonomic and biogeographic data during the past decade or so. This Atlas provides a comprehensive independant assessment from our viewpoint in Kentucky, which can readily be compared with the ‘official’ ranks if desired. Our use of lower case (g and s) for these global and state ranks indicates their unofficial nature, for contrast with the more official G and S ranks.

 

(i) Global abundance versus rarity. This coding is an elaboration of the traditional ‘G-ranks’ maintained by NatureServe (www.natureserve.org/explorer/ranking.htm), converting that 6-point scale into an 11-point scale. This conversion allows more precision for comparing species within east-central North America. It is still a provisional approximation, probably accurate to ± one unit in most cases, pending compilation of more data by J.T. Kartesz (in prep.), FNA (1993-2006), and others. For taxa with inadequate information, question marks indicate more speculative ranks, possibly off by ± two units.

g0        Probably globally extinct; see ‘GX’

g1        Probably extant but not seen since 1950; see ‘GH’

g2        Less than 5 large viable populations (with A or B quality in the Natural Heritage System) estimated to exist; equivalent to the Heritage Program’s ‘G1’

g3        About 5-10 large viable populations estimated to exist; see ‘G1G2’

g4        About 10-20 large viable populations estimated to exist; equivalent to the Heritage Program’s ‘G2’

g5        About 20-50 large viable populations estimated to exist; see ‘G2G3’

g6        About 50-100 large viable populations estimated to exist; equivalent to the Heritage Program’s ‘G3’

g7        Secure locally but largely restricted to 1-3 ecological regions (e.g., Interior Low Plateaus, Appalachian Plateaus, Ridge-and-Valley, Gulf Coastal Plain); or more widely distributed but restricted to uncommon or threatened habitats; probably 100-250 large viable populations; see ‘G3G4’

g8        Widespread and secure locally but concentrated in a few (ca. 3-6) ecological regions; or more widely distributed but concentrated in uncommon or threatened habitats; similar to the Heritage Program’s ‘G4’ but not closely correlated

g9        Widespread and common within a broad climatic zone of eastern North America (e.g., throughout the southeastern states, or the northeastern states, or the mid-western states); see ‘G4G5’

g10      Widespread throughout most of eastern North America or a similar sized area; similar to the Heritage Program’s ‘G5’

(ii) State abundance versus rarity. Parallel to the global ranks outlined above, this coding of state ranks is an elaboration of the traditional ‘S-ranks’ maintained by State Heritage Programs, which underlie the ‘endangered’ (mostly S1), ‘threatened’ (mostly S2) and ‘special concern’ (mostly S3) categories (e.g., KSNPC 2000), converting that 6-point scale into an 11-point scale.

s0         Probably extinct within Kentucky; see ‘SX’

s1         Probably extant but not seen since 1950; see ‘SH’

s2         Less than 5 large viable populations (A/B quality in heritage system) estimated to exist; equivalent to the Heritage Program ‘S1’

s3         About 5-10 large viable populations estimated to exist; see ‘S1S2’

s4         About 10-20 large viable populations estimated to exist; equivalent to the Heritage Program ‘S2’

s5         About 20-50 large viable populations estimated to exist; see ‘S2S3’

s6         About 50-100 large viable populations estimated to exist; equivalent to the Heritage Program ‘S3’

s7         Secure locally but largely restricted to a few regions (e.g., Interior Low Plateaus, Appalachian Plateaus, Ridge-and-Valley, Gulf Coastal Plain); or more widely distributed but restricted to uncommon or threatened habitats; about 100-250 large viable populations estimated to exist; see ‘S3S4’

s8         Widespread and secure locally but concentrated within a less than half of the state; or more widely distributed but concentrated in uncommon or threatened habitats; similar to the Heritage Program’s ‘S4’ but not closely correlated.

s9         Widespread and secure across most of the state, but uncommon or absent in one or more regional sections (e.g., Coastal Plain, Shawnee Hills, Bluegrass, Appalachian Plateaus); see ‘S4S5’

s10       Widespread and secure in all regions of the state; similar to the Heritage Program’s ‘S5’

(iii) Estimated trend: increased versus decreased ranks since settlement. These provisional estimates are based on information in the previous columns, combined with general knowledge and interpretation. In most cases, the estimated levels of increase or decrease are probably accurate to ± one unit. Question marks indicate more speculative assignments. These indices can be worked into various applications of the ‘conservatism’ concept for each species, including overall ‘floristic quality’ for a particular site (Ladd 1993, Taft et al. 1997). However, local applications of that concept will benefit from a more focused estimation of ranks that considers details of local distribution within each region. Although numerical estimates of changes are provided here, the real range of increases or decreases may be even wider; outer limits may be more accurate in most cases.

-6         Probably or possibly extinct, generally not seen since 1950; these have state rank of s0 or s1

-5         Probably declined by ´ 1/1000 to 1/10,000+; most are highly conservative species of native grassland or open grassy woodland in Class 10 or its transitions, including annuals or short-lived perennials that cannot recolonize into old fields or otherwise recover

-4         Probably declined by ´ 1/100 to 1/1000; some of these are conservative grassland or woodland species with slow recovery by seed, though often persistent vegetatively; others are typical within remnants of unusual vegetation types that have been very much reduced or degraded, especially in Classes 10, 8 and 7, and especially on fertile soils

-3         Probably declined by ´ 1/10 to 1/100; many of these are typical of vegetation types that have been much reduced or degraded, especially in Classes 12, 9 and 6, and especially on fertile soils.

-2         Probably declined by ´ ½ to 1/10; many of these are typical of widespread forest types on average soils, with general reduction across the state, especially in Classes 11, 5 and 4.

-1         Probably declined by up to ´ ½; many of these are typical of widespread forest types in more hilly regions, often with relatively infertile soils, especially in Classes 11, 5 and 4; in such areas, deforestation has been less extensive; other species are typical of more reduced natural vegetation types, but are able to survive or expand into more disturbed artificial habitats

0          Probably little changed, with a factor between ´ ½ and ´ 2; the best examples are typical of stable habitats with little significant degradation, especially rock outcrops and other sites in protected ravines or at high elevation; others are presumed to have a balance of natural habitat degradation versus expansion into artificial habitats, but distinction from ranks –1 and +1 remains speculative

+1        Probably increased by up to ´ 2; many occur in natural habitats that have been degraded and reduced, but are able to survive and prosper locally in farmland, yards, rights-of-way, and other artificial habitats

+2        Probably increased by ´ 2 to 10; most of these are widespread in farmland or other artificial habitats

+3        Probably increased by ´ 10 to 100+; most of these are abundant in farmland or other artificial habitats

+4        Alien that is rare or uncommon (equivalent to s1 to s6)

+5        Alien that is thinly scattered or only locally abundant (equivalent to s7 or s8)

+6        Alien that is widespread across much of state (equivalent to s9 or s10)

*          For codes +4 and +5, asterisks indicate aliens with current rapid increase, which are expected to spread throughout the state; for code +6, asterisks indicate the worst problematic species for natural areas.

<          For codes +4 and +5, ‘less than’ symbols indicate aliens that appear to have increased then decreased, as detailed in footnotes and the full database; most of these species have probably changed from +5 to +4, due to changes in agricultural practices such as weed seed inspections, decline in sheep-rearing, introduction of Festuca arundinacea, and advances of industrial agricultural systems.

Figure 1. General relationship of habitat classes to hydrology. Cliffs and other rocky openings occur on the   upper side; aquatic openings occur on the lower side. Between these extremes, more open, stressed or disturbed vegetation tends to occur to the right, but there are exceptions. See text for further details.

Figure 2. General relationship of upland forest types to the pH-related gradient (left to right), and to mesic versus xeric conditions (lower to upper). This typical gradient in hilly landscapes from mesic to xeric conditions (Class 05 to 11 to 12) is similar to that on deeper soils in farmed landscapes, from forest to thickets to open woodland and old fields (Classes 07 to 08 to 10), but there are significant differences; see text.

Figure 3. Ecoregional Sections of Kentucky.

Figure 4. Abbreviations of Kentucky Counties.

Table 3.  Kentucky County Abbreviations

ADAI     Adair

ALLE     Allen

ANDE    Anderson

BALL     Ballard

BARR    Barren

BATH    Bath

BELL     Bell

BOON    Boone

BOUR    Bourbon

BOYD    Boyd

BOYL    Boyle

BRAC    Bracken

BREA    Breathitt

BREC    Breckinridge

BULL     Bullitt

BUTL     Butler

CALD    Caldwell

CALL     Calloway

CAMP    Campbell

CARL    Carlisle

CARR    Carroll

CART    Carter

CASE     Casey

CHRI     Christian

CLAR    Clark

CLAY    Clay

CLIN      Clinton

CRIT      Crittenden

CUMB   Cumberland

DAVI     Daviess

EDMO   Edmonson

ELLI      Elliott

ESTI       Estill

FAYE     Fayette

FLEM    Fleming

FLOY     Floyd

FRAN     Franklin

FULT     Fulton

GALL     Gallatin

GARR    Garrard

GRAN    Grant

GRAV    Graves

GRAY    Grayson

GREE     Green

GRNP     Greenup

HANC    Hancock

HARD    Hardin

HARL    Harlan

HARR    Harrison

HART    Hart

HEND    Henderson

HENR    Henry

HICK     Hickman

HOPK    Hopkins

JACK     Jackson

JEFF       Jefferson

JESS       Jessamine

JOHN     Johnson

KENT     Kenton

KNOT    Knott

KNOX    Knox

LARU    Larue

LAUR    Laurel

LAWR   Lawrence

LEE        Lee

LESL      Leslie

LETC     Letcher

LEWI     Lewis

LINC      Lincoln

LIVI       Livingston

LOGA    Logan

LYON    Lyon

MADI    Madison

MAGO   Magoffin

MARI    Marion

MARS    Marshall

MART   Martin

MASO    Mason

MCRA   McCracken

MCRE   McCreary

MCLE    McLean

MEAD   Meade

MENI     Menifee

MERC   Mercer

METC    Metcalfe

MONR   Monroe

MONT   Montgomery

MORG   Morgan

MUHL   Muhlenberg

NELS     Nelson

NICH     Nicholas

OHIO     Ohio

OLDH    Oldham

OWEN   Owen

OWSL    Owsley

PEND     Pendleton

PERR     Perry

PIKE      Pike

POWE    Powell

PULA     Pulaski

ROBE    Robertson

ROCK    Rockcastle

ROWA   Rowan

RUSS     Russell

SCOT     Scott

SHEL     Shelby

SIMP      Simpson

SPEN      Spencer

TAYL     Taylor

TODD    Todd

TRIG      Trigg

TRIM     Trimble

UNIO     Union

WARR   Warren

WASH   Washington

WAYN   Wayne

WEBS    Webster

WHIT    Whitley

WOLF    Wolfe

WOOD   Woodfor