Invasive Plants 

Field and 

Reference Guide:

An Ecological 

Perspective of 

Plant Invaders 

of Forests and 

Woodlands

United States

Department of

Agriculture

Forest Service

Northeastern 

Research Station

Eastern Region 

Northeastern Area 

State & Private 

Forestry

NA-TP-05-04
Cover photograph: Berberis thunbergii invading a disturbed forest

(C. Huebner).
Invasive Plants Field and 

Reference Guide: 

An Ecological Perspective 

of Plant Invaders of 

Forests and Woodlands

Cynthia D. Huebner1,2

with 

Cassandra Olson3

and

Heather C. Smith2

1 Author for Correspondence (chuebner@fs.fed.us)

2 USDA Forest Service

Northeastern Research Station

Morgantown, WV 26505

3 USDA Forest Service

Forest Inventory and Analysis

North Central Research Station

St. Paul, MN 55108

i

INTRODUCTION

Purpose of This Field Guide: 

There are many field guides available about invasive plants 

and their identification. The purpose of this particular field 

guide is to give a scientific synthesis of what is known about 

the behavior of such species in managed, disturbed, and 

pristine forested systems in addition to key information for 

accurate identification. Such information will be helpful when 

prioritizing research questions and choosing the best control 

strategies. Control methods for each species are not provided. 

The most successful control methods are most often sitespecific; over-generalizing control methods might lead to poor 

management and frustrating outcomes. This is not to say that 

the information that is available should not be used; this guide 

just could not do it justice and still achieve its primary goals. 

Four Goals:

1. While there is a great deal of publicly available information 

about many invasive plants, much of this information lacks 

corresponding citations for verification. The first goal of 

this guide is to help provide such information, using mostly 

peer-reviewed scientific publications and other primary 

sources. If information about a species provided in other 

guides or Web pages could not be verified by such sources, 

it was not included in the species description. As research 

and information about each species improve and increase in 

volume, it is likely that some of the research synthesis will 

need to be organized for each species in a separate reference 

book that will accompany this guide in the future.

2. Despite the scientific nature of Goal 1, the second goal is to 

make this guide accessible to a variety of people, including 

private landowners and managers as well as researchers. 

We do this by providing simple, cursory descriptions that 

can then be read about in more detail (if desired) using the 

corresponding citations. A glossary is provided to help 

explain some of the more botanical terms and phrases that 

could not be avoided. 
ii 

3. The third goal is to emphasize the dynamic nature of 

invasions and science. Species will be added and updated 

on an annual basis. Because this can become quite costly in 

terms of complete hardcopy publications, new species will 

be provided on an individual basis rather than by printing 

new field guides annually. 

4. The last goal is to provide a useful, true-to-form field guide 

that can be used extensively in field situations. This requires 

that the guide be small and weatherproof. In addition, not 

every species included will necessarily be of interest to each 

user at a given point in time, which is why we chose to have 

removable pages. This also allows one to easily add new 

pages to their guidebook, ordering one or many new pages at 

a time as they become available.

A Collaborative Effort:

The USDA Forest Service recognizes that the threat of invasive 

plant species to public and private forests is serious and that 

successful progress in prevention, control, research, and 

restoration from the negative impacts of such invasions is only 

possible using a concerted and organized effort. Thus, the 

Forest Service-Eastern Region (Region 9), the Northeastern 

Area (NA) State and Private Forestry, and the Northeastern 

Research Station have joined forces to put this guide together 

for the public. This guide would not have been possible without 

the guidance and funding provided by Nancy Berlin (Region 9) 

and several Region 9 botanists (see acknowledgements) as well 

as Don Dagnan and Noel Schneeberger (NA State and Private 

Forestry). 

This is a work in progress; information and comments from 

the users of this guide are very welcome.

Guide Organization:

This guide contains 15 species (see list after the Introduction) 

that are potential invaders of forests and woodlands. Such 

invaders pose a threat by invading forests of various ages, 

ranging from a recently harvested forest or woodland to an oldgrowth forest. New species will be added using a prioritized 
iii

list of 50 species, which will likely grow in number. Currently, 

the next 15 species to be included (in alphabetical order) 

are Akebia quinata, Ampelopsis brevipedunculata, Cirsium 

arvense, Coronilla varia, Euonymus alatus, Euonymus fortunei, 

Euphorbia esula, Hesperis matronalis, Ligustrum obtusifolium, 

Lonicera morrowii, Rumex acetosella, Ranunculus ficaria, 

Rhamnus frangula, Vinca minor, and Vincetoxicum nigrum.

Because this guide is dynamic in terms of new additions over 

time, there are no page numbers. Instead, this guide is organized 

by habit type (herb, vine, shrub, or tree) and then alphabetically 

by scientific name, making it easy to insert new species as 

they become available. A list of species by both scientific and 

common name is provided in this current guide and will be 

updated as new species are added. Notification of new species 

syntheses will be posted on the Web pages of the Northeastern 

Area State and Private Forestry, Forest Service-Eastern Region 

(Region 9), and Northeastern Research Station. You may also 

request to be placed on a list for such notifications by contacting 

Cynthia Huebner at chuebner@fs.fed.us.

Within each species description, the more common Latin 

synonyms are provided, but are not comprehensive. Also, the 

Natural Resources Conservation Service species code, which 

is commonly used by Forest Inventory and Analysis and other 

organizations, is also provided. Citations, which are referenced 

in the text with small numbers, are listed in the back (after the 

glossary) by habit and by species’ Latin names. The order of the 

numbers in the descriptions reflects source importance. 

Acknowledgements:

We greatly appreciate the comments of the following Region 

9 botanists/ecologists: Marquita Sheehan, Erin Larson, April 

Moore, Harry Pawelczyk, Kirk Larson, Ian Shackleford, 

Jan Schultz, Jack Greenlee, and Mary Beth Deller. William 

McWilliams (Forest Inventory and Analysis) also provided 

useful comments and support for the project. We also thank 

Michelle Frank and Rick Turcotte (NA State and Private 

Forestry) for their comments and Victoria Evans, Nancy Lough, 

Sandy Fosbroke, and Patty Dougherty (NA State and Private 

Forestry) for layout, editing, and publishing of the guide. 
iv 

Species List

By Scientific Name

Herbs

Alliaria petiolata – garlic mustard

Fallopia japonica – Japanese knotweed

Heracleum mantegazzianum – giant hogweed

Microstegium vimineum – Japanese stilt grass

Vines

Celastrus orbiculatus – oriental bittersweet

Lonicera japonica – Japanese honeysuckle

Polygonum perfoliatum – mile-a-minute weed

Pueraria montana var. lobata – kudzu vine

Shrubs

Berberis thunbergii – Japanese barberry

Elaeagnus umbellata – autumn olive

Lonicera maackii – amur honeysuckle, bush honeysuckle

Rhamnus cathartica – common buckthorn

Rosa multiflora – multiflora rose

Trees

Acer platanoides – Norway maple

Ailanthus altissima – tree of heaven
v

Species List

By Common Name

Herbs

Garlic mustard – Alliaria petiolata

Giant hogweed – Heracleum mantegazzianum

Japanese knotweed – Fallopia japonica

Japanese stiltgrass – Microstegium vimineum

Vines

Japanese honeysuckle – Lonicera japonica

Kudzu vine – Pueraria montana var. lobata

Mile-a-minute weed – Polygonum perfoliatum

Oriental bittersweet – Celastrus orbiculatus

Shrubs

Amur honeysuckle, bush honeysuckle – Lonicera maackii

Autumn olive – Elaeagnus umbellata

Common buckthorn – Rhamnus cathartica

Japanese barberry – Berberis thunbergii

Multiflora rose – Rosa multiflora

Trees

Norway maple – Acer platanoides

Tree of heaven – Ailanthus altissima

GARLIC MUSTARD

[Alliaria petiolata (M. Bieb.) Cavara & Grande; 

 A. officinalis Andrz.]

ALPE4

Habit: Erect, shade-tolerant,4 biennial herb.8

Reproduction: By seed;8 preferentially outcrosses but may 

self.6 

Leaves: Lower leaves kidney shaped with palmate venation, 

2-12 cm (3⁄4 to 43⁄4 in) long, arranged in a basal rosette that forms 

the first year and persists through winter; upper leaves alternate, 

triangular, toothed.15,8,4,7

Stems: Up to about 1 m (3 ft); one flowering stem per rosette, 

but up to six;4,17 may branch.8,4,7

Flowers: Second year; numerous; 5-7 mm (1⁄4 in) in diameter, 

white, 4-petaled; mostly in a cluster at the top of the stalk but 

may occur in leaf axils;8,4 bloom late April-June;15,8 open from 

bottom to top; remain open for 2-3 days, but produce nectar 

primarily in day 1; pollinated by medium-sized, short-tongued 

bees and flies, which visit one or two flowers per plant.6

Fruits/Seeds: As many as 3,000 seeds per plant; seeds dark 

brown to black,8,4 enclosed in long (4-7 cm; 11⁄2-23⁄4 in), narrow 

capsules (siliques);8,7 dispersed in late summer; cold stratification required to break dormancy;2,10 germination best in dark, 

moist conditions and lower temperatures (1-5o C; 34-41oF); small 

seed banks are formed; large, persistent seed banks unlikely 

due to high germination rates2 but may be more likely in drier 

conditions;3 germinants do not compete well with second-year 

rosettes;2,11 animal and water dispersed.4

Habitat: Native to Europe; first U.S. introduction in 1868;12 in 

upland or floodplain forest, savannas, roadsides, trail edges, and 

disturbed areas; shady, mesic areas with alkaline soils3,18 but 

found in high light, xeric areas with acidic soils.3,1

Comments: Nutritious;9 rare native butterfly oviposits on 

it but larvae cannot feed;13 herbivory defenses increase with 

wounding; levels vary among populations;5 allelopathic impact 

documented,14 possibly via effect on native plant mycorrhizae.16

Similar Native Species: Basal leaves of Thaspium, Zizia, 

Senecio, Viola spp; similar fruiting structures with several other 

mustards.8 

HERB
GARLIC MUSTARD

Alliaria petiolata (M. Bieb.) Cavara & Grande
JAPANESE KNOTWEED

[Fallopia japonica (Houtt.) Ronse Decraene; 

Polygonum cuspidatum Sieb. & Zucc.; Reynoutria 

japonica Houtt.]

POCU6

Habit: Perennial, herbaceous shrub 3 m (10 ft) or taller;8,20,21,7 

shoots survive one season; rhizomes survive decades; circular 

clonal stands formed in native habitat, senescing centrally.1

Reproduction: Primarily vegetatively via rhizome or shoot 

fragments;4,19,7,3 by seed;8,20,6,7 dioecious;8,6 or gynodioecious;3,7

viable, fertile hybrid (F. x bohemica) result of cross with F.

sachalinensis (also non-native and invasive).11,3

Leaves: Simple and alternate; broadly ovate, 8-15 cm long, 

5-12 cm wide (31⁄8-6 in x 2-43⁄4 in), with abruptly pointed tip, 

truncate base;8,6 stipule (ocrea), a tubular, membranous sheath.6,3

Stems: Round, sometimes ridged,8 glaucous, often mottled;6

hollow internodes7 with swollen nodes.20,6

Flowers: Mid-late summer; small (2-3 mm or 1⁄8 in),3 greenishwhite,20,8,6 1,000s/plant;7

 narrow inflorescences at middle/upper 

nodes;20,8,6 fly and bee pollinated; copious nectar,3 from which 

bees produce a dark, quality honey.2,17

Fruits/Seeds: Fruits 3-winged, 8-9 mm or 1⁄4-3⁄8 in; seeds (3-4 

mm or 1⁄8 in) dark, glossy;8,20,6 germination rate 61-95% in light 

and room temperature; no apparent cold stratification requirement;7

 wind,13 possibly water dispersed (like rhizome and shoot 

fragments); at least one bird species eats the seeds.3

Habitat: Native to Asia; introduced to the U.S. in the mid-late 

1800s;7 disturbed and riparian areas, roadsides, woodlands; 

shade intolerant;3 native substrate volcanic14 with low pH; grows 

in a variety of pH levels and soil types; preference for wet 

habitats;3 seedling survival dependent on water; adults tolerate 

drier conditions;13 USDA hardiness zones 4-8.5 

Comments: Tetraploid, hexaploid, or octoploid;12 polyploidy 

may increase genetic diversity;10 translocates N to radial clones 

until clones take up own N;1 most N in roots if N is limiting;9 C 

remobilized to rhizomes prior to shoot senescence;18 treatment 

for skin disorders, hepatitis, inflammations, natural estrogen 

substitute;15 exudes large quantity of guttation fluid;16 grazed by 

sheep, cattle, horses; F. japonica var. compactum also escapes.3

Similar Native Species: Virginia knotweed (P. virginianum); 

not a shrub; ocreae with bristles; inflorescence a slender spike.8,20

HERB
Fallopia japonica (Houtt.) Ronse Decraene

JAPANESE KNOTWEED
GIANT HOGWEED

[Heracleum mantegazzianum Sommier & Levier.]

HEMA17

Habit: Biennial or perennial herb with a deep (40-65 cm or 

16-26 in), branching tap root;9 blooms once then dies (monocarpic);4,9 some perennials have survived after flowering.9

Reproduction: By seed; cut stems may re-sprout;self-compatible, outcrossing, protandrous hermaphrodite.9

Leaves: Alternate;4,8,9 up to 3 m (10 ft) long, three-parted and 

pinnate;4 pubescent beneath;8 petiole base enlarged, surrounding 

the stem;4 upper leaves become gradually smaller.8

Stems: Often purple-mottled, 2-5 m (7-16 ft) tall, up to 10 cm 

(4 in) in diameter; hollow and ridged.4, 8, 9

Flowers: Compound umbel up to 1.5 m (5 ft) in diameter with 

50 to 150 rays; white ;4,9 most plants flower in third or fourth 

year;2,9 June-August;8,9 pollinated by a variety of insects, mostly 

bees and flies; self-pollination may occur between different 

umbels.5,9

Fruits/Seeds: Fruit elliptic, ridged, and winged, 8-15 mm (3⁄8-5⁄8 

in) in length on elongate stalks; splits in half, each half with one 

seed; shed Aug.-Oct.;4,8,9 over 100,000 seeds per plant possible;9

water, wind, or human dispersed, mostly within 10 m (33 ft) of 

mother plant;2,6,9 germination appears to require moisture and 

cold stratification7,9 and will occur in light or dark; viability 

ranges between 2-15 years.9

Habitat: Native to southwestern Asia;9 introduced as an 

ornamental;2 waste places, roadsides, disturbed woodlands, and 

streambanks;2,8 may invade a range of habitats,3,10 but possible 

preference for open, mesic, and seasonally cold environments.7,9

Comments: Sap has secondary compounds (furanocoumarins) 

that may cause blistering and rashes on humans, with sun 

exposure;3,9 some of the same substances inhibit insect herbivory 

by generalists but specialist insect herbivory is common;1 cattle, 

sheep, goat, pig, mollusc, and snail grazing are common; fungal 

pathogens are noted;9 an H. mantegazzianum x H. sphondylium

(native to UK) hybrid with low fertility is rare in the UK.5,6

Similar Native Species: Cow-parsnip (H. lanatum); flower 

usually has only 15-30 rays and stem reaches only 3 m (10 ft).4

HERB
GIANT HOGWEED

Heracleum mantegazzianum Sommier & Levier.
JAPANESE STILT GRASS

[Microstegium vimineum (Trin.) A. Camus; Andropogon 

vimineus Trin.; Eulalia viminea (Trin.) Kuntze]

MIVI

Habit: Reclining, loosely branching annual to 1 m (3 ft).16,8, 6,11

Reproduction: By seed;6,16, 8 may root at lower nodes.10

Leaves: Mid-vein of leaf blade offset from center; a line 

of silvery hairs runs down the mid-vein on upper surface; 

lanceolate, tapering at both ends, 5-10 mm (1⁄4-3⁄8 in) wide, 3-8 

cm (11⁄8-33⁄ in) long; pale green; leaf sheath collars with ciliate 

hairs.16, 8,14

Stems: Reclining and branching; nodes glabrous and slightly 

swollen.6,8,14

Flowers: Late summer/early fall; terminal spike-like, branching 

inflorescence up to 7 cm (23⁄4 in) long with paired, hairy spikelets; 

in one variety, one spikelet is awned; another variety both are 

awnless;5 may have both cleistogamous and chasmogamous 

flowers;18 chasmogamous flowers associated with populations in 

high light1 and under water stress;7 shade populations primarily 

cleistogamous; potentially highly selfed;7 flowering plants tend to 

be larger than non-flowering plants.7

Fruits/Seeds: Ellipsoid grain 2.8-3.0 mm (1⁄8 in) long; abundant 

seed production may occur infrequently;7 seed bank of at least 

3 years;1,7 seeds mature and are dispersed in late fall16 when they 

appear to be dormant; cold stratification may be required for 

germination;7 water and animal dispersed.14

Habitat: Native to tropical Asia;17,16,8 introduced into the U.S. in 

1919;5 shade tolerant;19 preference for shady areas (closed canopy 

forests, especially riparian areas) but found in high light areas 

(roadsides, ditches, forest borders, and fields);15 possible preference for bare ground, disturbed and acidic soil.1,20, 11,15

Comments: C4 photosynthesis;19,2,3 may acquire more light 

energy using sun flecks;9 has a lower capacity to photosynthesize 

in high light;19 forms a thick thatch of litter, which may prevent 

establishment of natives and itself;7 may alter soil conditions to 

its benefit by increasing pH, nitrification, and nitrate;1,4 association with non-native earthworms possibly due to increased litter 

decomposition or an agricultural connection.12,13

Similar Native Species: Leersia virginica; has hairy nodes, is a 

perennial, and flowers earlier.14

HERB
JAPANESE STILT GRASS

Microstegium vimineum (Trin.) A. Camus
ORIENTAL BITTERSWEET

[Celastrus orbiculatus Thunb.]

CEOR7

Habit: Deciduous, woody, twining vine.6

Reproduction: By seed and vegetatively6 by root suckering;3

dioecious; some perfect;6 some plants monoecious.8,3

Leaves: Alternate, entire leaves with shallow teeth; shape variable but most often less than twice as long (5-12.5 cm; 2-5 in) as 

wide, round with an abruptly pointed tip.6,2

Stems: Light brown;2 may reach up to 5-10 cm (2-4 in) in 

diameter and up to 18 m (59 ft) long;2,9,4 may impact host species 

by reducing photosynthesis and causing structural damage.3,11

Flowers: Short, axillary inflorescences with only a few flowers 

(whitish-greenish); each inflorescence subtended by a leaf longer 

than the inflorescence;6 May-June;15 insect or wind pollinated.1,21

Fruits/Seeds: Fruit, adjacent to a vegetative bud, matures late 

summer/early fall, often remaining on the vines through winter; 

most fall to the ground by early winter;7 yellowish outer skin 

covers a red, fleshy aril containing 3-6 seeds;3 dispersal by 

humans, birds (131 m; 430 ft),16 or small mammals;3 most often 

ingested in winter; defleshed seeds have higher germination 

rates than scarified (due to ingestion) seeds;7 no apparent seed 

bank.19

Habitat: Native to Japan, China, and Korea; not a forest species 

in native habitat;17 introduced into the U.S. in 1860;7 open areas; 

early/late-successional forests;3,17 USDA hardiness zones 4-7.2

Comments: “Sit and wait” strategy of invasion; germinates best 

in shade, but prolific growth may not take place without light;7,13

hybridizes with the native C. scandens, producing a sparingly 

fertile hybrid20 that is more vigorous than the native;14 very low 

root pressure, but conducts water at rates equal to native vines 

with high root pressure; despite evident embolism, keeps leaves 

for ~1 month after first frost; greater rates of secondary growth 

than a native grape species;18 may facilitate grape vine growth;5

evidence of medicinal properties for rheumatoid arthritis12 and 

reversing cancer cell resistance to treatment drugs.10

Similar Native Species: Bittersweet (C. scandens); leaves tend 

to be more than twice as long as wide; inflorescences terminal 

and not next to a vegetative bud.8,15

VINE
ORIENTAL BITTERSWEET

Celastrus orbiculatus Thunb.
JAPANESE HONEYSUCKLE

[Lonicera japonica Thunb.]

LOJA

Habit: Perennial, semi-evergreen to evergreen9,3,5 vine that 

trails or climbs to 7 m (23 ft).15 

Reproduction: Vegetatively; stem cuttings;1 by seed;12,21,11,3,30

may be pollinator limited;1,16,5,22 obligatory outcrosser.16

Leaves: Opposite, entire, oblong, 4-8 cm (11⁄2-31⁄8 in); base 

round/triangular; may have lobes or teeth; lower surface often 

lighter green than upper; surfaces may have a few hairs. 9,15,5

Stems: Young stems pubescent, reddish/light-brown;9,15,12

climber internodes shorter in length than those of trailers.28

Flowers: May-June;15,21,16 white-cream-pink,15,12 yellows with 

age;21,15 2.5-3.8 cm (1-11⁄2 in), 2 reflexed lips; stamens extend 

beyond lips; paired at each node;15 fragrant,19 tubular, pubescent 

inside11 with glandular hairs bearing nectar;22 open at dusk, 

maximizing visits from diurnal (bees) and nocturnal (moths) 

pollinators; nocturnal pollinators disperse pollen further;19 may 

be best adapted for hawkmoths,16,20 which are attracted to rhythmic linalool emission (highest first midnight of 2-day flowering 

period20); diurnal pollinators remove more pollen but with less 

efficiency than hawkmoths.19

Fruits/Seeds: Sept.-Oct.;15,21 black, glossy fruit 0.6 cm (1⁄4 in) 

long15,12 with 4-10 brown-black seeds;15 viability may be low;13 

dispersed by deer, rabbits, bobwhites, turkeys,10 and other 

birds.15

Habitat: Woods, fields, disturbed areas, roadsides, bottomlands, 

and fence rows;12, 21, 27 tolerates shade but most growth in full 

sun; rarely flowers in low light;23 not as shade tolerant as some 

native vines;4 sensitive to dry conditions;2 responds positively to 

an increase in CO2;

24 USDA hardiness zones 4-9.9

Comments: Native to E. Asia;12 introduced into the U.S. in 

mid 1800s 17,26 for horticultural purposes and soil stabilization;15

negative impacts (lower leaf N, photosynthesis, growth) on a 

native host all primarily due to root competition;6,7,8 diploid; less 

genetic diversity than L. sempervirens (tetraploid, native vine);25

greater annual carbon gain than this native;27 forage for deer,31,29

but this native preferred; herbivory increases growth;26 antiinflammatory18 and anti-bacterial/viral properties;14 6 cultivars.9

Similar Native Species: L. sempervirens; leaves glaucous (both 

surfaces), flowers terminal, and connate terminal leaves.12,21

VINE
JAPANESE HONEYSUCKLE

Lonicera japonica Thunb.
MILE-A-MINUTE WEED

[Polygonum perfoliatum L.; Ampelygonum perfoliatum

(L.) Roberty & Vautier; Tracaulon perfoliatum (L.) 

Greene9]

POPE10

Habit: Annual, shallow and fibrous-rooted, climbing vine4,15 to 

6-8 m (20-26 ft);10,15 may behave as a perennial (with a tap root) 

in subtropical climates.12,15

Reproduction: By seed;12 perennials may root at the nodes.15

Leaves: Alternate, simple, entire, glaucous, and glabrous; with 

recurved prickles on lower veins and petioles;5 triangular in 

shape and bright to pale green in color (sometimes reddish when 

young); 3-8 cm (13⁄-33⁄ in) long and 5-9 cm (2-31⁄2 in) wide; 

peltate with the petiole;3,4 sheathing stipule (ocrea) at petiole 

base encircles the stem (perfoliate).4,3,9,2,12

Stems: Wiry, slender, and armed with small, recurved prickles;2,8 climbing or reclining on other plants;12 becoming reddish 

with age.8,15

Flowers: Small (1.5 mm or 1⁄ in);15 10-15 in terminal or 

axillary spike-like racemes 1-2 cm (3⁄8-3⁄4 in) long; perfect, 

greenish-white to pink;4,12 bloom early summer to fall.6,5

Fruits/Seeds: Round, shiny, black achene;12,15 perianth 3-5 

mm (1⁄8-1⁄4 in), persistent, thickening to a fleshy, berry-like, 

iridescent blue covering;4,12 water, bird, small mammal, and 

human dispersed;12,15,6 buoyant;2,12 dispersed July-November;9,12

germination mid-March to April;6 cold stratification may be 

required in colder environments1,8 but detrimental in warmer 

areas; scarification may promote germination;8 persistent seed 

bank for at least 3 years.13

Habitat: Native to E. Asia;2,4,6 introduced into the U.S. in 

1890s10 near Portland, OR, with no local spread;13 found in 

Pennsylvania in the mid 1930s; along streams, in flood plains,12

roadsides, disturbed sites (harvested forests), and open woodlands;7,11 possible preference for moist soils.5

 

Comments: Easily spread in nursery stock;5 relatively shade 

tolerant compared to native congeners;5 performs better in open 

areas;9 numerous native insects feed on it with little effect.9,14

Similar Native Species: P. sagittatum and P. arifolium; leaves 

not glaucous, peltate, or triangular and ocreae not perfoliate.5,11,12

VINE
MILE-A-MINUTE WEED

Polygonum perfoliatum L.
KUDZU VINE

[Pueraria montana (Lour.) Merr. var. lobata (Willd.) 

Maesen & Almeida; P. lobata (Willd.) Ohwi.]

PUMOL

Habit: Perennial, twining vine2 that trails or climbs to 30 m (98 

ft);10,5,19 legume (with nitrogen-fixing bacteria).9

Reproduction: Tubers, root suckers,15 and runners that root at 

the nodes;8,15 by seed;8,10,5 successful seed production rare in cold 

climates;19,16 stem cutting propagation not successful.16

Leaves: Alternate, trifoliate, up to 18 cm (7-8 in) long with long 

petioles; dark green and a pubescent underside; leaflets with 

smooth or lobed margin; middle leaflet usually with three lobes 

and equal base; side leaves 1-2 lobes and unequal bases.19,10, 5

Stems: Up to 2.5 cm (1 in) diameter (some 10 cm (4 in));15

brownish; young stems with tan/bronze hairs;10,19,5,15 may grow 

30 cm (12 in)/day; die back in fall/winter;15 some overwinter.28

Flowers: In leaf axils in elongate, branching racemes to 20 cm 

(8 in) long; up to 2.5 cm (1 in) wide; papilionaceous, reddishpurple, upper petal base yellow;4,15 grape-scented odor;10,19,5,15

May-Nov. on vertically growing plants8 in direct sunlight.15

Fruits/Seeds: Early/late fall; flat, pubescent pod 4-5 cm (11⁄2-2 

in) long;10,15 seeds kidney shaped, 3-4 mm (1⁄8-3⁄ in);30 germination at 15-35o

C (59-95oF), in light or darkness, and best after 

scarification24 (mechanical27 or sulfuric acid25); cold stratification not required;25 mammal and bird dispersed.8

Habitat: Native to China, E. Asia;11,19 introduced into the U.S. 

in late 1800s;11 forest edges, roadsides, old fields, and disturbed 

areas; shade intolerant,2 but found in forests;6,7 not tolerant of 

cold;3 tolerant of compacted, nutrient-poor soils,26 though P is 

limiting;14 older plants with deep roots are more drought tolerant;23 positive growth response to increasing CO2

.

21

Comments: Uses include erosion control,19,15 feed,14,4 fiber,29 

ornamental,11 starch,1 and treatment for alcoholism,18,13 colds, 

asthma, diarrhea, fever, and anemia;23 used to improve soil 

in South American humid tropics;20 high genetic diversity in 

the U.S.;17 continental U.S. has this variety; P. montana var. 

chinensis is in Hawaii; variety hybridization occurs;12 isoprene 

emission possible ozone source;22 shading reduces root growth.9

Similar Native Species: Dioclea multiflora; fruit 2-winged 

along upper suture; flower smaller.10

VINE
KUDZU VINE

Pueraria montana (Lour.) Merr. var. lobata (Willd.) Maesen & 

Almeida 
JAPANESE BARBERRY 

[Berberis thunbergii DC.]

BETH

Habit: Spiny, deciduous shrub, to 2.5 m (8 ft) tall and wide.7,2,10

Reproduction: By seed;7 cut stumps and stems;2,16 may self or 

cross.2

Leaves: Alternate, simple, entire, spatulate, 1.3-2 cm (1⁄2-3⁄4 in) 

long; bright green above, lighter below;2,7,6 in clusters at each 

node;2,10 fall color red to purple depending on the cultivar.2

Stems: Numerous, reddish-brown, angled or grooved, glabrous, 

usually with stiff, short, single spines 1.3 cm (1⁄2 in) long at 

the nodes;2,17 older stems gray; inner bark yellow;17,10 may be 

replaced every few years.16,4

Flowers: Small, perfect, 6 yellow petals, stalked; 1-4 in umbellike clusters at the nodes; April–May;7,2,10 nectaries located on 

both sides of the 6 stamens, which have a tripping mechanism; 

the first visit removes over 50% of the sticky pollen;11 pollinated 

by bees.12

Fruits/Seeds: Early to late summer; bright red, dry, oblong to 

rounded berry 1-1.3 cm (3⁄8-1⁄2 in); 1 seeded;6,10,7 may remain 

on the shrubs through winter;2 production highest in highintermediate light levels;16 cold stratification and alternating 

temperatures improve germination;1,14,16 over 90% of fruit falls 

within 1 m (3 ft) of each shrub but has been mapped as far as 

80 m (262 ft); dispersed by birds (not a preferred food16), deer, 

turkey, and grouse;3 fruit removal may be highest in low light.16

Habitat: Introduced into the U.S. between 1864-18792,16 from 

Japan; full sun to full shade; most soil types and habitats; dry 

ridgetops to wetlands and roadsides to closed canopy forests.13,4

Comments: Growth minimal in low light; survival of seedlings 

drops from 90% in intermediate-high light to 40% in low light, 

but survivors persist; leafs out before canopy and retains leaves 

after canopy leaf fall;16 deer herbivory minimal,3 but severe 

winter damage from rabbits;18 mortality of stems or shrubs not 

related to population density;4,16 at least 47 cultivars;2 may alter 

soil conditions to its benefit by increasing pH, nitrification, and 

nitrate;5 association with non-native earthworms possibly due to 

increased litter decomposition or an agricultural connection.8,9

Similar Native Species: American barberry (B. canadensis); 

toothed leaves and usually 3-pronged spines.15

SHRUB
JAPANESE BARBERRY

Berberis thunbergii DC.
AUTUMN OLIVE

[Elaeagnus umbellata Thunb.]

ELUM

Habit: Deciduous, shrub or small tree up to 6 m (20 ft) tall and 

up to 9 m (30 ft) wide;8,13,12,4 nitrogen-fixing actinorhizal (by the 

actinomycete Frankia9) root nodules present.15

Reproduction: Primarily by seed;8 propagation by stump 

sprouting, by roots, and, for the Ellagood cultivar, by cuttings.16

Leaves: Alternate, simple, oval, entire and wavy; gray-green, 

silvery scaly beneath, with a shimmering appearance;12,8,13 young 

leaves may be silvery on both sides.4

Stems: Twigs silvery or golden brown; often thorny12,8,13 with 

brownish scales, giving a speckled appearance.4,12

Flowers: Clusters of 1-8 in leaf axils; fragrant, tubular, cream 

to light yellow;12 April-June;12,8,13 exterior silvery-scaly; 4 petals 

and stamens.13,12

Fruits/Seeds: Fruit 6-8 mm (1⁄4 in), fleshy, silvery with brown 

scales when immature, ripening to a speckled red12,8,13 or yellow;6

Sept.-Nov.;13,12 edible (bitter to semi-sweet);12 one seeded;8,12,13

dispersed by birds (not preferentially eaten14) and water;11 fruit 

high in lycopene;6 cold stratification improves germination; 

optimal germination at alternating temperatures of 20-30/10oC 

(68-86/50o F);1 persistent seed bank possible (seeds of E. 

angustifolia, a related non-native species, remain viable for 3 

years in the lab and have a dormancy period10); one study found 

that despite relative abundance of the species (compared to 31 

years ago), E. umbellata seeds were not in the seed bank.3

Habitat: Native to Asia; open woods, forest edges, roadsides, 

riparian areas, fence rows, meadows, pastures, sand dunes, mine 

spoils, other disturbed areas;12,13,3,14,11 possibly shade tolerant; 

tolerant of infertile and dry soils;5 acidic soils may reduce 

survival of seedlings;2 USDA hardiness zones 4-8.4

Comments: At least 5 cultivars;16 increases soil nitrogen, which 

may be beneficial to black walnut7 or have a negative impact by 

increasing invasion of other exotics or by changing ecosystem 

properties, neither of which is confirmed for this species yet.

Similar Native Species: Silver-berry (E. commutata); but 

thornless with egg-shaped, shorter leaves that are silver-brown 

and scaly on both sides.8 

SHRUB
AUTUMN OLIVE

Elaeagnus umbellata Thunb.
AMUR HONEYSUCKLE

[Lonicera maackii (Rupr.) Herder;

L. maackii (Rupr.) Maxim.]

 LOMA6

Habit: Deciduous shrub up to 5 m (16 ft) tall and wide.6,12,5,19,16

Reproduction: By seed; main stem may re-sprout; cut young 

stems and bare roots may root.6,5,22

Leaves: Opposite, inverted egg-shaped; broadest in middle, 

tapering at both ends, 3.5-8.5 cm (11⁄2-31⁄2 in) long, tip abruptly 

pointed; petiole short, pubescent;6,12,19 margin entire, fringed with 

hairs; upper surface dark green, paler below; both surfaces have 

hairs on the veins;6,12,2 leaf-out before most woody deciduous 

species;12 senescence later than several species.4

Stems: Grayish-brown, with short hairs on young stems; broad 

ridges and grooves appearing striped on older stems; internodes 

hollow; nodes and young stems may have white-tan pith.12,5

Flowers: Usually two at a node; 15-20 mm (1⁄2-3⁄4 in) long;6,12

May-June; on 5- to 8-year-old plants;3 white, aging to 

yellow;6,12,19 flower stems shorter than leaf petioles;12,19 two 

lipped, anthers longer than the lips;12 nectar mostly sucrose, 

attracting primarily bees; ~21,000 flowers/shrub, ~34 g sugar/

day/shrub.21

Fruits/Seeds: Bright red at maturity in late summer-fall;19,12

pulpy berry with 1-6 seeds (pers. obs.); may require cold, warm, 

or no stratification; optimal germination at 25 or 15oC (77 or 

59oF) in light;1,9 54-81% germinate in warm, moist conditions in 

light (30-55% in dark); seed bank unlikely;15,14 bird dispersed; 

the lipid-poor fruit not preferred;11,23 small mammals consume 

seeds despite bitter seed coat; not a significant part of diet.24,25

Habitat: Native to Eurasia; introduced into the U.S. late 

1850s;5,13 urban areas, old fields, flood plains, upland/lowland 

forests (early-late successional), disturbed ground, wood edges, 

and roadsides;12,19 USDA hardiness zones 3-8.5

Comments: Stratification inconsistencies may be cultivardependent (rem-red requires cold; cling-red does not);22 higher 

growth rates and fitness in high light;10,14,17,13 removal tests show 

increased survival/fecundity of associated natives;8,7,18 American 

robins preferentially nest in L. maackii despite higher predation 

rates (nests are lower) compared to native shrubs.20

Similar Native Species: L. canadensis; smaller; flowers not 

strongly bilabiate; leaves not abruptly pointed.6

SHRUB
AMUR HONEYSUCKLE

Lonicera maackii (Rupr.) Herder
COMMON BUCKTHORN

[Rhamnus cathartica L.]

RHCA3

Habit: Deciduous shrub or tree to 8 m (26 ft) and as wide.7,17,4,3

Reproduction: Primarily by seed; may sucker from base; 

dioecious.7,17,4

Leaves: Elliptic to oblong/obovate, 3-7 cm (11⁄4-23⁄4 in); 

twice as long as wide; pinnately veined, lateral veins curving 

upward;7,14,17 opposite, some alternate, often abruptly pointed 

with rounded teeth (each bearing a gland14,18) on the margin;7,17

upper surface dark green, lower light green;7,14,3 yellow/brown in 

fall;3,8 downy beneath if young;8 early leaf-out,9 late senescence; 

leaf lifespan exceeds that of native shrubs by 58 days.9,1,3

Stems: Branches opposite (or nearly) at right angles to trunk;8

some twigs end in a short thorn;7,17,8,3,14 grayish/yellowish-brown, 

glabrous;3,14 trunk becomes scaly with age.18,14

Flowers: Male 2-6 per cluster with 4 yellowish-greenish petals 

and sepals, 4 stamens; female 2-15 per cluster, usually without 

petals, if present, linear and yellowish-brown, 4 vestigial 

stamens, 4 green sepals shorter than those in the male;14,7,8

April-June;14 appear with the leaves;7,17 females at a 6 to 1 ratio 

to males; honey-scented; calyx tube with nectarial lining; insect 

pollinated (bees and flies).8

Fruits/Seeds: Glossy black at maturity in late summer/early 

fall; fruit 0.5 cm (1⁄4 in) in diameter; drupe contains 3-4 

seeds;8,4,7,3,17 cold stratification may2 or may not1 be required; 

optimal germination at 20 or 30oC (68 or 86oF);2 most fruit 

falls beneath females; bird dispersed, but not preferred (even in 

native habitat);8 seedling establishment more likely on ground 

with little herb cover;6 dormancy and seed bank still unclear.1,13

Habitat: Native of Eurasia; introduced into the U.S. in 1880s;21,1

on calcareous soils in native habitat;8,1 open/shaded areas, 

roadsides, woodlands, riverbanks (not flooded5), pastures,1,12,17,19

and mature forests;9 tolerates various soil conditions.1

Comments: An alternate host for oat crown/leaf rust15 and an 

overwintering host of soybean aphid;20,16 early leaf-out more 

important than late senescence for carbon gain;9 growth rates 

higher in light;10 leaves decompose more rapidly than some 

natives and are high in nitrogen;11 USDA hardiness zones 3-7.3

Similar Native Species: R. caroliniana; flowers perfect and in 

parts of 5; leaves alternate.7,17 

SHRUB
COMMON BUCKTHORN

Rhamnus cathartica L.
MULTIFLORA ROSE

[Rosa multiflora Thunb.]

ROMU

Habit: Perennial, deciduous shrub up to 5 m (16 ft) and as 

wide; long, slender, arching branches.11,16,15

Reproduction: By seed; stem sprouts,11,16 shallow root sprouts, 

and layering (rooting cane tips that touch the ground);23 colonial;16 may self-fertilize or outcross (also with other roses, i.e., 

R. wichuraiana (non-native tetraploid)); male-donor-tetraploid 

crosses have larger fruit, more seeds.17

Leaves: Alternate, pinnately compound with 5-11 elliptic 

to obovate leaflets 2.5 cm (1 in) long with fine teeth;16,23,31

underside of leaflets with hairs and paler than upper surface;23,31

base of leaves have a fringed stipule.16,23,31 

Stems: Flexible, green-red; rigid, recurved thorns with a wide 

base;23 thornless cultivar exists.8,4

Flowers: White or slightly pink, numerous, arranged in terminal panicles; 1-4 cm (3⁄8–11⁄2 in) wide; May-June.15,23,16

Fruits/Seeds: Clustered, hard, maturing to red; 5-7 mm (1⁄4 in) 

wide, egg-shaped; glossy, smooth; Sept.-Oct.,23,11 lasting into 

winter;11 yellowish seeds;23 dispersed by turkeys, deer mice,36

birds (some migratory);27,28 seed bank24 active up to 20 years;21,25 

cold stratification required;1,4 germinates best in light (~60%); 

less than 10% in dark;37 optimum germination at 10-20oC (50-

68oF)38 or 5oC (41oF)1,38 after a long period.38

Habitat: Streambanks, pastures, roadsides,9 forest canopy 

gaps, disturbed areas, and mature forests;20,31 introduced into 

the U.S. from Asia ~1886;25 promoted in 1930s and ‘40s by 

government agencies as a ‘living fence,’ a soil stabilizer, and 

wildlife food/cover;9,12,19 tolerates a variety of soils;11,33,26 USDA 

hardiness zones 5-8.11

Comments: Used as rootstock for other roses32,34 but not the 

best choice for longevity and flower production;22 repeated 

herbivory lethal;3,25 rose rosette disease (also affects other 

cultivated and native roses;5,14) may be lethal;10,13,19 responds 

well to mycorrhizal inoculation;6,7,30 preferential nest site for 

veeries,18 other birds,35 and mammals, such as rabbits;2 at least 3 

cultivars;11 371 rose species considered its progeny.29

Similar Native Species: R. carolina (pasture rose) and R. 

blanda (smooth rose); stipules not fringed.16

SHRUB
MULTIFLORA ROSE

Rosa multiflora Thunb.
NORWAY MAPLE

[Acer platanoides L.]

ACPL2

Habit: Deciduous tree reaching 30 m (98 ft) in height.4,2

Reproduction: By seed;4 can be propagated from roots9 or 

cuttings.2, 9

Leaves: Opposite, green to bronze, smooth, 5-7 lobed with few 

teeth and broad bases up to 18 cm (7 in) wide; wider than long; 

petioles with milky juice, best seen at base of petiole;4,3 fall leaf 

color tending to be bright yellow.2

Stems: Trunk with widely spreading branches (cultivar-dependent; some have narrow canopies) and close bark;3

 twigs are 

olive brown and leaf scars meet to form a sharp angle; buds are 

plump, fleshy, and green to maroon.2

Flowers: Stalked, yellow-green, and perfect; appearing before2 

or with the leaves in spring, in loose clusters (corymbs).3 

Fruits/Seeds: Fruits appear in late spring through summer; 

two-winged (samaras) with the wings almost horizontally (180o

angle) divergent;3 seeds are wind dispersed with low fall rates 

(estimated dispersal distance of 50 m or 164 ft);7 germination 

with cold stratification for 90-120 days;2,1 seed bank potential.5 

Habitat: Introduced into the U.S. from Europe in 175611 and 

planted extensively in urban and suburban areas;2 now also 

found on roadsides and waste places, in hedgerows and roadside 

thickets;3 also spreading into early4 and some late-successional 

forests;6,11,10 withstands sandy, clayey, and acidic and calcareous 

soils; somewhat resistant to drought and tolerates ozone and 

sulfur dioxide; overplanted tree with a splitting bark problem; 

USDA hardiness zones 4 to 7.2

Comments: Shade tolerant;11,8 leaves hold late in autumn and 

root system tends to be shallow;8,2 at least 36 cultivars; susceptible to wilt, anthracnose, tar spot, and leaf scorch;2 regenerates 

prolifically under its own canopy, possibly reducing overall 

diversity;6,11 greater investment in foliage than stem and roots 

may make it less competitive in drier, open environments.8

Similar Native Species: Sugar maple (A. saccharum); does not 

produce milky juice; samara wings are at an angle, and leaf scars 

do not meet.4,3,2 

TREE
NORWAY MAPLE

Acer platanoides L.
TREE OF HEAVEN

[Ailanthus altissima (Mill.) Swingle, A. glandulosa Desf.]

AIAL

Habit: Deciduous tree to 30 m (98 ft) in height.4

Reproduction: By seed and vegetatively via root suckering.8

Leaves: Pinnately compound to 1 m (3 ft) in length with 11-41 

leaflets, each with a ‘thumb’ or lobe at the base; leaflet teeth have 

glands.8

Stems: Bark gray and smooth; younger twigs covered with a 

light brown to reddish brown pubescence; thick or chubby tips; 

may grow up to 2 m (6.5 ft) in one season.4

Flowers: Dioecious; hermaphrodites exist;8 bloom June to July; 

may flower as early as 6 weeks after germination;6 typically 

insect pollinated by a variety of pollinators, including bees.3

Fruits/Seeds: Two-winged samara (180o) with one central seed; 

sets late summer; over 300,000 seeds per tree documented;1 seeds 

may remain on the tree through winter;4,14 wind dispersed; cold 

stratification not required for germination;11 seed bank formation 

thought unlikely,16 though found as part of an urban forest seed 

bank.15

Habitat: Native to China; several introductions into the U.S. 

since 1784;7 found in poor and rich soil on steep and shallow 

slopes, urban areas, open fields and woodlands, and closed 

canopy forests; often associated with disturbed habitats;12,14

USDA hardiness zones 4-8.4

Comments: Extensive cloning;16 at least 5 cultivars;4 crushed 

leaves, stems, and roots smell of rancid peanut butter; allelopathic 

properties present,13 though negative effects may be less severe if 

previously exposed;17 tolerant of several air pollutants;10,18,20 seedlings are drought resistant;21 quassinoid compounds may deter 

some insect herbivory9 and frugivory,19 but seeds and tissue may 

be preyed upon by deer, mice, and voles,5,19,2 even preferentially 

in the case of some invertebrates.2

Similar Native Species: Sumacs (Rhus glabra, R. typhina); 

walnuts (Juglans nigra, J. cinerea); none of the crushed leaves or 

broken stems of these have the same odor; fruits not samaras.8

TREE
TREE OF HEAVEN

Ailanthus altissima (Mill.) Swingle
GLOSSARY

Acidic: pH less than 7; releases protons (hydrogen ions, H+) in 

water.

Actinorhizal: symbiotic relationship of nitrogen-fixing bacteria 

with plant roots; less common than the rhizobia (Rhizobium and 

Bradyrhizobium) nitrogen-fixing bacteria that are often associated with legumes.

Alkaline: pH greater than 7; releases hydroxyl ions (OH-) in 

water.

Allelopathic: ability to inhibit the growth of another plant 

species using toxic chemical substances.

Annual: a plant that completes its life cycle in 1 year— 

germinating from seed, flowering, setting seed, and dying in one 

growing season.

Anther: enlarged terminal pollen-bearing portion of the stamen.

Aril: fleshy, often brightly colored, tissue covering some seeds.

Asexual: reproduction without union of gametes (i.e., union of 

sperm (in the pollen) and egg (in the ovule) in plants); includes 

vegetative and clonal growth.

Awn: bristle-like structure; often associated with grass flowers.

Axillary: the point where the leaf base or leaf petiole meets the 

stem.

Biennial: grows vegetatively for the first year, then flowers and 

dies the next.

Bilabiate: 2-lipped (petals of a flower); bilaterally symmetrical 

in shape.

C4: photosynthetic pathway that uses CO2 more efficiently (at a 

higher energy cost) by allowing storage of CO2 in bundle sheath 

cells and reducing photorespiration; there is less need for gas 

exchange and open stomates; C4 plants are well adapted to high 

light, high temperatures, and low moisture. 

Chasmogamous flower: open; may outcross.

Cleistogamous flower: closed; must self-fertilize.

Clonal: producing vegetative offshoots that can survive on their 

own from the same parent.

Collar: the leaf margin at the intersection of blade and sheath 

surrounding the stem.

Compound: two or more similar parts of the same structure 

(such as flowers or leaflets).
Congener: belonging to the same genus.

Connate: united or fused parts.

Corymb: a flat-topped inflorescence with outer flowers on 

longer pedicels compared to the inner flowers; central flower is 

the youngest.

Cultivar: a variety of a plant species occurring only under 

cultivation (though they may escape into the wild).

Cuneate: wedge shaped (or triangular), narrowing to the point 

of attachment.

Cyme: a flat- or round-topped (or scorpioid) inflorescence 

where the central (or upper) flowers are older and the outer (or 

lower) flowers are youngest.

Deciduous: all leaves shed each year.

Dioecious: male and female unisexual flowers on separate 

plants.

Diploid: having two complete chromosome sets (2n).

Dormancy (for seeds): arrested growth, requiring either further 

embryo development or an environmental cue for germination 

to occur. 

Drupe: fleshy, one- to several-seeded fruit with a stony inner 

layer.

Embolism: filling of vascular tissue (vessels and tracheids) with 

air after water columns rupture (cavitation); such air pockets 

prevent the flow of water.

Evergreen: with leaves that persist for more than one growing 

season.

Fecundity: ability to reproduce; number of offspring produced.

Fertilization: two reproductive haploid cell nuclei (each with 

one chromosome set or 1n) fuse together, forming a zygote (with 

two sets of chromosomes or 2n).

Frugivory: consumption of fruit.

Fruit: the mature ovary of a plant containing seeds.

Generalist: an organism seeking a broad range of resources, 

such as in pollination of flowers, herbivory, or frugivory by 

insects.

Germination: beginning or resumption of growth (usually in 

reference to a seed).

Glabrous: smooth, no hairs.
Glaucous: waxy, bluish green.

Grain: dry, one-seeded fruit, characteristic of grasses. 

Guttation: water expelled from leaf tissue, often along the 

margins, caused by root water pressure.

Gynodioecious: female flowers and perfect flowers on separate 

plants.

Habit: general look or growth form of a plant.

Herbivory: consumption of live plant tissue.

Hermaphrodite: one flower having both functional sexes; same 

as perfect.

Hexaploid: having 6 complete chromosome sets (6n).

Inflorescence: a flower cluster.

Internode: section of stem between two nodes.

Leaf scar: scar left on a twig from a fallen leaf.

Limiting: scarce resource, i.e., N limiting, means nitrogen is 

scarce.

Linalool: a fragrant liquid alcohol.

Lycopene: red carotenoid pigment; an antioxidant; commonly 

found in tomatoes.

Mesic: wet or moist.

Monocarpic: flowering and fruiting once, then dying; also 

called semelparous; opposite is polycarpic or iteroparous, where 

organisms reproduce more than once before dying.

Monoecious: male and female unisexual flowers contained on 

one plant.

Mycorrhiza: a fungus and plant root mutually beneficial 

association (symbiosis); mycorrhizae is plural.

N: nitrogen (nitrate, nitrite, ammonium).

Native: plant species naturally occurring in a given range, not 

introduced to an area by humans.

Nectaries: glands that secrete nectar.

Nitrate: NO3-; one of the preferred forms of nitrogen for uptake 

by plants.

Nitrification: oxidation of ammonium ions (NH4+) or ammonia 

(NH3) to nitrate (NO3-) by free-living soil bacteria.

Nitrogen fixation: conversion of gaseous nitrogen (N2) into 

nitrogen compounds by free-living and symbiotic bacteria; more 

appropriately called dinitrogen fixation.
Node: place of attachment of leaf to stem.

Nodule: swellings on the roots of legumes and other plants 

inhabited by nitrogen-fixing bacteria.

Oblong: sides parallel with ends rounded; longer than broad.

Obovate: egg-shaped but connected at the narrow end.

Ocrea: stipular stem sheath above the leaf base; ocreae is plural.

Octoploid: having 8 complete chromosome sets (8n).

Outcrosser: an individual (i.e., plant) that may be fertilized 

by another individual of the same species (but not of the same 

clone), receiving new genetic material.

Palmate: radiating out from a central axis.

Panicle: a branching inflorescence with pediceled flowers; 

flowers mature at the base first, then upwards.

Papilionaceous: butterfly-shaped; common in pea or bean 

flowers.

Pedicel: stalk that bears a single flower.

Peltate: petiole attached at or near the middle of the underside 

of a leaf, but not going through the leaf.

Perennial: living 2 years or longer.

Perfect: bisexual, having both male and female reproductive 

organs; usually referring to flowers.

Perfoliate: leaf surrounds the stem or petiole; stem or petiole 

goes through the leaf blade.

Perianth: petals and sepals of a flower collectively; most often 

used when petals and sepals look very similar.

Petiole: leaf stalk.

pH: measure of acidity and alkalinity that is the negative 

logarithm of the effective hydrogen ion concentration.

Pinnate: arranged on opposite sides of a central axis; i.e., a 

column of leaflets or veins on each side.

Pollination: pollen transferred from an anther to a stigma (or 

archegonium neck of gymnosperms); may lead to fertilization. 

Polyploidy: having three or more complete chromosome sets.

Protandrous: with male reproductive organs maturing prior to 

those of the female (pollen dispersing before female structure is 

receptive).

Pubescent: with hairs.
Raceme: unbranching, prolonged inflorescence producing 

stalked flowers, maturing from the base upward or outward.

Ray: one of the branches of an umbel.

Recurved: curved backwards.

Reflexed: bent backwards.

Rhizomatous: with rhizomes (underground stems that can send 

up new shoots).

Rosette: radiating cluster of leaves at ground level.

Samara: closed, dry fruit with wings.

Scarification: seed coat degradation that often facilitates 

germination.

Seed: fertilized ovule with a hard coat, embryo, and sometimes 

endosperm (food storage for embryo). 

Seed bank: seeds present in the soil and persisting for various 

time periods (longer than one season).

Self-compatible: individual that is capable of fertilizing itself.

Selfed: self-fertilized.

Senescence: life cycle stage from full maturity to death; can be 

used to describe a whole plant or parts of a plant (such as the 

leaves).

Shade intolerant: grows well or preferentially in high light 

conditions and less well in low light conditions.

Shade tolerant: grows well or preferentially in low light conditions.

Sheath: leaf base surrounding the stem.

Silique: dry fruit, splitting with each half or valve separating 

from the other and leaving a central thin septum.

Simple: only one, or not divided.

Spatulate: spatula-shaped; with rounded, broad top portion and 

narrowing to the base.

Specialist: an organism seeking a specific resource (narrow 

range), such as in pollination of flowers, herbivory, or frugivory 

by insects.

Spike: unbranched inflorescence with flowers without pedicels 

(sessile).

Spikelet: a small, prolonged spike subtended by one or two 

bracts (in grasses and sedges).
Stamens: male sex organ of a flower that produces pollen; 

composed of anther and filament.

Stock: a plant part united with another plant part (the scion) 

of the same or a different species and supplying mostly underground parts; uniting stocks to scions is grafting.

Stratification: seed exposure to different (often colder) 

temperatures to promote germination.

Successional: directional pattern of plant community regeneration or colonization; i.e., going from bare ground or old field 

to young forest (early successional) to mature forest (late 

successional).

Sucker: root or stem offshoot emerging from beneath the soil to 

produce a new plant.

Suture: the line or seam where a mature fruit splits.

Tendril: modified leaf in the form of a narrow, coiling structure, 

providing climbing support for a plant.

Tetraploid: having 4 complete chromosome sets (4n).

Translocate: to transport over a long distance water, minerals, 

or food within one individual or among ramets (individuals) of 

a clone.

Truncate: straight or flat-based as if cut off.

Umbel: flowers of a flat-topped or rounded inflorescence with 

equal length pedicels arising from a single point.

Variety: in the taxonomic hierarchy, a lower than species division being either equivalent to subspecies level or less; naturally 

formed (not cultivated).

Vegetative: propagation using asexual means; non-reproductive 

plant parts.

Venation: vein pattern found in leaves.

Viability: possibility of survival (i.e., of a seed to form a plant).

Xeric: dry.
Invasive Plants Field and 

Reference Guide: 

An Ecological Perspective 

of Plant Invaders of 

Forests and Woodlands

NA-TP-05-04

Citations and Photograph

Information Section

Cynthia D. Huebner1,2

with 

Cassandra Olson3

and

Heather C. Smith2

1 Author for Correspondence (chuebner@fs.fed.us)

2 USDA Forest Service

Northeastern Research Station

Morgantown, WV 26505

3 USDA Forest Service

Forest Inventory and Analysis

North Central Research Station

St. Paul, MN 55108

Citations and Photograph Information

Alphabetized by Habit and Species Latin Name

Herbs

Alliaria petiolata

Text Citations:

1. Anderson, R.C. and T.M. Kelley. 1995. Growth of garlic 

mustard Alliaria petiolata in native soils of different acidity. 

Transactions of the Illinois State Academy of Science 88(3 

and 4): 91-96.

2. Baskin, J.M. and C.C. Baskin. 1992. Seed germination 

biology of the weedy biennial Alliaria petiolata. Natural 

Areas Journal 12(4): 191-197.

3. Byers, D.L. and J.A. Quinn. 1998. Demographic variation 

in Alliaria petiolata (Brassicaceae) in four contrasting 

habitats. Journal of the Torrey Botanical Society 125(2): 

138-149.

4. Cavers, P.B., M.I. Heagy and R.F. Kokron. 1979. The 

biology of Canadian weeds, 35. Alliaria petiolata (M. 

Bieb.) Cavara and Grande. Canadian Journal of Plant 

Science 59: 217-229.

5. Cipollini, D. 2002. Variation in the expression of chemical 

defenses in Alliaria petiolata (Brassicaceae) in the field and 

common garden. American Journal of Botany 89(9): 1422-

1430.

6. Cruden, R.W. and A.M. McClain. 1996. Pollination biology 

and breeding system of Alliaria petiolata (Brassicaceae). 

Bulletin of the Torrey Botanical Club 123(4): 273-280.

7. Fernald, M.L. 1970. Gray’s Manual of Botany, 8th ed. D. 

Van Nostrand Company. New York, NY.

8. Gleason, H.A. and A. Cronquist. 1993. Manual of Vascular 

Plants of Northeastern United States and Adjacent Canada, 

2nd ed. The New York Botanical Garden. Bronx, NY. 

9. Guil-Guerrero, J.L. 1999. Nutritional composition of wild 

edible crucifer species. Journal of Food Biochemistry 23: 

283-294.

10. Lhotska, M. 1975. Notes on the ecology and germination of 

Alliaria petiolata. Folia Geobotanica and Phytotaxonomica 

10(2): 179-183.
11. Meekins, J.F. and B.C. McCarthy. 2002. Effect of 

population density on the demography of an invasive 

plant (Alliaria petiolata, Brassicaceae) population in a 

southeastern Ohio Forest. American Midland Naturalist 

147: 256-278.

12. Nuzzo, V.A. 1993. Current and historic distribution of 

garlic mustard (Alliaria petiolata) in Illinois. The Michigan 

Botanist 32(1): 23-33.

13. Porter, A. 1994. Implications of introduced garlic mustard 

(Alliaria petiolata) in the habitat of Pieris virginiensis

(Pieridae). Journal of the Lepidopterists’ Society 48(2): 

171-172.

14. Prati, D. and O. Bossdorf. 2004. Allelopathic inhibition of 

germination by Alliaria petiolata (Brassicaceae). American 

Journal of Botany 91(2): 285-288.

15. Rhoades, A.F. and T.A. Block. 2000. The Plants 

of Pennsylvania. University of Pennsylvania Press. 

Philadelphia, PA.

16. Roberts, K.J. and R.C. Anderson. 2001. Effect of garlic 

mustard (Alliaria petiolata (Beib.Cavara & Grande)) 

extracts on plants and arbuscular mycorrhizal (AM) fungi. 

American Midland Naturalist 146: 146-152.

17. Susko, D.J. and L. Lovett-Doust. 1998. Variable patterns 

of seed maturation and abortion in Alliaria petiolata

(Brassicaceae). Canadian Journal of Botany 76(10): 1677-

1686.

18. Welk, E., K. Schubert and M.H. Hoffmann. 2002. Present 

and potential distribution of invasive garlic mustard (Alliaria 

petiolata) in North America. Diversity and Distributions 8: 

219-233.

Photograph Information:

Basal leaves (photographer: Jil M. Swearingen, USDI National 

Park Service); flowers (photographer: John Randall, The 

Nature Conservancy); plant (photographer: Dan Tenaglia, 

www.missouriplants.com); mature fruit (photographer: 

Britt Slattery, USFWS). Photographs reproduced from 

www.forestryimages.org or www.invasive.org (mature fruit). 
Fallopia japonica

Text Citations:

1. Adachi, N. 1996. Nitrogen translocation via rhizome 

systems in monoclonal stands of Reynoutria japonica in 

an oligotrophic desert on Mt. Fuji: Field experiments. 

Ecological Research 11: 175-186.

2. Andros, C.F. 2000. Polygonum cuspidatum (Japanese 

knotweed, Mexican bamboo). American Bee Journal 

140(11): 854.

3. Beerling, D.J., J.P. Bailey and A.P. Conolly. 1994. 

Biological Flora of the British Isles: Fallopia japonica

(Houtt.) Ronse Decraene (Reynoutria japonica Houtt.: 

Polygonum cuspidatum Sieb. & Zucc.). Journal of Ecology 

82(4): 959-979.

4. De Waal, L.C. 2001. A viability study of Fallopia japonica

stem tissue. Weed Research 41: 447-460.

5. Dirr, MA. 1998. Manual of Woody Landscape Plants: 

Their Identification, Ornamental Characteristics, Culture, 

Propagation and Uses, 5th ed. Stipes Publishing L.L.C. 

Champaign, Illinois.

6. Fernald, M.L. 1970. Gray’s Manual of Botany, 8th ed. D. 

Van Nostrand Co. New York, NY.

7. Forman, J. and R.V. Kesseli. 2003. Sexual reproduction 

in the invasive species Fallopia japonica (Polygonaceae). 

American Journal of Botany 90(4): 586-592.

8. Gleason, H.A. and A. Cronquist. 1993. Manual of Vascular 

Plants of Northeastern United States and Adjacent Canada, 

2nd ed. New York Botanical Garden. Bronx, NY.

9. Hirose, T. and K. Kitajima. 1986. Nitrogen uptake and 

plant growth I. Effect of nitrogen removal on growth of 

Polygonum cuspidatum. Annals of Botany 58: 479-486.

10. Hollingsworth, M.L. 2000. Evidence for massive clonal 

growth in the invasive weed Fallopia japonica (Japanese 

knotweed). Botanical Journal of the Linnean Society 133: 

463-472.

11. Hollingsworth, M.L., P.M. Hollingsworth, G.I. Jenkins, J.P. 

Bailey and C. Ferris. 1998. The use of molecular markers 

to study patterns of genotypic diversity in some invasive 

alien Fallopia spp. (Polygonaceae). Molecular Biology 7: 

1681-1691.

12. Kim, J.Y. and C-W. Park. 2000. Morphological and 

chromosomal variation in Fallopia section Reynoutria

(Polygonaceae) in Korea. Brittonia 52(1): 34-48.
13. Maruta, E. 1976. Seedling establishment of Polygonum 

cuspidatum on Mt. Fuji. Japanese Journal of Ecology 26: 

101-105.

14. Maruta, E. 1983. Growth and survival of current-year 

seedlings of Polygonum cuspidatum at the upper distribution 

limit on Mt. Fuji. Oecologia 60: 316-320.

15. Matsuda, H., H. Shimoda, T. Morikawa and M. Yoshikawa. 

2001. Phytoestrogens from the roots of Polygonum 

cuspidatum (Polygonaceae): Structure-requirement of 

hydroxyanthraquinones for estrogen activity. Bioorganic & 

Medicinal Chemistry Letters 11: 1839-1842.

16. Mizuno, N., A. Takahashi, T. Wagatsuma, T. Mizuno and H. 

Obata. 2002. Chemical composition of guttation fluid and 

leaves of Petasites japonicus v. giganteus and Polygonum 

cuspidatum growing on ultramafic soil. Soil Science Plant 

Nutrition 48(3): 451-453.

17. Peterson, B. and W. Peterson. 2000. Japanese knotweed, 

Polygonum cuspidatum, as a honey plant. American Bee 

Journal 140(7): 518.

18. Price, E.A.C., R. Gamble, G.G. Williams and C. Marshall. 

2002. Seasonal patterns of partitioning and remobilization 

of 14C in the invasive rhizomatous perennial Japanese 

knotweed (Fallopia japonica (Houtt.) Ronse Decraene). 

Evolutionary Ecology 15: 347-362.

19. Pysek, P., J.H. Brock, K. Bimova, B. Mandak, V. Jarosik, 

I. Koukolikova, J. Pergl and J. Stepanek. 2003. Vegetative 

regeneration in invasive Reynoutria (Polygonaceae) taxa: 

The determinant of invasibility at the genotype level. 

American Journal of Botany 90(10): 1487-1495.

20. Rhoads, A.F. and T.A. Block. 2000. The Plants of 

Pennsylvania. University of Pennsylvania Press. 

Philadelphia, PA.

21. Seiger, L.A. and H.C. Merchant. 1997. Mechanical 

control of Japanese Knotweed (Fallopia japonica [Houtt.]) 

Ronse Decraene): effects of cutting regime on rhizomatous 

reserves. Natural Areas Journal 17(4): 341-345.

Photograph Information:

Leaves (Jack Ranney, University of Tennessee); leaves and 

flowers (Britt Slattery, U.S. Fish and Wildlife Service). 

Photographs reproduced from www.invasive.org.
Heracleum mantegazzianum

Text Citations:

1. Berenbaum, M. 1981. Patterns of furanocoumarin 

distribution and insect herbivory in the Umbelliferae: Plant 

chemistry and community structure. Ecology 62(5): 1254-

1266.

2. Caffrey, J.M. 1999. Phenology and long-term control of 

Heracleum mantegazzianum. Hydrobiologia 415: 223-228.

3. Dawson, F.H. and D. Holland. 1999. The distribution 

in bankside habitats of three alien invasive plants in the 

U.K. in relation to the development of control strategies. 

Hydrobiologia 415: 193-201.

4. Gleason, H.A. and A. Cronquist. 1993. Manual of Vascular 

Plants of Northeastern United States and Adjacent Canada, 

2nd ed. The New York Botanical Garden, Bronx, NY.

5. Grace, J. and M. Nelson. 1981. Insects and their pollen 

loads at a hybrid Heracleum site. New Phytologist 87: 

413-423.

6. Pysek, P. 1991. Heracleum mantegazzianum in the Czech 

Republic: Dynamics of spreading from the historical 

perspective. Folia Geobotanica et Phytotaxonomica 26(4): 

439-454.

7. Pysek, P., M. Kopecky, V. Jarosik and P. Kotkova. 1998. 

The role of human density and climate in the spread of 

Heracleum mantegazzianum in the Central European 

landscape. Diversity and Distributions 4: 9-16.

8. Rhoads, A.F. and T.A. Block. 2000. The Plants of 

Pennsylvania. University of Pennsylvania Press. 

Philadelphia, PA.

9. Tiley, G.E.D, F.S. Dodd and P.M. Wade. 1996. Heracleum 

mantegazzianum Sommier & Levier. Journal of Ecology 84: 

297-319.

10. Willis, S.G. and P.E. Hulme. 2002. Does temperature 

limit the invasion of Impatiens glandulifera and Heracleum 

mantegazzianum in the UK? Functional Ecology 16: 530-

539.

Photograph Information:

Leaves (photographer: Donna R. Ellis, University of 

Connecticut), inflorescence (photographer: Terry English, 

USDA APHIS PPQ), and seeds (photographer: USDA APHIS 

Archives). Photographs reproduced from www.invasive.org.

Microstegium vimineum

Text Citations:

1. Barden, L.S. 1987. Invasion of Microstegium vimineum 

(Poaceae), an exotic, annual, shade-tolerant, C4 grass, into 

a North Carolina floodplain. American Midland Naturalist 

118(1): 40-45.

2. Barden, L.S. 1996. The linear relation between stand 

yield and integrated light in a shade-adapted annual grass. 

Bulletin of the Torrey Botanical Club 123(2): 122-125.

3. Barden, L.S. 1996. A comparison of growth efficiency of 

plants on the east and west sides of a forest canopy gap. 

Bulletin of the Torrey Botanical Club 123(3): 240-242.

4. Ehrenfeld, J.G., P. Kourtev and W. Huang. 2001. Changes 

in soil functions following invasions of exotic understory 

plants in deciduous forests. Ecological Applications 11(5): 

1287-1300.

5. Fairbrothers, D.E. and J.R. Gray. 1972. Microstegium 

vimineum (Trin.) A. Camus (Graminae) in the United States. 

Bulletin of the Torrey Botanical Club 99: 97-100.

6. Fernald, M. L. 1970. Gray’s Manual of Botany, 8th ed. D. 

Van Nostrand Company. New York, NY.

7. Gibson, D.J., G. Spyreas and J. Benedict. 2002. Life 

history of Microstegium vimineum (Poaceae), an invasive 

grass in southern Illinois. Journal of the Torrey Botanical 

Society 129(3): 207-219.

8. Gleason, H.A. and A. Cronquist. 1993. Manual of Vascular 

Plants of Northeastern United States and Adjacent Canada, 

2nd ed. The New York Botanical Garden. Bronx, NY. 

9. Horton, J.J. and H.S. Neufeld. 1998. Photosynthetic 

responses of Microstegium vimineum (Trin.) A. Camus, a 

shade-tolerant, C4 grass, to variable light environments. 

Oecologia 114: 11-19.

10. Hoshikawa, K. 1969. The underground organs of the 

seedlings and the systematics of gramineae. Botanical 

Gazette 130(3): 192-203.

11. Hunt, D.M. and R.E. Zaremba. 1992. The northeastward 

spread of Microstegium vimineum (Poaceae) into New York 

and adjacent states. Rhodora 94(878): 167-170.

12. Kourtev, P.S., J.G. Ehrenfeld and W.Z. Huang. 1998. 

Effects of exotic plant species on soil properties in hardwood 

forests of New Jersey. Water, Air, and Soil Pollution 105: 

493-501.
13. Kourtev, P.S., W.Z. Huang and J.G. Ehrenfeld. 1999. 

Differences in earthworm densities and nitrogen dynamics 

in soils under exotic and native plant species. Biological 

Invasions 1: 237-245.

14. Mehrhoff, L. 2000. Perennial Microstegium vimineum

(Poaceae): An Apparent Misidentification? Journal of the 

Torrey Botanical Society 127(3): 251-254.

15. Redman, D.E. 1995. Distribution and habitat types for 

Nepal Microstegium [Microstegium vimineum (Trin.) 

Camus] in Maryland and the District of Columbia. Castanea 

60(3): 270-275.

16. Rhoads, A.F. and T.A. Block. 2000. The Plants of 

Pennsylvania. University of Pennsylvania Press. 

Philadelphia, PA. 

17. Stewart, R.R. 1945. The grasses of northwest India. 

Brittonia 5(4): 404-468.

18. Tanaka, H. 1975. Pollination of some Graminae (2). 

Journal of Japanese Botany 50: 25-31.

19. Winter, K., M.R. Schmitt and G.E. Edwards. 1982. 

Microstegium vimineum, a shade adapted C4 grass. Plant 

Science Letters 24: 311-318.

20. Zampella, R.A. and K.J. Laidig. 1997. Effect of watershed 

disturbance on Pinelands stream vegetation. Journal of the 

Torrey Botanical Society 124(1): 52-66.

Photograph Information:

Flower, leaf, and sheath close up (photographer: David. 

J. Moorehead, The University of Georgia); population 

(photographer: Chuck Bargeron, the University of Georgia). 

Photographs reproduced from www.invasive.org.
Vines

Celastrus orbiculatus

Text Citations:

1. Brizicky, G. 1964. The genera of Celastrales in the 

Southeastern United States. Journal of Arnold Arboretum. 

45: 206-218.

2. Dirr, MA. 1998. Manual of Woody Landscape Plants: 

Their Identification, Ornamental Characteristics, Culture, 

Propagation and Uses, 5th ed. Stipes Publishing L.L.C. 

Champaign, Illinois.

3. Dreyer, G.D., L.M. Baird and C. Fickler. 1987. Celastrus 

scandens and Celastrus orbiculatus: Comparisons of 

reproductive potential between a native and an introduced 

woody vine. Bulletin of the Torrey Botanical Club 114(3): 

260-264.

4. Fernald, M.L. 1970. Gray’s Manual of Botany, 8th ed. D. 

Van Nostrand Company. New York, NY.

5. Fike, J. and W.A. Niering. 1999. Four decades of old 

field vegetation development and the role of Celastrus 

orbiculatus in the northeastern United States. Journal of 

Vegetation Science 10: 483-492.

6. Gleason, H.A. and A. Cronquist. 1993. Manual of Vascular 

Plants of Northeastern United States and Adjacent Canada, 

2nd ed. New York Botanical Garden. Bronx, NY.

7. Greenberg, C.H., L.M. Smith and D.J. Levey. 2001. Fruit 

fate, seed germination and growth of an invasive vine – an 

experimental test of ‘sit and wait’ strategy. Biological 

Invasions 3: 363-372.

8. Hou, D. 1955. A revision of the genus Celastrus. Annals 

Missouri Botanical Garden 42: 215-302. 

9. Hutchinson, M. 1992. Vegetation management guideline: 

round-leaved bittersweet (Celastrus orbiculatus Thunb.). 

Natural Areas Journal 12(3): 161.

10. Kim, S.E., Y.H. Kim, J.J. Lee and Y.C. Kim. 1998. A new 

sesquiterpene ester from Celastrus orbiculatus reversing 

multidrug resistance in cancer cells. Journal of Natural 

Products 61: 108-111. 

11. McNab, H. and M. Meeker. 1987. Oriental bittersweet: a 

growing threat to hardwood silviculture in the Appalachians. 

Northern Journal of Applied Forestry 4: 174-177.
12. Min, K.R., B.Y. Hwang, H-S. Lim, B-S. Kang, G-J. 

Oh, J. Lee, S-H. Kang, K.S. Lee, J.S. Ro and Y. Kim. 

1999. (-)-Epiafzelechin: Cycloxygenase-1 inhibitor and 

anti-inflammatory agent from aerial parts of Celastrus 

orbiculatus. Planta Medica 65(1999): 460-462.

13. Patterson, D.T. 1975. Photosynthetic acclimation to 

irradiance in Celastrus orbiculatus Thunb. Photosynthetica 

9(2): 140-144.

14. Pooler, M.R., R.L. Dix and J. Feely. 2002. Interspecific 

hybridizations between the native bittersweet, Celastrus 

scandens, and the introduced invasive species, C. 

orbiculatus. Southeastern Naturalist 1(1): 69-76.

15. Rhoads, A.F. and T.A. Block. 2000. The Plants of 

Pennsylvania. University of Pennsylvania Press. 

Philadelphia, PA.

16. Robinson, G.R. and S.N. Handel. 1993. Forest restoration 

on a closed landfill: rapid addition of new species by bird 

dispersal. Conservation Biology 7(2): 271-278.

17. Silveri, A., P.W. Dunwiddie and H.J. Michaels. 2001. 

Logging and edaphic factors in the invasion of an Asian 

woody vine in a mesic North American forest. Biological 

Invasions 3: 379-389.

18. Tibbetts, T.J. and F.W. Ewers. 2000. Root pressure and 

specific conductivity in temperate lianas: Exotic Celastrus 

orbiculatus (Celastraceae) vs. native Vitis riparia (Vitaceae). 

American Journal of Botany 87(9): 1272-1278.

19. Van Clef, M. and E.W. Stiles. 2001. Seed longevity in three 

pairs of native and non-native congeners: assessing invasive 

potential. Northeastern Naturalist 8(3): 301-310.

20. White, O.E. and W.M. Bowden. 1947. Oriental and 

American bittersweet hybrids. Journal of Heredity 38(4): 

125-127.

21. Wyman, D. 1950. Fruiting habits of certain ornamental 

plants. Arnoldia 10(13): 81-85.

Photograph Information: 

Leaves, close up fruit, and plant with fruit (photographer: 

James H. Miller, USDA Forest Service, SE Research Station). 

Photographs reproduced from www.forestryimages.org.
Lonicera japonica

Text Citations:

1. Andrews, E.F. 1919. The Japanese honeysuckle in the 

Eastern United States. Torreya 19: 37-43.

2. Bell, D.J., I.N. Forseth and A.H. Teramura. 1988. Field 

water relations of three temperate vines. Oecologia 74: 

537-545.

3. Carter, G.A. and A.H. Teramura. 1988. Nonsummer stomatal conductance for the invasive vines kudzu and Japanese 

honeysuckle. Canadian Journal of Botany 66: 2392-2395.

4. Carter, G.A. and A.H. Teramura. 1988. Vine photosynthesis 

and relationships to climbing mechanics in a forest understory. American Journal of Botany 75(7): 1011-1018.

5. Carter, G.A., A.H. Teramura and I.N. Forseth. 1989. 

Photosynthesis in an open field for exotic versus native 

vines of the southeastern United States. Canadian Journal of 

Botany 67: 443-446.

6. Dillenburg, L.R., D.F. Whigham, A.H. Teramura and 

I.N. Forseth. 1993. Effects of below- and aboveground 

competition from the vines Lonicera japonica and 

Parthenocissus quinquefolia on the growth of the tree host 

Liquidambar styraciflua. Oecologia 93: 48-54.

7. Dillenburg, L.R., D.F. Whigham, A.H. Teramura and I.N. 

Forseth. 1993. Effects of vine competition on availability 

of light, water, and nitrogen to a tree host (Liquidambar 

styraciflua). American Journal of Botany 80(3): 244-252.

8. Dillenburg, L.R., A.H. Teramura, I.N. Forseth and D.F. 

Whigham. 1995. Photosynthetic and biomass allocation 

responses of Liquidambar styraciflua (Hamamelidaceae) to 

vine competition. American Journal of Botany 82(4): 454-

461.

9. Dirr, MA. 1998. Manual of Woody Landscape Plants: 

Their Identification, Ornamental Characteristics, Culture, 

Propagation and Uses, 5th ed. Stipes Publishing L.L.C. 

Champaign, Il.

10. Dyess, J.G., M.K. Causey and H.L. Stribling. 1994. Effects 

of fertilization on production and quality of Japanese 

honeysuckle. Southern Journal of Applied Forestry 18(2): 

68-71.

11. Fernald, M.L. 1970. Gray’s Manual of Botany, 8th ed. 

D.Van Nostrand Company. New York, NY.
12. Gleason, H.A. and A. Cronquist. 1993. Manual of Vascular 

Plants of Northeastern United States and Adjacent Canada, 

2nd ed. New York Botanical Garden. Bronx, NY.

13. Haywood, J.D. 1994. Seed viability of selected tree, shrub, 

and vine species stored in the field. New Forests 8: 143-

154.

14. Houghton, P.J., Z. Boxu and Z. Xisheng. 1993. A clinical 

evaluation of the Chinese herbal mixture ‘Aden-I’ for 

treating respiratory infections. Phytotherapy Research 7: 

384-386.

15. Kurz, D. 1997. Shrubs and Woody Vines of Missouri, 

Missouri Department of Conservation. Jefferson City, MO.

16. Larson, K.C., S.P. Fowler and J.C. Walker. 2002. Lack of 

pollinators limits fruit set in the exotic Lonicera japonica. 

American Midland Naturalist 148: 54-60.

17. Leatherman, A.D. 1955. Ecological life history of Lonicera 

japonica Thunb. Dissertation. University of Tennessee. 

Knoxville, TN.

18. Lee, S.J., K.H. Son, H.W. Chang, S.S. Kang and H.P. Kim. 

1998. Anti-inflammatory activity of Lonicera japonica. 

Phytotherapy Research 12: 445-447.

19. Miyake, T. and T. Yahara. 1998. Why does the flower 

of Lonicera japonica open at dusk? Canadian Journal of 

Botany 76: 1806-1811.

20. Miyake, T., R. Yamaoka and T. Yahara. 1998. Floral scents 

of hawkmoth – pollinated flowers in Japan. Journal of Plant 

Research 111: 199-205.

21. Rhoads, A.F. and T.A. Block. 2000. The Plants of 

Pennsylvania. University of Pennsylvania Press. 

Philadelphia, PA.

22. Roberts, A.V. 1979. The pollination of Lonicera japonica. 

Journal of Agricultural Research 18(2): 153-158.

23. Robertson, D.J., M.C. Robertson and T. Tague. 1994. 

Colonization dynamics of four exotic plants in a northern 

Piedmont natural area. Bulletin of the Torrey Botanical 

Club 121: 107-118.

24. Sasek, T.W. and B.R. Strain. 1990. Implications of 

atmospheric CO2 enrichment and climatic change for the 

geographical distribution of two introduced vines in the 

USA. Climatic Change 16: 31-51.
25. Schierenbeck, K.A., J.L.Hamricks and R.N. Mack. 1995. 

Comparison of allozyme variability in a native and an 

introduced species of Lonicera. Heredity 75(1): 1-9.

26. Schierenbeck, K.A., R.N. Mack and R.R. Sharitz. 1994. 

Effects of herbivory on growth and biomass allocation in 

native and introduced species of Lonicera. Ecology 75(6): 

1661-1672.

27. Schierenbeck, K.A. and J.D. Marshall. 1993. Seasonal and 

diurnal patterns of photosynthetic gas exchange for Lonicera 

sempervirens and L. japonica. American Journal of Botany 

80(11): 1292-1299.

28. Schweitzer, J.A. and K.C. Larson. 1999. Greater 

morphological plasticity of exotic honeysuckle species may 

make them better invaders than native species. Journal of 

the Torrey Botanical Society 126(1): 15-23.

29. Sheldon, J.J. and M.K. Causey. 1974. Deer habitat 

management: Use of Japanese honeysuckle by white-tailed 

deer. Journal of Forestry 72(5): 286-287.

30. Slezak, W.F. 1976. Lonicera japonica Thunb. an aggressive 

introduced species in a mature forest ecosystem. M.S. 

Thesis, Rutgers University. New Brunswick, NJ.

31. Sotala, D.J. and C.M. Kirkpatrick. 1973. Foods of whitetailed deer, Odocoileus virginianus, in Martin County, 

Indiana. American Midland Naturalist 89(2): 281-286.

Photograph Information:

Leaves and flowers (photographer: Chuck Bargeron, University 

of Georgia); leaves (photographer: Ted Bodner, Southern 

Weed Science Society); fruit (photographer: Ted Bodner, 

Southern Weed Science Society). Photographs reproduced from 

www.invasives.org.

Polygonum perfoliatum

Text Citations:

1. Baskin, C.C. and J.M. Baskin. 2001. Seeds: Ecology, 

Biogeography, and Evolution of Dormancy and 

Germination. Academic Press. San Diego, CA.

2. Cusick, A.W. and M. Ortt. 1987. Polygonum perfoliatum L. 

(Polygonaceae): A Significant New Weed in the Mississippi 

Drainage. SIDA Contributions to Botany 12(1): 246-249.

3. Fernald, M.L. 1970. Gray’s Manual of Botany, 8th ed. 

Illustrated. D. Van Nostrand Company. New York, NY. 

4. Gleason, H.A. and A. Cronquist. 1993. Manual of Vascular 

Plants of Northeastern United States and Adjacent Canada, 

2nd ed. New York Botanical Garden. Bronx, NY.

5. Hickman, J.C. and C.S. Hickman. 1977. Polygonum 

perfoliatum: a recent Asiatic adventive. Bartonia 45: 18-23.

6. McCormick, L.H. and N.L. Hartwig. 1995. Control of the 

noxious weed mile-a-minute (Polygonum perfoliatum) in 

reforestation. Northern Journal for Applied Forestry 12: 

127-132.

7. Oliver, J. D. and N.C. Coile. 1994. Polygonum perfoliatum

L. (Polygonaceae), the Mile-a-minute weed. Florida 

Department of Agriculture & Consumer Services Division 

of Plant Industry. Botany Circular No. 29. November/

December 1994. 

8. Oliver, J.D. 1996. Mile-a-minute weed (Polygonum

perfoliatum L.), an invasive vine in natural and disturbed 

sites. Castanea 61(3): 244-251.

9. Park, C.W. 1986. Nomenclatural typifications in 

Polygonum Section Echinocaulon (Polygonaceae). Brittonia 

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Leaves and flowers (photographer: Leslie J. Mehrhoff, 

University of Connecticut); fruit (photographer: Barry Rice, 

sarracenia.com); plant (photographer: Jil M. Swearingen, 

USDI National Park Service). Photographs reproduced from 

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Leaves, flowers, and shrub in flower (photographer: James 

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www.invasive.org.
Lonicera maackii

Text Citations:

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Fruit (photographer: Chuck Bargeron, The University of 

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Rhamnus cathartica

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Rosa multiflora 

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Photograph Information:

Flowers and fringed stipule (photographer: James H. 

Miller, USDA Forest Service, SE Research Station); fruit 

(photographer: David J. Moorhead, University of Georgia). 

Photographs reproduced from www.invasive.org.
Trees

Acer platanoides

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Photograph Information:

Leaf and flowers (photographer: Paul Wray, Iowa State 

University), and samara (photographer: Bill Cook, 

Michigan State University) Photographs reproduced from 

www.forestryimages.org.
Ailanthus altissima

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1. Bory, G. and D. Clair-Maczulajtys. 1980. Production, 

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of extracts from tree of heaven (Ailanthus altissima). 

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Photograph Information:

Leaves and twig with mature fruit (photographer: James Miller, 

USDA Forest Service, SE Research Station); fruit with reddish 

color, not all cultivars have this color (photographer: Chuck 

Bargeron, University of Georgia). Photographs reproduced from 

www.forestryimages.org.
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