Biological Invasions (2006) 8:1501–1510 Ó Springer 2006

DOI 10.1007/s10530-005-5842-1

Status, invasiveness and environmental threats of three tropical American

invasive weeds (Parthenium hysterophorus L., Ageratum conyzoides L.,
Lantana camara L.) in India

Ravinder K. Kohli*, Daizy R. Batish, H. P. Singh & Kuldip S. Dogra

Centre for Environment, Department of Botany, Panjab University, Chandigarh 160014, India; *Author for
correspondence (e-mail: rkkohli45@yahoo.com; fax: +91-172-2541022)

Key words: Ageratum conyzoides, allelopathy, invasive weeds, Lantana camara, Parthenium hysterophorus,
tropical America

Abstract

Invasive weeds have threatened the integrity of ecosystems throughout the world. They affect not only the
species diversity of native areas but also their biological integrity. In India, a number of invasive exotic
weeds have been reported but some viz. Parthenium hysterophorus, Lantana camara and Ageratum cony-
zoides, especially those from tropical America are troublesome and have caused adverse ecological, eco-
nomic and social impact. These weeds can be seen growing in different landscapes but are luxuriantly
localized in unattended forests and cultivated areas. Parthenium hysterophorus (Asteraceae, commonly
known as congress grass) is perhaps the most troublesome and noxious weed of urban and rural India.
Besides rapidly colonizing areas replacing the native vegetation, it is also known to cause a number of
human health problems such as skin allergy, rhinitis and irritation to eyes of the residents in the vicinity.
Likewise, it causes fodder scarcity in addition to being unpalatable and toxic to livestock. Lantana camara
(Verbenaceae), another serious tropical American pest, has encroached upon large areas of land, especially
the forests where it has virtually replaced the forest floor vegetation and reduced tree growth. Also because
of its bushy and spreading type of growth it obstructs forest operations. The third weed, Ageratum co-
nyzoides (Asteraceae, Billy goat weed) has invaded agricultural fields. It interferes with crops and causes
yield reductions of major staple crops of India. When it invades rangeland areas, it out competes native
grasses causing scarcity of fodder. These weeds have similar growth strategies such as fast growth rates,
short life-cycles, greater reproductive potential, high competitive abilities and allelopathy that make them
successful invaders of native habitats. Mechanical, chemical, biological and cultural control tactics have
failed individually, though integrated approaches combining all these approaches along with community
participation and proper land management have been relatively successful. This paper presents various
aspects of biology, ecology, hazards and control measures of these weeds.

Introduction of the most serious threats to biodiversity, caus-

Invasion by non-native exotic species in alien
environments poses a major threat to native
plant communities and alters fundamental struc-
tures and functions of ecosystems. It poses one
ing major changes in vegetation at a global level
(Vitousek et al. 1996; Mack et al. 2000). Though
biological invasion has been a problem for a long
time, interest in this field surged in the 1980s
when SCOPE (Scientific Committee on the
1502
Problems of the Environment) identified the
problems of biological invasions as a globally
significant issue (Simberloff 2003). Studies have
shown that invasion of plant species in alien
environments causes ecosystem imbalance and
thus is detrimental to ecological integrity and
economies of the invaded area (Pimentel et al.
2001). The impact, however, varies from place to
place depending upon the physio-geographical
features, human population density and diversity
of landscape in the area of potential invasion
(Pysek and Prach 2003). Keane and Crawley
(2002) pointed out that invasive species often
possess specific spreading dynamics such as a
tendency to escape from natural predators and
the ability to occupy vacant niches. Further, they
have not co-evolved with native flora and fauna.
As a result, the native plants lack the adaptive
strategies to the chemicals released by the inva-
sive alien plants. In addition, invasive plant spe-
cies generally possess unique traits such as fast
vegetative and reproductive growth, wide adapt-
ability and ecological adaptations to withstand
environmental stress in the alien environment
(Baker 1974). The success of plant species in the
alien environment has been attributed to lack of
natural predators or possession of novel weapons
such as allelopathy (a phenomenon in which a
plant releases chemicals into soil that may ad-
versely affect the growth of other plants) (Heirro
and Callaway 2003). Invasive species are also
able to undergo genetic changes due to selection
pressure and respond quickly to anthropogenic
disturbances (Sakai et al. 2001). There is need to
investigate on the extent of disturbance caused
by the invasive species in the alien ecosystems,
and the factors facilitating invasion so that
appropriate management steps can be taken.
Status of invasive plants in India
India is one of the richest centres of biodiversity
due to a wide range of variations in climate,
geography and landscape (UNEP-WCMC 2000).
It is the seventh largest country in the world and
second in Asia with a land frontier of 15,200 km
and coastline of 7516 km. Physically, it has four
well defined regions viz. Himalayan mountains,
Gangetic river plains, Southern Deccan Plateau
and Islands of Lakshdeep and Nicobars. The
Himalayas, extending from east to west in the
North of India, are one of the 34 hotspots of
biodiversity and home to a variety of plants
exhibiting great diversity and richness (Conserva-
tion International 2005). It represents a unique
combination of endemics and exotics, some of
which have proven to be useful as crop plants,
for instance. However, the regional diversity is
under enormous threat from various biotic, abi-
otic and anthropogenic induced pressures. Nota-
ble among these is the presence of a large
number of exotic plant species that have either
been introduced deliberately or have entered
accidentally. As per a report in 1977, nearly 18%
of the Indian flora is constituted by adventive
aliens of which 55% are American (Nayar 1977).
There are, however, no recent data regarding the
exact number and incidence of these introduced
invasive species in India. Nevertheless, some
troublesome noxious known weeds, namely Lan-
tana camara, Ageratum conyzoides, Mikania mi-
crantha, Eupatorium odoratum and Parthenium
hysterophorus, are invasive in nature and threaten
the integrity of the native ecosystems (Saxena
1991). Among these, the three South American
and tropical American weeds such as Lantana ca-
mara, Parthenium hysterophorus, Ageratum cony-
zoides (hereafter Lantana, Parthenium, and
Ageratum, respectively) are the most problematic
(Kohli et al. 2004). They have established them-
selves not only in the plains but also in hilly
areas, particularly in the Himalayan ecosystems
(Kohli et al. 2004).
History of introduction in India
Lantana was introduced into India in 1809 as an
ornamental hedge in Calcutta’s gardens. From
there it escaped and became invasive in nature.
Parthenium, on the other hand, is believed to
have entered India between 1950s and 1960s
along with food grains imported from the USA.
It was first spotted in 1955 on the outskirts of
Pune (Maharashtra) and described botanically in
1956 by Rao. However, the workers have ques-
tioned the time and mode of entry into India
(Kohli and Rani 1994); reports indicate the pres-
ence of Parthenium in India as early as 1810

(Bennet et al. 1978), a view supported by the her-
barium specimens, catalogues and notes available
about the weed in recognized national herbaria
and institutes. Based on these differing views
regarding the entry of Parthenium, it was con-
cluded that though the weed occurred in India
long before 1955, it probably remained uncom-
mon, then spread rapidly throughout the plains
and into hilly tracts. As regards the mode and
time of Ageratum entry into India, no informa-
tion is available. However, it was known to exist
in India well before 1882 as reported in ‘‘The
Flora of British India’’ (Hooker 1882). Further-
more, probably it was introduced as an orna-
mental plant and later it escaped and assumed a
weedy habit, though no authentic report is avail-
able in this regard.
Biotic feature favouring invasiveness
The weeds Lantana, Ageratum and Parthenium
are native to tropical America and have spread
to other tropical and subtropical regions of the
world including India. Lantana (commonly
known as lantana, prickly lantana, wild sage or
invading lantana), being shade tolerant prefers to
grow under the canopy of forest trees or planta-
tions. It is also common in pastures, grasslands,
along riverbanks and roadsides, and even agri-
cultural fields. Parthenium (commonly known as
congress grass, carrot grass, ragweed parthenium
and feverfew) is primarily a weed of wastelands
with a potential to invade any habitat. It is com-
mon in pastures, agricultural areas, vacant urban
areas, roadsides, and railway tracts. It has star-
ted invading higher altitudinal ranges and is rap-
idly encroaching upon hilly tracts after becoming
established in the lower and middle Himalayas
(Kohli et al. 2004). Ageratum (commonly known
as billy goat weed, bill goat weed, goat weed or
tropical white weed) is a weed of arable land
found mostly in cultivated areas, grasslands and
pastures interfering with the growth and develop-
ment of crops, grasses and forage crops. It is a
serious invader of the medicinally and aromati-
cally important herbal under storey of the forest
floors and greatly threatens the abundance and
diversity of these herbal species. Morphologi-
cally, these weeds are quite distinct and show
1503
diverse life-forms, habit, morphology and repro-
ductive biology. The area fully infested by each
weed along with their close-up are shown in Fig-
ure 1. Various biotic features of these weeds are
described in Table 1.
Reasons for successful invasion
Growth Strategies
Lantana flowers throughout the year. It produces
enormous (number ranges from 10,000 to 12,000
per plant) fruits varying in size from 3 to 6 mm
in diameter and contain 1–2 seeds that are dis-
persed to other areas through bird droppings or
by goats and sheep or accidentally by even hu-
mans. These germinate quickly and form dense
impenetrable thickets that can survive even after
forest fires. The weed also spreads by suckering.
Seed germination and suckering are stimulated
by burning and slashing, both roots and shoots
can coppice after cutting, browsing or herbicidal
treatment.
Parthenium also possesses an enormous ability
to grow and establish fast. It can grow in almost
all types of soils, though less so in lateritic soil.
Its seeds germinate throughout the year (irrespec-
tive of photoperiod, temperature variations, or
seasons) and, if moisture is favourable, form
dense monocultural stands. It grows luxuriantly
in the rainy season and in more humid areas and
climates. Under water-stressed conditions, it re-
mains a rosette and bolts with the onset of the
rains or moisture. Further, it regenerates quickly
from the root stumps or parts, even petioles or
midribs left in the soil (Kohli and Rani 1994). It
flowers profusely, producing abundant seeds (ap-
prox. 25,000 per plant; Navie et al. 1996) and
forms an enormous seed bank in soil. Seeds,
small and light, are easily disseminated by wind
or water for quick colonization in uninvaded
areas.
Ageratum is highly adaptable and can grow in
a variety of habitats. It prefers sandy soil with
sufficient moisture in the cultivated fields, grass-
lands and disturbed sites. It produces a large
number (8000–10000/plant) of fruits (achene)
that have a pappus. These are easily dissemi-
nated by air, water and animals. The seeds are
1504
Figure 1. Pictures of Lantana camara, Parthenium hysterophorus and Ageratum conyzoides. Column (a) shows fully infested area
whereas (b) shows a close up of each plant.
photoblastic and remain viable for one year. The
optimum temperature range for the germination
of seeds is 20–25 °C. It also spreads fast vegeta-
tively through stolons.
Allelopathy
Allelopathy, the release of chemicals by a plant
into the environment that adversely affects the
growth of other plants, is one of the mechanisms
that enables weeds to successfully colonize an
alien environment (Bais et al. 2003; Heirro and
Callaway, 2003). Native plants are not evolution-
arily adapted to the chemicals released by alien
species and hence fail to establish and ultimately
respond with reduced density, stunted growth
and decreased numbers. Even soil microbes lack
the detoxification and efficient utilization mecha-
nism for these ‘new’ molecules (Callaway and
Aschehoug 2000). Each of these three weeds
exhibits strong allelopathy. Parthenium affects
the emergence, early growth and development of
crops and associated species by releasing pheno-
lic acids and sesquiterpenes from fresh plant
parts (Kanchan and Jayachandra 1980; Kohli
and Batish 1994) and residues (Batish et al. 2002;
Singh et al. 2003). Likewise, Lantana affects
other species by releasing a variety of chemicals
(both volatile and non-volatile) from its fresh
parts, residues and essential oils (Arora and
Kohli 1993; Ambika et al. 2003). Recently, Ager-
atum has also been reported to affect the crops
Table 1. Biotic characteristics of 3 invasive weeds under study.

Feature Lantana Parthenium Ageratum

Life forms Scrambling aromatic shrub Annual herb with a tendency to be Aromatic annual
perennial
Habit Erect in the open and scrambling in Erect Erect
scrubland. Perennial
Plant Height Usually up to 2 m but can even grow to Grows up to 1.5 m, occasionally reaching Grows up to 1 m or more sometimes
5m 2m
Root type Tap root short with a mat of many shal- Tap root Tap root
low side roots
Root length (cm) 15–30 14–18 13–21
Stem Woody, squarish in cross-section and Stem is greenish white, hairy, longitudi- Young stems pubescent. Often red and has
hairy when young but become cylindrical nally grooved long white hairs
and up to 150 mm thick with age.
Leaf shape Ovate, entire with serrate margin Rhomboidal and dissected with short fine Ovate or rhombic-ovate
hairs (trichomes)
Leaf arrangement Occur in opposing pairs along the stem Alternate Opposite
Number of trichomes in
Stem – 75.0±13.3 32.0±9.3
Leaves – 112.0±10.54 55.5±11.7
Inflorescence and colour Axillary heads; Flowers ranging in colour Capitulum; Flowers creamy-white Capitulum; flowers white or blue-purplish
of flowers from cream to yellow to orange, pink,
purple and red
Flowering period Throughout the year Throughout the year October to April
Flowers per plant 20–40 flowers per head 70–120 per head 60–100 per head
Fruit Fleshy drupe, 3–6 mm in diameter and Cypsela with persistent appendages and Cypsela with aristate pappus
containing 1–2 seeds (1.5 mm long). Fruits style; dark brown to black in colour
mature rapidly and change colour from
dark green to black
Seeds per plant 10,000–12,000 10,000–25,000 8,000–10,000
Source: Ambika et al. (2003); Kohli and Rani (1994); Okunade (2002).
1505
1506
and other native species by releasing a number of
phenolic acids and volatile oils from its leaves
and flowers (Kong et al. 2002; Singh et al. 2003).
Under environmental stress, these weeds release
greater amounts and types of secondary metabo-
lites that provide them extra selective advantage
over the natives, enabling them to colonise
quickly at the cost of natives.
Phytochemistry
These weeds are virtually immune to herbivory
owing to the presence of a wide array of phyto-
chemicals of diverse groups. Leaves of Lantana
are rich in volatile essential oils, phenolic com-
pounds and triterpenoids such as lantadenes and
a number of other phenolic compounds like
aesulin, quercetin, isorhamnetin, fisetin, gossype-
tin, tricin, aesculetin, and dicaffeoyl tartaric acid
(Table 2). These provide the weed a number of
properties such as allelopathy and toxicity. Par-
thenium leaves, on the other hand, contain a
number of phenolic acids, sesquiterpene lactones
like parthenin and coronopilin, alkaloids, sapo-
nins, flavonoids, tannins, and unsaturated sterols
(Table 2). Likewise, a number of phytochemicals
(flavonoids, alkaloids, chromenes, phenolics
and essential oils) belonging to different groups
have been reported in Ageratum (Table 2). Be-
side, it also contains 6,7-dimethoxy-2,2-dimethyl-
chromene, 6-demethoxyageratochromene, 6-vinyl-
demethoxyageratochromene, Ageratochromene,
Endo-borneol, Endo-bornyl-acetate, Ethyl-eugenol,
Ethyl-vanillin, Farnesol, fenchyl acetate, Friede-
lin, germacrene, HCN, Hexadecenoic-acid,
Kaempferol and its glucosides, Linoleic acid, ner-
olidal, and Quercetin and its glucosides (Sharma
and Sharma, 1995; Kong et al. 2002; Okunade
2002). Precocene I and II are the major constitu-
ents of the essential oils of Ageratum and are
insecticidal, and possess anti-juvenile hormonal
activity (Okunade 2002).
Environmental impact of the three weeds
The weeds are hazardous to ecosystems and the
natural environment of the region. They deplete
biodiversity by affecting density, frequency and
dominance of natives. As a result native plant
species fail to compete with them and hence are
significantly reduced. Lantana is one of the most
serious weeds among the top hundred invasive
species. The invasion of lantana in the forest
areas has produced a significant reduction in
density and biomass. Being shade tolerant, the
weed covers the whole understorey of the forest
floors, replacing native herbs and shrubs. Be-
cause of the dense and sprawling growth habit of
Lantana, even forest operations are difficult. In
rangelands, grasslands, and pastures, Lantana
infestation causes a severe fodder scarcity, thus
affecting livestock. It is toxic to livestock and if
its fruits are eaten by children, it causes severe
toxicity and even death. In India, Lantana infes-
tation in sandalwood forests has not only af-
fected the trees but also favours the spread of
sandal spike disease (Holm et al. 1977). Its woo-
dy stem and higher lignin content pose fire haz-
ards, increasing fire frequency.
Parthenium is perhaps the most noxious weed
of all, especially of urban India. It is a quick in-
vader and has invaded almost all accessible land.
It reduces biodiversity and affects landscape and
soil quality. In agricultural fields, the weed de-
pletes nutrients from and releases toxic sub-
stances into the soil. The growth, establishment
and yield of crops are reduced and at some pla-
ces, it is known to cause fodder and food scar-
city. Parthenium is hazardous to human health as
well as to livestock. Health impacts on humans
include contact dermatitis, skin irritation, nausea,
giddiness and respiratory problems like bronchi-
tis and asthma. Parthenium causes acute toxicity
in cattle and milk becomes bitter tasting due to
the presence of parthenin, which is also hepato-
toxic (Kohli and Rani 1994). Its presence in the
cattle milk shows its non-bio-degradability.
Similarly, Ageratum has significant negative
impacts. In arable and cultivated lands, it re-
duces the yields of wheat, corn and rice and
other important crops. The precise figures
regarding the percent loss of yield are not
known, yet it depends upon the intensity of
infestation of weed. Roder et al. (1998) have re-
ported that yield of rice is negatively associated
with the density of Ageratum. Farmers in the
lower Shivalik ranges of the Himalayas had even
abandoned their fields because Ageratum had
ruined their croplands. Its strong and pungent
Table 2. Major chemical constituents of Lantana, Parthenium and Ageratum and their biological activity.
Chemical nature
Essential oils (leaves)
Phenolic compounds
Sequiterpene lactones
Triterpenoids
Source: Ambika et al. (2003); Hart et al. (1976); Kohli and Rani (1994); Kong et al. (2002); Okunade (2002); Saleh (1974); Vidya (1994); Xuan et al. (2004).
Weed Chemical constituents Biological nature
Lantana p-cymene, a-phellandrene, a-pinene, diterpene, c- Provide allelopathic property to
terpinene, caryophyllene, cardinene, cineole, lin- plant; used as antiseptic against
alool, geraniol, a-terpinol, citral, phellandral wounds
Parthenium – –
Ageratum Precocene I and II, b-caryophyllene, a-bisabo- Allelopathic, anti-inflammatory,
lene, a-cubebene, a-farnesene, b-farnesene, antimicrobial
b-cubebene, germacrene, eupatoriochromene,
fenchyl acetate, sesquiterpene hydrocarbon, ner-
ol, caryophyllene oxide, bornyl formate, neroli-
dal, b-gurijunene, copane, camphene, b-elemene,
methanezulene, tricycoundecane, a-muurolene,
demethoxy-en-cecalin, audorenececalinol
Lantana (leaf, root and fruit) p-coumaric, caffeic, vanillic, o-coumaric, gentisic, Allelopathic
ferulic, syringic, salicylic, trans-cinnamic, proto-
catechuic, and p-hydroxybenzoic acid; quercetin,
aesulin, isorhamnetin, tricin, gossypetin, aescu-
letin, dicaffenyl tartaric acid
Parthenium (leaf, inflorescence, roots) caffeic, p-coumaric, gallic, p-hydroxybenzoic, Allelopathic
anisic, vanillic and ferulic
Ageratum (leaf, stem, root) Gallic, coumalic, catechin, p-coumaric, sinapic, p- Allelopathic
hydroxybenzoic, benzoic, and protocatechuic
acid
Lantana – –
Parthenium Parthenin, coronopilin, hymenin Allelopathic, toxic to cattle and
man, antiallergenic, autotoxic,
antibacterial, nematicidal, fungi-
cidal
Ageratum – –
Lantana (leaves) Lantadene A and B Toxic and allelopathic
Parthenium – –
Ageratum – –
1507
1508
smell causes giddiness and nausea as well as
allergic reactions in humans.
Management strategies
A number of physical, biological, chemical and
ecological methods have been tried in the past,
but none has succeeded so far. Manual re-
moval of the weeds like Parthenium and Agera-
tum is not feasible because of the enormity of
the area involved, multiple life forms (variabil-
ity in the growth stages from seedling to ma-
ture to seed setting stage) in the same area,
dangers of health problems, and the likelihood
of re-emergence. For heavily infested areas, cut-
ting with swords or use of shrub-masters has
also been tried at some places. When piles of
these plants are either burned or allowed to
decompose, the soil is contaminated. Labour
intensive removal methods harm the health of
workers and the weeds quickly regenerate from
the root stumps.
A number of synthetic herbicides such as atra-
zine, alachlor, paraquat, glyphosate, simazine,
2,4-D and 2,4,5-T have also been tried for the
management of these weeds. The environmental
dangers these herbicides pose on one hand, while
the plants regenerate from the underground parts,
like root stumps, stolons or suckers of the weeds,
on the other, make these efforts futile. Seeds in
the soil also germinate and thus the plants soon
emerge and colonize. Excessive dependence on
the herbicides has produced more resistant weeds.
Among the various biological methods, a num-
ber of pests, insects and fungal pathogens were
tried in the past, but none has provided a satis-
factory solution to this problem. Among various
biological methods, the use of Mexican beetle
(Zygogramma bicolorata) for the control of Par-
thenium caught the attention of the scientists in
the 1990s but unfortunately attacks on non-tar-
get species resulted in termination of the project.
Biological control of lantana included a number
of insects such as the lantana lace bug (Teleon-
emia scrupulosa) and fungal pathogens like rust
fungi (Prospodium tuberculatum) (Thomas and
Ellison 2000), but their effects on the non-target
plants and the failure in providing effective eradi-
cation have limited their use.
Eco-friendly strategies, such as ecological ap-
proaches through antagonistic plants like Cassia
sericea (Joshi 1991) or natural plant products like
essential oil from Eucalyptus for the control of
Parthenium (Kohli et al. 1998) are needed to
control these weeds. Joshi (1991) reported that
Cassia sericea has a good antagonistic effect
against Parthenium and can replace it from the
invaded areas. However, its success is limited and
has only been reported from one Indian state,
Karnataka. Eucalypt oils hold a great potential in
suppressing the emergence, growth and biomass
accumulation in Parthenium (Singh et al. 2005).
Integrated approaches following different meth-
ods coupled with proper land management and
community participation can effectively control
these weeds. In case of Parthenium, which has in-
fested large areas of India, an integrated ap-
proach coupled with community participation has
been quite successful. The area infested with the
weed is once cleared either with herbicide treat-
ment or manual uprooting followed by subjecting
it to use does not let Parthenium invade again.
The area cleared if managed by growing horticul-
tural plants or even native grasses, the weed den-
sity is severely reduced and it can be controlled.
This Herculean task is not possible without the
participation of the local community. For this
educating the masses assumes importance. The
success of this approach has been evidenced
through community participation by motivating
and educating people about the biology and ecol-
ogy of the weed and putting the land to use after
complete removal of the weed (Batish et al. 2004).
Acknowledgements
This work was carried under the B.P. Pal Envi-
ronment Fellowship Award of the Ministry of
Environment and Forests, Government of India,
to R.K. Kohli. The financial assistance of the
Ministry of Environment and Forests for this
award is gratefully acknowledged.
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