SPECIES RICHNESS AFTER LONICERA MAA CKII REMOVAL
FROM AN OLD CEMETERY MACROPLOT
ON DEAD HORSE KNOB, MADISON COUNTY, KENTUCKY
Ralph L. Thompson 1 2
1 Hancock Biological Station
Murray State University
Murray, Kentucky 42071
2 Berea College Herbarium, Biology Department
Berea, Kentucky 40404
ralph_thompson@berea.edu
Derick B. Poindexter
I.W. Carpenter, Jr. Herbarium
Appalachian State University, Biology Department
Boone, North Carolina 28608
poindexterdb@appstate.edu
ABSTRACT
The predominance of Lonicera maackii (Rupr.) Herder (Amur Honeysuckle) in central
Kentucky has made it a significant invasive for continued community interaction studies. To better
understand the dynamics of vegetation succession with respect to this species and overall species
richness, quantitative floristics of two macroplots were made at the summit (312 m) of Dead Horse
Knob (Rucker's Knob) near Berea, in Madison County, east-central Kentucky. An old abandoned
cemetery was cleared of L. maackii and a macroplot (20 x 12 m) served as a test plot while a second
macroplot was placed within a dense thicket of L. maackii to serve as a reference plot. Thirty quadrats
( i x 1 m) were randomly placed within each macroplot as a means to determine species frequency. A
full floristic survey was then conducted of each macroplot. Results from frequency data suggest that
native annual and perennial species will quickly recolonize an area after removal of Amur
Honeysuckle and become the most important components of the site. Likewise, floristic data
demonstrated nearly a three-fold increase in species richness (60 taxa) in the test macroplot after
removal of Amur Honeysuckle compared to species richness (21 taxa) in the reference macroplot. In
the absence of Amur Honeysuckle, vegetation has resulted in a significantly different floristic
assemblage between macroplots. Frequency data from the cleared test macroplot indicate that L.
maackii will remain viable through seedling recruitment, and without further control or disturbance, an
understory thicket of Amur Honeysuckle will undoubtedly be reestablished at this site.
KEY WORDS: Amur honeysuckle, exotics, invasives, Kentucky, Lonicera maackii, species richness
The most aggressive invasive species with the greatest deleterious impact on the total
vegetation and native plants in central Kentucky is undoubtedly Lonicera maackii (Rupr.) Herder
(Amur Honeysuckle). In east-central Kentucky, Campbell et al. (1995) noted that L. maackii was the
most abundant, widespread exotic shrub in successional forests at Raven Run Nature Sanctuary in
adjacent Fayette County. Thompson et al. (2005) and Thompson and Green (2010) reported the
abundance and aggressive impact of L. maackii on the floristic composition and habitats in two studies
of abandoned limestone quarries in the Outer Bluegrass Region of contiguous Clark County and
Garrard County. In neighboring Ohio, Amur Honeysuckle is considered the most abundant naturalized
shrub in the Greater Cincinnati region (Luken & Thieret 1995). Its severe impact on the flora and
natural vegetation of the eastern USA has become extremely detrimental in reducing native herbaceous
Thompson and Poindexter:
cover, shrub and tree seedlings, and species diversity (Gorchov & Trisel 2003; Rundle et al. 2007;
Clark & Weckman 2008).
Luken and Thieret (1995) presented a 150-year chronology of the interaction between people
and the introduction of Lonicera maackii, an invasive east Asian shrub from the Amur River region of
Manchuria. Amur Honeysuckle is a fast-growing, multi-stemmed, deciduous shrub up to 6 meters
high that produces attractive axillary flowers and red berries in the leaf axils of opposite simple leaves.
The fruits are readily disseminated by avian vectors (Ingold & Craycraft 1983; Bartuszevige &
Gorchov 2006). This shrub was introduced into North America in 1896 and was much used in
horticultural settings and as an ornamental landscaping plant; ironically, L. maackii was initially
promoted by the USD A Soil Conservation Service (Luken & Thieret 1995, 1996). Amur Honeysuckle
was much planted in arboreta, botanical gardens, paries, college and university campuses, government
plots, and by the general public until the 1960s. By 1994, Amur Honeysuckle had migrated into at
least 24 states (Trisel & Gorchov 1994). Unfortunately, even though its disastrous influences on
natural plant communities have been repeatedly documented, L. maackii continues to be sold by
numerous plant nurseries as a garden and landscaping plant in the USAaccording to an Internet search
of active websites.
Research focusing on competition and allelopathic effects of Lonicera maackii on native flora
and natural vegetation, its persistent migration, as well as mechanisms for the control of this invasive
species have been extensively documented in recent literature from Kentucky (e.g., Luken 1988;
Luken & Mattimiro 1991; Luken & Goessling 1995; Luken et al. 1995a, 1995b; Luken et al. 1997),
and Ohio (e.g., Ingold & Craycraft 1983; Trisel & Gorchov 1994; Hutchinson & Vankat 1997, 1998;
Medley 1997; Trisel 1997; Deering & Vankat 1999; Gould & Gorchov 2000; Gayek & Quigley 2001;
Collier et al. 2002; Gorchov & Trisel 2003: Hatlman & McCarthy 2004; Mller & Gorchov 2004;
Bartuszevige & Gorchov 2006; Doming & Cipollini 2006; Runkle et al. 2007).
Regardless of the large amount of current research (as exemplified above) that has provided
ample evidence for a focused management of Amur Honeysuckle, it is still only regionally regarded as
an invasive plant pest (MA-EPPC 2011; SE-EPPC 2011). It has yet to be recognized as a Kentucky or
Federal Noxious Weed (see USDA NRCS 2011). Nevertheless, this is not an unusual issue since
other severe invasive species [e.g., Pcrsicaria perfoiiata (L.) H. Gross (Miie-a-minute-vine)] have yet
to be Federally listed despite their rampant spread (Poindexter 20 10). In consideration of this, it
seems rational to continue to produce studies such as this one to help elucidate the impacts of Amur
Honeysuckle on local vegetation.
We conducted a quantitative floristic study on the summit of Dead Horse Knob in east-central
Kentucky, during the summer of 2011. Our objectives were to: 1) determine frequency of taxa through
30 (1 x 1 m) quadrats each within two 20 x 12 m macroplots (one a test plot and one a Lonicera
maackii reference plot) for species richness, species composition, and overall floristic diversity; 2)
survey the two macroplots for species not present in quadrats; 3) document the competitive effects of
L. maackii as a means to provide further data on its negative impact on native flora and vegetation; and
4) promote the need for elevating this species to the forefront of controlled management efforts in the
eastern USA
THE STUDY SITE
Dead Horse Knob (hereafter DHK or knob) is an isolated, dome-shaped, solitary 2.6-ha
"knob" near the city of Berea within southern Madison County, Kentucky (Figure 1). The DHK site is
a part of the 270-ha Berea College Farm and is surrounded by open agriculture and pasture land
(Thompson et al. 2008). It lies within the Knobs-Norman Upland Ecoregion of the Interior Plateau
Physiographic Province (Woods et al. 2002) at 37.583591°N and 84.298871°W (Figure 1). Vegetation
in this ecoregion is highly altered Oak-Hickory Forest (Woods et al. 2002).
The study site is located al the summit (312 m) of DHK where a dense thicket of Lonicera
maackii constitutes much of the understory cover throughout the highly disturbed Dry-Mesic Oak-
Hickory Forest. On the DHK summit, Coyler shaly silt loam is the shallow, strongly acid (4.5-5.0 pH),
excessively well-drained soil derived from weathered Devonian New Albany shale (Weir 1969).
While identified on topographic maps as Dead Horse Knob, it is also recognized as Rucker's Knob by
many local people due to a small private cemetery, the Pullins-Rucker Cemetery, on the knob crest.
The family cemetery began with the interment of a Revolutionary War veteran, Loftus Pullins, Jr.
(1764-1841), and some of his descendents.
The cemetery understory was covered by Amur Honeysuckle along with a few other invasive
wood) plants, a few pole-sized and mature canopy trees, and a very sparse herbaceous layer. The
shrubbery was partially removed from the cemetery in September of 1981 for a commemorative
ceremony held by the Madison County DAR and SAR (Daughters and Sons of the American
Revolution) organizations on November 8, 1981, to recognize the contributions of Loftus Pullins, Jr.
and to dedicate a flat marble monument to his memory.
Afterwards, the cemetery soon was abandoned and secondary succession was allowed to
occur. Over the next 30 years, the Pullins-Rucker Cemetery understory became dominated by a dense
thicket of Lonicera maackii and other invasive shrubs and vines including Lonicera japonica,
Euonymus fortunei, Ligustrum ohtusifohum . and Cdasirus orbiculatus. These five Old World woody
taxa meet the criteria as novel, invasive colonizers by the definition of Davis and Thompson (2000),
i.e., they are truly long-distance colonizers aggressively expanding their geographical range and
having severe impacts on their environment.
Figure 1. Dead Horse Knob (Rucker' Knob), in Madison County. Kentucky. Pullins-Rucker
Cemetery in red-outlined rectangle (not to scale). Map adapted from Google Earth, 2010
DigitalGlobe. Image from January 14, 2004.
Thompson and Poindexter: Lonicera maackii removal 4
METHODS
On September 10, 2010, the Amur Honeysuckle thicket was removed with a chain saw and
hand clippers in the cemetery area to expose the flat graveyard headstones (Figure 2A). The eventual
intent was to clear the cemetery of all woody plants, except healthy canopy trees, and to build a
wooden fence around the cemetery. At the time of brush removal, an inventory of all native and
exotic herbaceous and woody taxa observed was recorded. By the following April 14, 2011, the
cleared area had been colonized by the invasive Stellaria media and a lush regrowth of Lonicera
maackii from stump sprouts, new subterranean shoots, and emerging seedlings.
On May 17, 2011, an area encompassing 25 x 15 m in the cemetery was carefully cleared of
understory Amur Honeysuckle, other woody invaders, and all tree saplings by hand clippers and chain
saw. Cut-stumps were then treated with undiluted Tordoni® RTU Specialty Herbicide from Dow
Agro Sciences™. By mid-July 2011, all treated stumps essentially showed a 100% kill from the
active ingredients, Picloran (4-ammo-3. 5, 6-trichloropicolinic acid) and 2, 4-dichlorophenoxyacetic
acid.
Two 20 x 12 m macroplots were laid out with end steel rods and heavy cord twine on August
24 and 25, 2011. The first macroplot boundary 7 (Figure 2A) was placed within the initial 25 x 15 m
cleared cemetery (hereafter, cemetery test macroplot). The second macroplot (Figure 3A) was placed
in an adjacent dense understoiy thicket of non-altered Lonicera maackii (hereafter, the Amur
Honeysuckle reference macroplot).
Thirty 1 x 1 m cemetery test quadrats and 30 (1 x 1 m) Amur Honeysuckle reference quadrats
were randomly placed within the two macroplots. Frequency was determined for all living rooted
herbaceous and woody plants within the <*amph>i? quadiaK l\\\s quantitatr>o method was derived
from Wuetal. (2002):
Frequency = Number of q uadr ats in whic h a s pecies occu rs X 1 00
Total number of quadrats taken
Frequency is the proportion of the number of samples (i.e., 1 x 1 m quadrats, n=30 quadrats
per macroplot) in which a given species occurs expressed as a percentage. Frequency is a measure of
the abundance and distribution of a species within a stud}' area (i.e., 20 x 12 m macroplots, n=2
macroplots). A frequency value gives information to where the species is located and its probability
of occurring. After quadrat frequency data were gathered, the two macroplots were completely
inventoried for taxa not occurring within the 60 quadrats.
Afloristic similarity between macroplots was determined using the strategy of Barbour et al.
(1999), whom presented the S0renson's Community Coefficient (CC) or Index of Similarity (IS)
method to determine the degree of floristic congruence between two sites of vegetation based on
taxon presence only.
Sorenson's Community Coefficient (CC) = 2 X (number of species common to both A+B) X 100
Total species plot A+ Total species plot B
An annotated list of taxa in this study was compiled with four relevant attributes: 1) origin
(native or exotic), 2) growth duration and habit, 3) macroplot{s) occurrence; and 4) voucher specimen
documentation (see Appendix). Nomenclature follows Weakley (2011).
Figure 2 Cemetery test macroplot. (A) After removal of Lonicem maachi on September 10, 2010,
and (B) secondary successional plant colonization on September 17, 2011.
Figure 3. (A) Margin of the Amur Honeysuckle reference macroplot with a scarcity of plants within
the herb layer, and (B) margin of the Cemetery test macroplot with a substantial herb layer. Images
were taken on September 17, 2011.
Thompson and Poindexter:
RESULTS AND DISCUSSION
Cemetery Test Macroplot.
The 30 cemetery test quadrats contained 51 different species (19 exotics) from 46 genera in
26 families with a total count of 323 individual plants (Table 1). The most important colonizing taxa
in the test quadrats with 50% frequency or greater were Phytolacca americana, Oxalis stricta.
Solarium ptychanthum, Erechtites hieraciifolius, Persicaria longiseta, and Ailanthus altissima (Table
1; Figures 2B & 3B). An additional nine taxa, Carex blanda, Commelina diffusa, Eclipta prostrata,
Chamaesyce maculata, Panicum dichotomiflorum, Passiflora incarnata, Persicaria maculata, Smilax
bona-nox, and Vernonia gigantea, were collected within the cemetery macroplot but not recorded
within the 30 cemetery test quadrats. Thus, a total of 60 taxa were documented for the test macroplot.
Amur Honeysuckle Reference Macroplot.
The 30 Amur Honeysuckle reference quadrats had only 21 species present (seven exotics)
from 18 genera in 15 families with a total count of 110 individual plants (Table 2). The most
prominent taxa within the reference plot quadrats with at least 50% frequency were Lonicera maackii
and L. japonica. Twenty of 21 taxa in the Amur Honeysuckle quadrats were present in the cemetery
quadrats including all seven exotics (Table 3), A single Acer saccharum seedling in an Amur
Honeysuckle quadrat was not present in the cemetery quadrats (Table 3). No other species were
documented outside of the quadrats within this reference macroplot. The herbaceous layer is sparse
under the Amur Honeysuckle thicket (Figure 3A).
Floristic Comparisons.
Species richness was substantially higher in the cemetery test macroplot with 60 documented
taxa (Table 1) versus only 21 within the Amur Honeysuckle reference macroplot (Table 2). Despite
the difference in total taxa for each macroplot, both contained proportionately similar numbers of
exotics (cemetery test macroplot = 31.6%, Amur Honeysuckle reference macroplot = 33.3%).
The IS for these two study sites was 55.6% based on quadrat presence only, but 49,4% when
the additional collection records outside of the quadrats were included. Any two sites with an IS of
50% or greater are considered to be of the same vegetation type. The biological reality of the two
macroplots is best represented by the full floristic composition of both macroplots. Thus, the
differences between the two plots were great enough to consider them as significantly different based
on species composition.
Species Richness and Origin of Taxa.
The species richness from the two 20 x 12 m macroplots combined was 61 taxa (38 herbs. 23
woody), which included 20 naturalized species (32.79%). The 38 herbaceous taxa comprised 25
annuals/biennials (14 exotics), 12 perennials (one exotic), and one perennial vine. The 23 woody taxa
consisted of seven vines (three exotics), four shrubs (two exotics), and 11 trees (one exotic). Fourteen
of the 21 naturalized taxa are stale-listed by the KY-EPPC (2011). The nine "severe threat" invasive
plants are Ailanthus altissima, Carduus nutans, Celastrus orbiculatus, Euonymus fortunei, Ligustrum
obtusifolium, Lonicera japonica, Lonicera maackii, Microstegium vimineum, and Stellaria media.
Three "significant threat' 1 taxa with invasive characteristics are Persicaria longiseta, Persicaria
maculosa, and Setaria faberi, and the two "lesser threat"' taxa are Commelina communis and
Echinochloa crusgalli var. crusgalli.
Secondary Succession.
Several woody taxa observed after Amur Honeysuckle was cleared from the cemetery on
September 10, 2010, included Euonymus fortunei, Ligustrum obtusifolium, Lonicera japonica,
Lonicera maackii, Parthenocissus quinquefolia, Symphoriocarpos orbiculatus. Toxicodendron
radicans var. radicans, and Vitis vulpina. Eight of these nine shrubs and vines were present in
Thompson and Poindexter: Lonicera maackii removal 8
quadrats sampled from both the cemetery macroplot (Table 1) and Amur Honeysuckle macroplot
(Tables 2 & 3), except for Ligustrum obtusifolium m the cemetery macroplot. Some native and exotic
herbaceous and tree seedling volunteers had propagules present in the seed bank (seed pool) before
the September 2010 clearing. Seven herbs observed before Amur Honeysuckle removal in 2010 were
Bidens bipinnata, Commelina communis, Lobelia inflata, Oxalis striata, Persicaria longiseta,
Phytolacca americana, and Steilaria media. These woody and herbaceous taxa invariably had
propagules present in the seed bank before removal.
Volunteering taxa with light-weight, wind-carried fruits and seeds were able to quickly
colonize the cemetery after Amur Honeysuckle removal (Figures 2B & 3B). Native or exotic
herbaceous taxa found in the cemetery quadrats from one growing season included 14 members of
Asteraceae (Ambrosia artemisiifolia, Bidens bipinnata, Bidens frondosa, Cardans nutans,
Conoclinum coelestinum, Conyza canadensis var. canadensis, Erechtites hieraciifolius, Eclipta
prostrata, Eupatorium serotinum, Galinsoga quadriradiata, Sonchus asper, Taraxacum officinale,
Verbesina altemifolia, and Vernonia gigantea), seven taxa of Poaceae (Digitaria ischaemum,
Digitaria sanguinalis, Echinochloa crusgalli var. crusgalli, Microstegium vimineum, Panicum
dichotomiflorum, Setaria faberi, and Setaria pumila), Cyperus strigosus, Lobelia inflata, Oxalis
striata, Persicaria longiseta, Persicaria maculosa, Steilaria media, and Verbena urticifolia var.
urticifolia, among others (Table 1; Appendix). Commelina communis, Persicaria longiseta,
Microstegium vimineum, and Steilaria media were annuals with light-weight seeds within the non-
altered Amur Honeysuckle reference quadrats (Table 2). Tree seedlings with wind-carried samaras
•were Acer rubrum var. rubrum, Acer saccharum, Ailanthus altissima, and Fraxinus i
Trees with heavier diaspores distributed by gravity and/or animals (mainly birds and
mammals) are Celtis occidentalis, Gleditsia triacanthos, Nyssa syivatica, Quercus alba, Quercus
velutina, Primus serotina var. serotina, and Robinia pseudoacacia, while shrubs, woody vines, and
perennials with gravity- dispersed and/or animal-dispersed iruits and seeds include Ligustrum
obtusifolium, Lonicera Japonica, Lonicera maackii, Parthenocissus quinquefolia, Phytolacca
americana, Rubus pensilvanicus, Sambucus canadensis, Symphoriocarpos orbiculatus,
Toxicodendron radicans var. radicans, and Vitis vulpina. No state-listed taxa were encountered in the
two macroplots (KSNPC 2010).
CONCLUSIONS
Our quantitative floristic frequency data and observations support conclusions in literature
that removal of Lonicera maackii allows for a large increase in species richness and overall species
composition primarily because of increased insolation and decreased competition from Amur
Honeysuckle through shade-release. The dense growth habit and allelopathic defenses of this taxon
make it a more efficient competitor than many other invasive species and certainly all the native
species. As evidenced by the Lonicera maackii seedling recruitment in the cemetery test macroplot,
reestablishment of another understory thicket of Amur Honeysuckle is eminent in a short time without
further control measures. In short, well-known "novel" invasive plant colonizers, such as Lonicera
maackii, have severely influenced litter decomposition, disturbed soil nutrient cycles, replaced the
native flora and vegetation, clianged patterns of seedling germination, reproduction, and regeneration,
and altered overall secondary plant succession in their respective environments (Poindexter &
Thompson 2009).
Although our floristic quantitative study at DHK is determined from only a small sample of
quadrat frequency data and personal observations, the density ox* Lonicera maackii has significantly
lowered species richness and species diversity of native annual and perennial herbs, shrubs, and tree
seedlings to be expected in the Oak-Hickory Forest communities of east-central Kentucky. These
deleterious effects are markedly evident when. DHK is compared to a pristine area within the Berea
Thompson and Poindexter:
College Forest, the Anglin Falls Ravine in adjoining Rockcastle County, which has very high species
richness and is currently without any Amur Honeysuckle-infestation (Thompson & Fleming 2004).
ACKNOWLEDGEMENTS
We wish to thank Drs. Guy Nesom and J. Richard Abbott for their critical reviews of our
manuscript. This study is part of a complete floristic survey of the 2.6-ha Dead Horse Knob to further
add biodiversity documentation for an area of social and historical significance in Kentucky.
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USDA, NRCS. 2011. The PLANTS Database. National Plant Data Team, Greensboro, North Carolina.
<http://plants.usda.gov> Accessed 24 Sep 2011.
Weakley, AS. 2011. Flora of the Southern and Mid- Atlantic States. Working draft of 15 May 2011.
Univ. of North Carolina Herbarium, North Carolina Botanical Garden, Chapel Hill, North
Carolina.
Weir, GW 1967. Geologic map of the Berea Quadrangle east-central Kentucky. Map GQ-649. (Map
scale 1:24,000), U.S. Geol. Survey, Washington, DC.
Woods, A.J., J.M Omernik, W.H. Martin. G.J. Pond. W.M, Andrews, S.M. Call, J.A. Comstock, and
D.D. Taylor. 2002. Ecoregions of Kentucky (color poster with map, descriptive text,
summary tables, and photographs. (Map scale 1:1,000,000). U.S. Geo). Survey, Reston
Virginia.
Wu, X.B., RE. Smeins, and R.D. Slack. 2002. Fundamentals of Ecology Laboratory Manual (ed. 3).
edition. Kendall/Hunt Publishing Company, Dubuque, Iowa.
APPENDIX
The annotated list provides voucher records and selective characteristics of taxa documented
in this study. Taxonomic concepts and nomenclature follow Weakley (2011). Exotic taxa are
preceded by an asterisk (*) and invasive pest plants for Kentucky (KY-EPPC 2011) are indicated, by a
double asterisk (**). A plant found in the test macroplot, but not found in quadrats is indicated by a
dagger (|). Alter the scientific name, a vernacular name from either Weakley (2011) or USDA,
NRCS (2011) is given. Growth duration (annual, biennial, perennial) and habit (herb, graminoid,
herbaceous vine, woody vine, shrub, tree) are designated after the scientific name and are adapted
from USDA, NRCS (2011). Habit descriptors are followed by an occurrence designation for each
macroplot: CTP = cemetery test plot and ACP = Amur Honeysuckle reference plot. An italicized
voucher collection number ends each taxon entry.
ANGIOSPERMS: MONOCOTS
COMMELINACEAE, Spiderwort Family
**Commelina communis L., Asiatic Dayflower.
Annual herb; CTP, ACP. 11-650
m \*Commelina diffusa Burmann f. , Creeping
Dayflower. Annual herb; CTP. 11-652
CYPERACEAE, Sedge Family
\Carex blanda Dewey Eastern Woodland Sedge.
Perennial graminoid; CTP. 10-354
Cyperus strigosus L., Straw-Colored Flat-Sedge.
Perennial graminoid, CTP. 11-651
POACEAE, Grass Family
*Digita?ia ischaemum (Schreb.) Schreb. ex
Muhl., Smooth Crab-Grass. Annual graminoid;
CTP. 11-612
*Digitaria sanguinalis (L.) Scop., Hairy Crab-
Grass. Annual graminoid; CTP 11-644
**Echinochloa crusgalli (L.) P. Beauv. var.
crusgalli, Barnyard Grass. Annual graminoid;
CTP. 11-637
**Microstegiumvimineum (Trin.) A. Camas,
Nepalese Browntop. Annual graminoid; CTP,
ACP. 10-941
'\Panicum dichotomiflorum Michx., Fall Panic
Grass. Annual graminoid; CTP 11-636
**Setariafaberi RAW. Herrm., Nodding
Foxtail. Annual graminoid; CTP. 11-643
*Setaria pumila (Poir.) Roem. & Schiult,
Annual graminoid; CTP. 11-620
SMILACACEAE, Greenbrier Family
'\Smilax hona-nox L., Saw Greenbrier. Woody
vine; CTP. 10-1056
ANGIOSPERMS: DICOTS
ADOXACEAE, Moschatel Family
Samhucus canadensis L., Common Elderberry.
Shrub; CTP. 10-402
ANACARDIACEAE, Cashew Family
Toxicodendron radicans (L.) Kuntze var.
radicans, Eastern. Poison-Ivy. Woody vine; CTP
ACP. 10-220
ASTERACEAE, Aster Family
Ambrosia artemisiifolia L., Annual Ragweed.
Annual herb; CTP. 11-611
Bidens bipinnaiaL... Spanish Needles. Annual
herb; CTP. 11-618
Bidens frondosa L., Devil's Beggar-Ticks.
Annual herb; CTP. 11-641
**Carduus nutans L., Nodding Musk Thistle.
Annual/biennial herb; CTP. 10-424
Conoclinium coelesimum (L.) DC, Blue
Mistflower. Perennial herb; CTP. 11-653
Conyza canadensis (L.) Cronquist var. canadensis,
Canadian Horseweed. Annua! herb; CTP. 11-616
■f *Eclipta prostrata (L.) L., False Daisy. Annual
herb; CTP. 11-529
Erechtites hieraciifolius (L.) Raf. ex DC,
Fireweed. Annual herb; CTP. 11-645
Eupatorium serotinum Michx., Late-Flowering
Thoroughwort. Perennial herb; CTP. 11-638
*Galinsoga quadriradiata Ruiz & Pavon, Shaggy
Soldier. Annual herb; CTP. 11-647
*Sonchus asper (L.) Hill, Spiny-Leaved Sow-
Thistle. Annual herb; CTP. 10-415
^'Taraxacum, officinale EH. Wiggers, Common
Dandelion. Perennial herb; CTP. 10-26
Thompson and Poindexter:
Verbesina altemifolia (L.) Britton ex Kearney,
Common Wrngstem. Perennial forb, CTP. 11-
CANNABINACEAE, Hemp Family
Celtis occidentalis L., Northern Hackberry Tree;
CTPACP. 10-394
CAMPANULACEAE, Bellflower Family
Lobelia inflata L., Indian Tobacco. Annual herb;
CTP. 11-639
CAPRIFOLIACEAE, Honeysuckle Family
**Lonicera japonic a Thunb., Japanese
Honeysuckle. Woody vine; CTP, ACP 10-223
**Lonicera maackii CRupr.) Herder, Amur
Honeysuckle. Shrub; CTP, ACP. 10-215
Symphoricarpos orbiculatus Moench, Indian
Currant. Shrub; CTP, ACP. 10-1047
CARYOPHYLLACEAE, Pink Family
**Stellaria media (L.) Vill., Common
Chickweed. Annual herb; CTP, ACP. 10-24
CELASTRACEAE, Staff- Tree Family
**Celastrus orbiculatus Thunb., Oriental
Bittersweet. Woody vine; CTP, ACP. 10-805
**Euonymusfortunei (Turcz.) Hand.-Mazz. ,
Winter Creeper. Woody vine; CTP, ACP. 10-
1049
EBENACEAE, Ebony Family
Diospyros vitginiana I,.. Common Persimmon.
Tree; CTP, ACP. 10-398
EUPHORBIACEAE, Spurge Family
Acalypha rhomboidea Raf., Common Three-
Seeded Mercury. Annual herb; CTP. 11-627
^Euphorbia maculata L., Spotted Sandmat.
Annual herb; CTP. 11-640
FABACEAE, Legume Family
Gleditsia triacanthos L., Honey Locust. Tree;
CTP. 11-458
Robinia pseudoacacia L., Black Locust. Tree;
CTP. 11-452
FAGACEAE, Beech Family
Quercus alba L., White Oak. Tree; CTP, ACP
10-1034
Quercus velutina Lam.. Black Oak. Tree; CTP,
ACP. 10-1041
NYSSACEAE, Sourgum Family
Nyssa sylvatica Marsh., Blackgum. Tree; CTP.
10-403
OLEACEAE, Olive Family
Fraxinus americana I,., White Ash. Tree; CTP,
ACP. 10-1062
"Ligustrum obtusifolium Sieb. & Zucc, Border
Privet. Shrub; CTP 10-333
OXALIDACEAE, Wood-Sorrel Family
OxalisstrictaL., Common Yellow Wood-Sorrel.
Perennial herb; CTP. 11-628
PASSIFLORACEAE, Passion-FIower Family
iPassiflora incamata L., Maypops. Perennial
vine; CTP. 11-617
PHYTOLACCACEAE, Pokeweed Family
Phytolacca americana L., American Pokeweed.
Perennial herb; CTP, ACP. 10-868
POLYGONACEAE, Buckwheat Fam lly
**Persicaria longiseta (Bruijn) Katagawa,
Asiatic Smartweed. Annual herb; CTP, ACP. 10-
839
'\**Persicatia maculosa A. Gray, Lady's Thumb.
Annual herb; CTP. 10-878
ROSACEAE, Rose Family
Primus serotina Ehrh. var. serotina, Wild Black
Cherry. Tree; CTP, ACP. 11-117
Rubus pensihanicus Poir., Pennsylvania
Blackberry. Perennial herb; CTP, ACP. 11-466
SAPINDACEAE, Soapberry Family
Acer rubrumL. var. rabrum, Red Maple. Tree;
CTPACP 10-404
Acer saccharum Marsh., Sugar Maple. Tree;
ACP. 10-192
SIMAROUBACEAE, Quassia Family
**Ailanthus altissima (P. Mill.) Swingle, Tree-of-
Heaven. Tree; CTP. 10-822
SOLANACEAE, Nightshade Family
Solatium carolinense L. var. carolineiise, Horse-
Nettle. Perennial herb; CTP. 10-935
Solatium ptychanthum Dunal ex DC, West Indian
Nightshade. Annual herb; CTP. 11-615
VERBENACEAE, Vervain Family
Verbena urticifoliaL. var. urticifolia, White
Vervain. Perennial herb; CTP. 11-614
VITACEAE, Grape Family
Parthenocissus quinquefolia (L.) Planch.,
Virginia Creeper. Woody vine; CTP, ACP. 10-
343
:tp,
Thompson and Poindexter:
t Dead Horse Knob C
Commelinaceae
Celastraceae
Rosaceae
Verbenaceae
Celastraceae
Cannabaceae
Nyssaceae
Fabaceae
Oleaceae
Oleaceae
Fagaceae
Fagaceae
Number of Quadrats
Phytolaccaceae
Phytolacca americana
Oxalidaceae
Oxalis stricta
Solanaceae
Asteraceae
Erechtites hieraciifolius
Polygonaceae
**Persicaria longiseta
Simaroubaceae
* *Ailanthus altissima
Caprifoliaceae
**Lonicera maackii
Caprifoliaceae
Symphoriocarpos orbicula
Parthenocissus quinquefoh
Campanulaceae
Lobelia inflata
Fabaceae
Robiniapseudoacacia
Vitis vulpina
Caprifoliaceae
**Lonicerajaponica
Caryophyllaceae
**Stellaria media
Bidens bipinnata
Rubus pensihank
**M_icrostegium i
To;
'•adicans v
Verbena urticifolia var. urticifolia
**Celastrus orbiculatus
Celtis occidentals
Primus serotina var. serotina
Acer rubrum var. rubrum
**Carduus nutans
*Digitaria ischaemum
*Digitaria sanguinalis
^Taraxacum officii
Acalypha rhomboi
Diospyros Virginia
Conoclinium coeh
Cyperus strigosus
**Echinochloa en
*Gaiinsoga quadr,
Gleditsia triacanth
26 famines, 51 species (19 exotics), /-
: pest plant for Kentucky (K Y-EPPC 201 1
Thompson and Poindexter:
r Honeysuckle stand
Family
Species
Number o
f Quadrats
Frequency-/.
Caprifoliaceae
**Lonicera maackii
16
53.33
Caprifoliaceae
**Lonicerajaponica
50.00
Caprifoliaceae
Symphoriocarpos orbiculatus
10
33.33
Phytolacca ceae
Phytolacca americana
10
33.33
Vitaceae
Parthenocissus quinquefolia
23.33
Rosaceae
Rub u s pen sth am ens
20.00
Anacardiaceae
Toxicodendron radicans var. radicans
6
20.00
Cannabaceae
Celtis occidenialis
Celastraceae
**Euonymus forhmei
5
16.66
Sapindaceae
Acer rubrum var. rubrum
4
13.33
Commelinaceae
**Commel;na communis
4
13.33
Rosaceae
Prunus serotina var. serotina
4
13.33
Vitaceae
Vitis vulpina
4
13.33
* *Microstegium vimineum
10.00
Caryophyllaceae
**Stellaria media
3
10.00
Fagaceae
Quercusalba
6.66
Polygonaceae
6.66
Sapindaceae
Acer sac charum
1
3.33
Ebenaceae
3.33
Fagaceae
Quercus velutina
1
3.33
Oleaceae
Fraxinus americana
1
3.33
TOTALS:
15 families, 2
1 specs
7 exotics)
18 genera, 1
plants
r Kentucky (KY-EPPC 201 1
Floristic composition of two 20 x 12 m macroplots at Dead Horse Knob, in >
Amur Honeysuckle Reference
Euphorbiaceae
Acalypha rhomboidea
Acer rubrum var. rubrum
Sapindaceae
Acer saccharum
Simaroubaceae
* "Ailanthus altissima
Ambrosia artemisiifolia
Asteraceae
Bidens bipinnata
Bidens frondosa
Asteraceae
"Carduus nutans
Cjperaceae
]Carexblanda
Celastraceae
"Celastrus orbiculatus
Cannabaceae
Celtis occidenialis
Commelinaceae
"Commelina communis
Commelinaceae
\*Commelina diffusa
Asteraceae
Conoclinium coelestinum
Asteraceae
Conyza canadensis var. ca
Cyperaceae
Cyperus strigosus
Poaceae
-Digitaria ischaemum
Poaceae
*Digitaria sanguinalis
Poaceae
**Echinochloa crusgalli V
Asteraceae
fEcliptaprostrata
Erechtites hieraciifolius
Celastraceae
**Euonymus fortunei
Asieraceae
Eupatorium serotinum
Thompson and Poindexter:
Euphorbiaceae
"[Euphorbia maculata
Oleaceae
Asferaceae
*Galinsoga quadriradiata
Fabaceae
Gleditsia triacanthos
Oleaceae
* *Ligustrum obtusifolium
Campanulaceae
Lobelia inflata
Caprifoliaceae
* *Lonicerajaponica
Caprifoliaceae
**Lonicera maackii
Poaceae
* *Microstegium vimineum
Nyssaceae
Oxalidaceae
Oxalis stricta
Poaceae
\Panicum dichotomiflorum
Vitaceae
Parthenocissus quinquefolia
Passifloraceae
Polygonaceae
* *Persicaria longiseta
Polygonaceae
| * *Persicaria maculosa
Phytolaccaceae
Phytolacca americana
Rosaceae
Prurms serotina var. serotina
Fagaceae
Quercus alba
Fagaceae
Quercus velutina
Fabaceae
R ob inia pseudoacacia
Rosaceae
Rubus pensilvanicus
Adoxaceae
Sambucus canadensis
Poaceae
**Setariafdberi
*Setaria pumila
Smilacaceae
\Smilax bona-nox
Solanaceae
Solarium carolinense var. car
Solanaceae
Solarium ptychanihum
Asferaceae
*Sonchus asper
Caryophyllaceae
**Stellaria media
Celastraceae
Symphoricarpos orbiculatus
Aster aceae
^Taraxacum officinale
Toxicodendron radicans var.
Verbenaceae
Aster aceae
Verbesina alternifolia
\Vernonia gigantea
Vitaceae
Vitis vulpina
r Kentucky (KY-EPPC 20 1 1), (t)=found in macroplot I
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