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Scholars
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African Journal of Environmental Economics and Management ISSN 2375-0707 Vol. 11 (1), pp. 001-012, 
January, 2023. Available online at www.internationalscholarsjournals.org © International Scholars Journals 

 

Author(s) retain the copyright of this article. 
 
 
 

 

Full Length Research Paper 

 

Effects of Lantana camara Invasion on Riparian 

Vegetation in Uttarakhand: A Case Study from 

the Garhwal Himalayas 
 

Parveen Kumar Dobhal1,2*, Ravinder Kumar Kohli2 and Daizy Rani Batish2
 

 
1
Department of Botany, Government Degree College, Dakpathar, Vikasnagar, Dehradun – 248125, India. 

2
Department of Botany, Panjab University, Chandigarh - 160014, India. 

 
Accepted 17 September, 2023 

 
Lantana camara L. (Verbenaceae), an exotic from tropical America, has invaded vast areas of forests 
surrounding Nayar River in Garhwal Himalayas (Uttarakhand, India). Several factors like high 
reproductive potential, absence of preferred predators, evergreen nature and sufficient moisture 
provided by river water favour extraordinary growth of L. camara in this area. In this study, the impact of 
L. camara invasion on basal area cover, density, frequency and abundance of various plant species was 
determined. Invasion was able to change the quality (composition, distribution) and quantity (growth in 
number and size) of different species in this region. The invasion was also found to have some relation 
with native-exotic nature and different plant habits (like tree and herbs) of local flora. As determined by 
various ecological indices, there was significant loss of species richness and diversity in invaded 
localities. In total, there was a 28.4% decrease in species richness of invaded localities. Excluding L. 
camara, nearly 63% loss of basal area of vegetation was recorded in the invaded localities compared to 
not invaded ones. There was also an impression that L. camara favoured exotics over endemic species 
in this riparian zone. 

 
Key words: Invasion, Lantana camara, riparian, species richness, species diversity, Garhwal Himalaya. 

 
INTRODUCTION 

 
There is a plethora of biological invasions across the blue 

planet in the 21
st

 century. They affect ecological 

processes, which sooner or later contribute to the loss of 
biodiversity from native ecosystems (Vitousek, 1988; 
Baret et al., 2004; Lodge et al., 2006; Pauchard and 
Shea, 2006). After habitat destruction, this is the second 
most extensive threat to global biodiversity on continents, 
and second to none in the case of islands (Sharma et al., 
2005a). Risk is not only restricted to biodiversity loss, as  
invasive alien species (IAS) also threaten the environment,  
 
 
 
*Corresponding author. E-mail: dobhalp1@yahoo.co.in, 

kumar.prv1@rediffmail.com. 
 
Abbreviations: IAS, Invasive alien species. 

 
 
 

 
economies, and human welfare (Lodge et al., 2006). 
Therefore global efforts are being made tocontrol these 
invaders.  

In India, a large number of exotics are naturalized, 
affecting the distribution of native flora. A few among 
them have conspicuously altered the vegetation patterns 
of the country. Reddy (2008), reported 173 IAS from 
India, 80% introduced from the neotropics. The most 
prominent invasive species include Ageratum conyzoides 
L., Chromolaena odorata L., Eupatorium adenophorum 
Spreng, Lantana camara L., Mikania micrantha Kunth, 
Mimosa invisa (Mart.) Solms, Parthenium hysterophorus 
L. and Prosopis juliflora (Sw.) DC. among terrestrial 
plants, and Eichhornia crassipes (Mart.) Solms and Pistia 
stratiotes L., among aquatic IAS. All these species are 
principal threats to the native flora (Sharma et al., 2005b;  
Hajra and Negi, 2007). P. hysterophorus and A. conyzoides 



2 

 

 
 
 

 

are mainly invaders of agroecosystems and pastures 
throughout India (Kohli et al., 2006; Javaid et al., 2009); 
C. odorata , E. adenophorum, M. micrantha are mainly 
invaders of forest, pastures and wastelands of Himalayan 
region, north-east and south-east India (Ramakrishnan, 
1991; Kunwar, 2003; Muniappan et al., 2005). Specifically 
in the North-Western Himalayas, invasive plants such as 
L. camara, P. hysterophorus and A. conyzoides are the 
most problematic weeds (Kohli et al., 2004; Dogra et al., 
2009a, b). Amongst these L. camara is considered as 
one of world’s top 100 invasive species and top 10 worst 
weeds of the world (Sharma et al., 2005a; IUCN, 2009). It 
is a major invader of forests, pasture and wastelands 
throughout the India (Dobhal, 2010). The success of L. 
camara may be attributed to large number of fruits per 
plant (Kohli et al., 2006; Dobhal, 2010), its ability to grow 
under a wide range of climatic conditions (Day et al., 
2003), allelochemicals released by roots in the soil 
inhibiting the growth of neighbouring plants (Ambika et 
al., 2003; Yadav et al., 2004), and possession of some 
feeding deterrents that probably render them less 
acceptable to generalist herbivores than non-invasive 
plants (Jogesh et al., 2008; Sharma et al., 2007). Its 
dominance in invaded areas and possession of 
poisonous components like triterpenes leads to forage 
scarcity and bad health or deaths of animal (Morton, 
1994; Ghisalberti, 2000; Sharma et al., 2007). This 
species also possess ability to change socio-economic 
scenario of invaded regions. In India, its invasion is 
associated with estimated loss of US$ 924 million per 
year (Pimentel et al., 2000). This IAS also dominates 
traditional rain fed agriculture areas of mountainous 
Himalaya and the interest of the farmers towards 
agriculture is declining rapidly (Maikuri et al., 2001). 
These left out terraced agricultural fields, orchards and 
nurseries appears even more intensely dominated by this 
weed. Although it is well known that L. camara is a 
serious invader of various terrestrial ecosystems, there 
are few studies which emphasize the magnitude of the 
influence of this weed on the biodiversity of native 
ecosystems (Sharma et al., 2005a). For example, 
Sharma and Raghubanshi (2007) reported this invasive 
as being responsible for differential depletion of native 
trees. In Himalayas, this was found to be associated with 
marked decreased species density, frequency, 
abundance and basal area and biomass of other species 
of invaded community (Bhatt et al., 1994; Dobhal et al., 
2009). The explicit impact of L. camara invasion in 
riparian zones is also not reported, particularly in India. 
To fill this gap, a study was conducted in the Pauri-
Garhwal region of Uttarakhand, the Himalayan state in 
Northern India. Here, large areas surrounding the River 
Nayar are invaded by this weed. It was hypothesized that 
invasion of L. camara led to degradation of typical 
structure of valuable flora in this riparian ecosystem in 
terms of richness, diversity and native /exotic composition 
of species. 

 
 
 
 

 

STUDY AREA 
 
The study was conducted in the Pauri Garhwal (29°45´-
30°15´N Latitude and 78°24´ -79°23´E Longitude), a 
district of Uttarakhand state, situated in Northern India. At 
an altitude of 600 m asl, running amid two ranges of 
lesser Himalaya (Dudhatoli - Pauri range in the north-east 
and Lansdowne - Mussoorie range in the south-west), 
Nayar is a major tributary of the River Ganges (Figure 1). 
The characteristic riparian vegetation of this region is 
dominated by Acacia catechu Willd. and Dalbergia sissoo 

Roxb. trees with associated shrubs and herbs such as 
Carissa opaca Stapf ex Haines, Barleria cristata L., 
Cissampelos pareira L., Ichnocarpus frutescens (L.) R. 
Br., Justicia adhatoda L., Launaea acaulis (Roxb.) 
Babcock ex Kerr., Murraya koenigii (L.) Spreng., Sida 
cordata (Burm. f.) Borss. Waalk., Ziziphus mauritiana 
Lam. There is heavy invasion of L. camara on both sides 
of the Nayar River. 

 

METHODS 
 
Data collection 
 
The study area was divided into 3 sites; each covered a stretch of 
approximately 2.5 - 3 km along the river (Figure 1). At each site 4 

quadrats, each measuring 4 × 4 m
2
 were laid randomly in L. camara 

invaded localities. A corresponding “control quadrat” was laid in 
nearby non-invaded localities at 50 m away from each of these 
quadrats. In order not to miss any species, these studies were 
continued in four seasons namely spring (February to April), 
summer (May to July), monsoon (August to October) and winter 
(November-January). In this manner 96 quadrats were sampled 
during a year. In each quadrat, plants of each species were 
counted; basal area of each plant was calculated using a screw 
gauge for thin vegetation and vernier caliper or meter tape for thick 
stems. However, when there was high density of individuals of a 
particular herb species (e.g. grasses) and it was extremely difficult 
to separate each of them, the mean density of individual in 12 sub-

quadrats was calculated. All these sub-quadrats were 100 cm
2
 

each and arranged in a random systemic design (Barbour et al., 

1999) inside the larger quadrat measuring 1600 cm
2
. This mean 

density when multiplied with total area of larger quadrat gave total 
number of individuals present inside. In other case when there were 
100 individuals of a species in a quadrat, a mean basal area for 
randomly chosen 100 individual of that species from that specific 
quadrat was calculated and used as a representative basal area for 
that species in that particular quadrat. 

 
Data analysis 
 
The collected plants were identified to species level with the help of 
available floras Duthi (1911), Gaur (1999) and online annotated 
checklist of flora of Nepal available online at www.efloras.org. For 
comparison with type specimens, the herbarium of Botany 
Department, Panjab University and Botanical Survey of India 
(B.S.I.), Dehradun were also consulted. Species were characterized 
on the basis of their exotic-native status with the help of “flora of the 
District Garhwal North-West Himalaya” (Gaur, 1999) and 
“Germplasm Resources Information Network” (GRIN, 2008) an 
online database. Species were also categorised on the basis of 
their growth forms viz. climber, herb, shrub, tree and under-shrub. 

Quantitative analysis of plants for per cent basal area, 



3 

 

   
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 

Figure 1. Location map of study area showing different sites of study. 
 

 
abundance, density, frequency and importance value index (IVI) 
was done as per Misra (1968) and Ambasht (1990) . Qualitative 
analysis in terms of richness, diversity and evenness indices was 
done using different methods. Species richness was calculated 
according to Margalef (1958), Menhinick (1964) and Peet (1974) as 
given in Sagar and Singh (1999). Diversity was calculated in terms 
of Fisher's diversity index (Fisher et al., 1943), Shannon's index H' 
(Shannon and Weaver, 1963), Simpson's index (Simpson, 1949), 
Hill's number N1 and N2 (Hill, 1973), using Biodiversity Pro Version 
2.0 (McAleece, 1997). Distribution of diversity was calculated in 
terms of Evenness number E1 (Pielou, 1969) and E2 (Sheldon, 
1969) calculated per the method given in Sagar and Singh (1999), 
using the applicable ecological software package from Ludwig and 
Raynold (1988). Significance of difference between data was 
determined by Tukey-test using SPSS software. 

 

RESULTS 
 
A total of 89 plant species were recorded from the study 
area, of which 81 were found in non- invaded localities, 
58 in L. camara invaded localities and 50 were common 
to both localities (Table 1).  

There were 31 species (Table 1) found exclusively in 
non-invaded localities or absent from invaded localities. 
Standing out against these 31 sensitive species there 
were just 8 species, namely Chloris dolichostachya, 
Jatropha curcas, Neyraudia arundinacea, Phyllanthus 

 
 

 

virgatus, Ricinus communis, Saccharum spontaneum, 
Sida acuta and Woodfordia fruticosa which occurred 
exclusively in invaded areas, and may therefore be 
considered more resistant to L. camara invasion 
compared to all other species found in the study area. In 
comparison to non-invaded sites there was 28.4% 
reduction in species richness in invaded sites. Among 
species which were common to invaded and non-invaded 
localities Cyanotis vaga, Phyllanthus urinaria, Boerhavia 
diffusa and Vallaris solanacea were most severely 
influenced by invasion. Their IVI values in invaded 
localities were reduced respectively, to 25.1, 34.5, 43.0 
and 15.7% of non-invaded localities. The other species 
demonstrating strong negative impacts of L. camara 
invasion were: Oxalis corniculata and Murraya koenigii 
throughout the year; Cissampelos pareira and Launaea 
acaulis during spring; Gnaphalium hypoleucum, 
Geranium nepalens, Vallaris solanacea and Aerva 
sanguinolenta during summer; and Cyanotis cristata, 
Physalis minima, Dioscorea belophylla, Parthenium 
hysterophorus and Aerva sanguinolenta during monsoon. 
The greatest number of species shows negative impacts 
during the winter, including Mallotus philippensis, Aerva 
sanguinolenta, Securinega virosa, Ageratum conyzoides, 
Barleria cristata, Parthenium hysterophorus, Ipomoea 



4 

 

 
 
 

 
Table 1. Inventory of plants in the study area in decreasing order of their IVI in Lantana camara invaded localities followed by species found exclusively 

in non-invaded localities.  
 

S/No. Botanical name (Family) and habit Annual mean of importance value index Native and exotic 
 

  Invaded localities Non-invaded localities status* / native range 
 

1. Lantana camara L. (Verbenaceae) / S 89.43 ± 9.29 4.10 ± 1.08 E / Tropical America 
 

2. Murraya koenigii (L.) Spreng. (Rutaceae) / S 39.12 ± 12.18 65.08 ± 10.74 N / India, Asia 
 

3. Ziziphus mauritiana Lam. (Rhamnaceae) / S 18.94 ± 4.26 17.08 ± 2.97 N / India 
 

4. Dalbargia sissoo Roxb. (Fabaceae) / T 14.54 ± 2.77 14.99 ± 6.14 
N / India, Pakistan, 

 

Afghanistan, Myanmar  

    
 

5. Acacia catechu Willd. (Mimosaceae) / T 14.05 ± 0.08 15.92 ± 3.69 N / India, Malaysia 
 

6. Carissa opaca Stapf ex Haines (Apocynaceae) / S 10.57 ± 1.73 10.47 ± 2.53 
N / India, Pakistan, 

 

Sri-Lanka, Myanmar  

    
  

7. Euphorbia hirta L. (Euphorbiaceae) /H  
8. Cassia occidentalis L. (Caesalpiniaceae) / U-S  
9. Parthenium hysterophorus L. (Asteraceae) / H  

10. Eragrostis tenella (L.) P. Beauv. ex Roemer & Schultes 
(Poaceae) / H  

11. Ipomoea hederifolia L. (Convolvulaceae) / C  
12. Saccharum spontaneum L. (Poaceae) / H  
13. Oxalis corniculata L. (Oxalidaceae) / H 

 
14. Corchorus aestuans L. (Tilliaceae) / H 

  
8.64 ± 0.14 4.11 ± 2.09 N / Pantropics 

 

7.56 ± 3.92 3.03 ± 1.59 E / Tropical America 
 

6.98 ± 2.43 10.15 ± 5.77 E / Tropical America 
 

6.23 ± 1.66 1.36 ± 0.46 
N / India, Tropical and 

 

Temperate Asia, Africa  

  
 

6.16 ± 4.75 4.48 ± 3.82 E / America 
 

6.11 ± 4.87 ---- N / India, Pantropics 
 

5.03 ± 1.17 16.98 ± 8.03 O / Cosmopolitan 
 

4.39 ± 5.07 2.35 ± 4.70 
O / Neotropics 

 

(probably)  

  
  

15. Ageratum conyzoides L. (Asteraceae) / H 

 

16. Aerva sanguinolenta (L.) Blume (Amaranthaceae) /H 

 

17. Cissampelos pareira L. (Manispermaceae) / C 
 

18. Pyrus pashia Buch. - Ham. ex D. Don (Rosaceae) / T 

 

19. Ichnocarpus frutescens (L.) R. Br. (Apocynaceae) / C 

 

20. Celtis australis L. (Ulmaceae) / T  

21. Launaea acaulis (Roxb.) Babcock ex Kerr (Asteraceae) /  
H 

 
22. Phyllanthus virgatus G. Forst. (Euphorbiaceae) / H 

 

23. Martynia annua L. (Martyniaceae) / H 
 

24. Sida cordata (Burm. f.) Borss. Waalk. (Malvaceae) / H  
25. Cassia tora L. (Caesalpiniaceae) / U-S 

 
26. Neyraudia arundinacea (L.) Henrard (Poaceae) / H 

 
27. Trichosanthes cordata Roxb. (Cucurbitaceae) / C  
28. Solanum incanum L. (Solanaceae) / S 

 
29. Capparis zeylanica L. (Capparaceae) / S 

 
30. Boerhavia diffusa L. (Nyctaginaceae) / H  

31. Securinega virosa (Roxb. ex Willd.) Baillon 
(Euphorbiaceae) / S 

 
32. Woodfordia fruticosa (L.) Kurz (Lythraceae) / S 

 

33. Triumfetta rhomboides Jacq. (Tiliaceae) / U-S 

 
 

4.38± 1.24 8.15± 6.87 

3.99 ± 2.09 6.82 ± 2.19 

3.82 ± 1.17 5.54 ± 1.29 

3.38 ± 0.50 1.02 ± 0.32 

3.26 ± 0.61 1.71 ± 1.70 

3.16 ± 0.43 2.98 ± 0.84 

2.57 ± 1.34 2.13 ± 0.46 

2.47 ± 1.70 ---- 

2.46 ± 2.84 1.91 ± 2.59 

2.45 ± 0.84 2.56 ± 1.53 

2.28 ± 2.63 1.17 ± 1.19 

2.19 ± 2.12 ---- 

1.80 ± 2.08 0.78 ± 1.56 

1.72 ± 1.17 2.21 ± 0.18 

1.69 ± 0.46 1.41 ± 0.34 

1.38 ± 0.84 3.21 ± 5.95 

1.24 ± 0.25 1.60 ± 0.49 

1.21 ± 0.23 ---- 

1.12 ± 1.29 0.46 ± 0.93  

 
E / Tropical America  
N / India, Bhutan, 
China, Myanmar, 

Malaysia, Nepal  
E / America  
N / India, Bhutan,  
Myanmar  
N / India, Sri-Lanka,  
Thailand, Indonesia  
E / Africa, South-  
Europe, Turkey  
N / India, Afghanistan,  
Pakistan  
N / India, Sri-Lanka,  
China  
E / North America,  
Mexico  
E / America  
E / South America  
N / India, Tropical-Asia, 

Africa  
N / India, Myanmar 

N / India, Africa  
N / India, Tropical 

and Temperate Asia  
O / Widely Naturalized 
 
N / India, Asia, Africa 
 
N / India, Burma,  
China, Africa  
N / India, China,  
Malaysia, Africa 



5 

 

 
 
 

 
Table 1 Contd.  
 

 

34. Phyllanthus urinaria L. (Euphorbiaceae) / H 1.09 ± 0.85 3.16 ± 5.98 

35. Vallaris solanacea (Roth) Kuntze (Apocynaceae) / C 1.04 ± 0.71 6.64 ± 1.58 

36. Chloris dolichostachya Lagasca (Poaceae) / H 1.01 ± 0.70 ---- 

37. Gnaphalium hypoleucum DC. (Asteraceae) / H 1.00 ± 1.15 3.70 ± 6.48 

38. Cyanotis cristata (L.) D. Don (Commelinaceae) / H 0.93 ± 1.07 1.50 ± 2.66 

39. Ipomea cairica (L.) Sweet (Convolvulaceae) / C 0.91 ± 0.64 0.21 ± 0.43 

40. Jatropha curcas L. (Euphorbiaceae) / S 0.91 ± 0.64 ---- 

41. Physalis minima L. (Solanaceae) / H 0.86 ± 0.99 2.04 ± 2.39 

42. Pupalia lappacea (L.) Juss. (Amaranthaceae) / H 0.83 ± 1.66 0.55 ± 1.10 

 
 
N / Indian 

subcontinent, Indo-

china, China, Japan  
N / India, Tropical and 

Temperate Asia  
N / India, Afghanistan, 

China, Myanmar, Sri-

Lanka  
N / Indian Himalayas 

Japan, China, Bhutan  
N / India, Tropical 

Asia, Tropical Africa  
E / Australia, America, 

New-Zealand  
E / Tropical America  
E / Tropical America, 

Africa  
N / India, Pakistan, 

S.E. Asia, Africa, 

Madagascar  

43. Cryptolepis buchananii Roem. & Schult. (Asclepiadaceae) 
/ C  

44. Mallotus philippensis (Lam.) Muell. Arg. (Euphorbiaceae) 
/ T 

 
45. Ampelocissus latifolia (Roxb.) Planchon (Vitaceae) / C 

 

46. Holoptelea integrifolia (Roxb.) Planch. (Ulmaceae) / T 

   

0.72 ± 1.43 1.52 ± 0.55 
N / India, China, 

 

Malaysia  

  
 

0.71 ± 1.42 6.06 ± 1.67 E / Australia 
 

0.67 ± 0.77 0.46± 0.91 
N / India, Indian 

 

subcontinent  

  
 

0.66 ± 1.32 3.79 ± 0.59 
N / India, Nepal, 

 

Myanmar, Sri-Lanka, 
 

  Indo-China 
  

47. Asparagus filicinus Buch.-Ham. ex D. Don 
(Asparagaceae) / S 

 
48. Geranium nepalense Sweet (Geraniaceae) / H 

 

49. Cyanotis vaga (Lour.) J. A. & J. H. Schult. 
(Commelinaceae) / H 

 
50. Cypreus niveus Retz. (Cyperaceae) / H 

 
51. Gomphrena globosa L. (Amaranthaceae) / H  

52. Dioscorea belophylla (Prain) J.O. Voigt. ex Haines 
(Dioscoreaceae) / C 

 
53. Rhus parviflora Roxb. (Anacardiaceae) 

 
54. Peristrophe bicalyculata (Retz.) Nees (Acanthaceae) / H  
55. Sida acuta Burm. (Malvaceae) / H  
56. Euphorbia helioscopia L. (Euphorbiaceae) / H 

 
57. Barleria cristata L. (Acanthaceae) / U-S 

 
58. Ricinus communis L. (Euphorbiaceae) / S 

 
59. Abrus prectoris L. (Fabaceae) / C 

 

60. Anisomeles indica (L.) Kuntze (Lamiaceae) / H 

 

61. Arthraxon prionodes (Steudel) Dandy (Poaceae) / H 
 
62. Asclepias curassavica L. (Asclepiadacea) / H 

 
 

0.65 ± 0.48 1.05 ± 0.63 
N / India, China, Indo- 

 

China, Myanmar  

  
 

0.61 ± 0.44 1.13 ± 1.34 
N / India, Tropical and 

 

Temperate Asia  

  
 

0.46 ± 0.53 1.83 ± 3.65 
N / India, Pakistan, 

 

Burma, Africa  

  
 

0.41 ± 0.48 0.15 ± 0.30 
N / India, S. and S. E. 

 

Asia, Tropical Africa  

  
 

0.34 ± 0.39 0.11 ± 0.22 E / Tropical America 
 

0.33 ± 0.39 0.89 ± 0.36 N / Indian Himalayas 
 

0.32 ± 0.64 0.16 ± 0.32 
N / India, Nepal, Sri- 

 

Lanka  

  
 

0.26 ± 0.52 0.29 ± 0.58 N / India, Nepal 
 

0.26 ± 0.52 ---- E / Africa 
 

0.26 ± 0.52 0.16 ± 0.31 E / Eurasia 
 

0.24 ± 0.47 0.41 ± 0.81 
N / India, Myanmar, 

 

Indo-China  

  
 

0.22 ± 0.44 ---- E / Tropics, Africa 
 

---- 0.97 ± 0.71 
N / India, Old World 

 

Tropics  

  
 

---- 0.65 ± 1.30 
N / India, China, 

 

Malaysia  

  
 

---- 3.00 ± 6.00 
N / India, Tropical and 

 

Temperate Asia, Africa  

  
 

---- 0.17 ± 0.34 E / Tropical America 
  

  



6 

 

 
 
 

 
Table 1. Contd.  

 

63. Atylosia scarabaeoides (L.) Benth. (Fabaceae) / H 

 

64. Brachiaria reptans (L.) Gardner & C. E. Hubbard 
(Poaceae) / H 

 
65. Butea monosperma (Lam.) Kuntze (Fabaceae) / T 

 

 
 

---- 0.10 ± 0.21 

---- 3.97 ± 4.65 

---- 12.62 ± 0.92 

 
 
N / India, China,  
Malaysia  
N / India, Tropical 

and sub-tropical Asia, 

Africa  
N / India, Nepal, 

Pakistan, Malaysia, 

Indo-China  
66. Cannabis sativa L. (Cannabaceae) / H ---- 0.37 ± 0.73 N / India, Central Asia 

 

67. Cassia fistula L. (Caesalpiniaceae) / T ---- 6.84 ± 0.55 N / India, Myanmar 
 

68. Circium verutum (D. Don) Sprengel (Asteraceae) / H ---- 0.41 ± 0.83 
N / India, Pakistan, 

 

Afghanistan  

    
 

 
Coccinia grandis (L.) Voigt (Cucurbitaceae) / C ---- 

 N / India, Tropical and 
 

69. 0.54 ± 1.07 Temperate Asia , 
 

    Africa 
  

70. Dioscorea bulbifera L. (Dioscoreaceae) / C ---- 0.12 ± 0.24 
 

71. Eupatorium adenophorum Spreng. (Asteraceae) / S ---- 0.54 ± 0.37 
 

72. Flacourtia indica (Burm. F.) Merrill (Flacourtiaceae) / S ---- 0.12 ± 0.24 
 

73. Galium asparifolium Wallich (Rubiaceae) / H ---- 0.98 ± 1.96 
 

74. Galium elegans Wall. ex Roxb. (Rubiaceae) / H ---- 2.27 ± 4.54 
 

75. Gloriosa superba L. (Liliaceae) / H ---- 0.21 ± 0.42 
 

76. Grewia optiva J. R. Drumm. ex Burret (Tiliaceae) / T ---- 0.56 ± 0.43 
 

77. Jasminum arborescens Roxb. (Oleaceae) / C ---- 1.19 ± 2.38 
 

78. Justicia adhatoda L. (Acanthaceae) / S ---- 3.21 ± 0.58 
 

79. Lactuca dissecta D. Don (Asteraceae) / H ---- 1.38 ± 2.75 
 

80. 
Malvastrum coromandelianum (L.) Garcke (Malvaceae) / 

---- 0.61 ± 0.71  
H 

 
 

    
 

81. Momordica dioica Roxb. Ex Willd. (Cucurbitaceae) / C ---- 0.45 ± 0.89 
 

82. Naringi crenulata (Roxb.) Nicolson (Rutaceae) / T ---- 2.07 ± 0.38 
 

83. Oenothera rosea L'Herit. ex Aiton (Onagraceae) / H ---- 1.98 ± 3.95 
 

84. Rhamnus virgatus Roxb. (Rhamnaceae) / S ---- 0.20 ± 0.40 
 

85. Saussurea candicans (DC.) Sch. Bip. (Asteraceae) / H ---- 0.63 ± 1.26 
 

86. Solanum nigrum L. (Solanaceae) / H ---- 2.44 ± 1.31 
 

87. Syzygium cumini (L.) Skeels (Myrtaceae) / T ---- 0.40 ± 0.48 
 

88. Toona serrata (Royle) M. Roemer (Meliaceae) / T ---- 0.78 ± 1.55 
 

89. Urena lobata L. (Malvaceae) / U-S  ---- 0.71 ± 0.96 
  

 
N / Pantropics  
E / North America, 

Mexico  
N / India, Tropical and 

Temperate Asia, Africa  
N / India, Myanmar, 

Sri-Lanka  
N / India, Nepal, 

Bhutan, W. China  
N / Tropical Asia, 

Africa  
N / India, Nepal  
N / India, Pakistan, 

Sri-Lanka  
N / India, Indo-China  
N / India, Tropical and 

Temperate Asia  
E / Southern USA, 

Mexico, Central 

America  
N / India, Malaysia, 

Sri-Lanka  
N / Indian 

subcontinent, 

China, Indo-China  
E / America  
N / India, Temperate 

Asia  
N / India  
E / Eurasia  
N / India, Tropical-

Asia  
N / India, Myanmar 

E / Australia 
 

Given after family: C for climber, H for herb, S for shrub, T for tree and U-S for under-shrub; - - - - Species absent; *Status based on native and exotic status 

where; N, E and O stand for Native, Exotic and Obscure status respectively; ± is standard deviation. 
 
 

 

hederifolia, Holoptelea integrifolia and Sida cordata. On 

the other hand, among species which were common to 

 
 
 

 

invaded and non-invaded localities, only a few species 

were growing more luxuriously in invaded localities. 



7 

 

  
 
 

 
Table 2. Mean number of plant species in terms of climber, herb, shrub, tree and under-shrub among species present in non-

invaded, exclusively in non-invaded, commonly in invaded and non-invaded, in invaded and exclusively in invaded localities.  
 

 

Habit type 
In non- 

In Invaded 
Common in Exclusively in Exclusively in 

 

 invaded both non-invaded invaded  

 

localities  

  localities localities localities localities  

   
 

Number of 
Climber 9.33 ± 2.52 6.33 ± 0.58 5.67 ± 1.53 5.50 ± 0.71 1.00 ± 1.41 

 

Herb 24.67 ± 2.08** 13.00 ± 4.36 10.33 ± 4.73 14.33 ± 3.21** 2.67 ± 0.58  

species 
 

Shrub 8.67 ± 2.08 8.33 ± 3.21 6.67 ± 2.08 2.00 ± 0.00 1.67 ± 1.15  

representing plant  

Tree 7.33 ± 2.31* 4.33 ± 0.58 4.33 ± 0.58 3.00 ± 1.73* 0 
 

habit  

Under-shrub 4.00 ± 1.00 2.67 ± 1.53 2.00 ± 1.00 2.00 ± 1.73 0.67 ± 0.58 
 

 
 

Total species  54.00 ± 7.00* 34.33 ± 2.08 29.00 ± 2.65 25.00 ± 9.17* 5.33 ± 0.58 
 

 
*Significantly different at level P < 0.05** and 0.1* from associated L. camara invaded localities, as estimated by t-test , ± is standard deviation. 

 
 

 
Table 3. Percentage of sum of % basal area for all species, sum of frequency for all species, sum of density for all species, sum 

of abundance for all species in invaded compared with non-invaded localities during four seasons.  
 

  Percentage against non-invaded localities  

Season Sum of % basal area Sum of frequencies Sum of densities Sum of abundance 

 covered by all species of all species of all species of all species 

Spring 83.15 70.29 66.25 52.88 

Summer 74.45 89.64 75.25 88.30 

Monsoon 73.04 62.98 45.22 75.96 

Winter 78.42 56.84 49.81 51.75 

Annual mean 77.27 ± 4.537 69.9 ± 14.2 59.1 ± 14 67.2 ± 17.9 
 
 

 

These were Martynia annua and Cassia tora specifically 
during summer; Euphorbia hirta and Cassia occidentalis 
during summer as well as monsoon seasons; and 
Eragrostis tenella and Pyrus pashia throughout the year.  

In terms of plant habit L. camara invasion seems to 
have maximum impact on herbs and trees. In comparison 
to 24.67 ± 2.08 herb species in non-invaded localities 
there were significantly less, 13.00 ± 4.36 species in L. 
camara invaded localities (Table 2). Similarly, compared 
to 7.33 ± 2.31 tree species in non-invaded localities there 
were significantly less (4.33 ± 0.58) species in L. camara 
invaded localities. The trend continued among those 
species which were exclusive to either invade or non-
invaded localities. There were fewer herbaceous and no 
tree forms among species specific to invaded localities in 
comparison to species present specifically in non-invaded 
localities.  

The values for invaded localities in terms of sum of per 
cent basal area (secondary growth of stem), sum of 
frequencies (probability of finding any species), sum of 
densities (overall density) and sum of abundance (local 
density) of all species were very low compared to non-
invaded sites (Table 3).  

The negative impact of L. camara on local flora is 

illustrated by the lower percentage values for invaded 
localities in terms of various parameters mentioned above 

varied with different seasons. The annual mean 

 
 

 

value of overall basal area of all plants from invaded 
localities was 77.27 ± 4.54% of non-invaded localities. It 
was lowest (73.04%) during winter and highest (83.15%) 
during spring. The sum of frequencies of all species in 
invaded localities was reduced to 69.9 ± 14.2% of non-
invaded localities (Table 3). 

Thus, with changing seasons there were some large 
fluctuations in impact of L. camara invasion on probability 
of finding any plant species. The maximum negative 
impact was found during winter season, when overall 
frequency of finding any species in invaded localities was 
reduced to just 56.84% compared to non-invaded 
localities. Relatively lesser negative impact was observed 
during summer, when frequency for invaded localities 
was 89.64% of non-invaded ones. Overall density of plant 
species in invaded localities was just 59.1 ± 14% of non-
invaded localities; it was most negatively affected feature 
of plant distribution in invaded localities. The impact on 
density was most severe during monsoon followed by 
winter when overall density of plants in invaded localities 
was reduced to nearly half of corresponding non-invaded 
localities. Similarly, overall abundance of all species from 
invaded localities was very low; it was 67.2 ± 17.9% of 
non-invaded localities (Table 3). 

Among all parameters mentioned above, overall value 

of basal area demonstrated a comparatively lesser 

impact of invasion. However, this was not the case if the 



8 

 

 
 
 

 

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300    
b 

  
 

280 
     

 

      
 

260       
 

240       
 

220       
 

200       
 

180       
 

160       
 

140       
 

120       
 

100       
 

80   
ab 

 ab 
ab 

 

60 ab    
 

40 a 
    

 

     
 

20       
 

 A B C D E F 
  

 

Specific group 

 
 

Figure 2. Percentage of exotics in relation to native species present in non-invaded (A), 
exclusively in non-invaded (B), commonly in invaded and non-invaded (C), exclusively in 
invaded (D), in invaded localities (E) and in whole area under study (F) . Means denoted by 
different alphabets are significantly different at P < 0.1, as estimated by Tukey-test, ± is 
standard deviation. 

 
 

 

basal area of L. camara is excluded from the analysis. In 
invaded localities L. camara had about 62.9 ± 0.86% 
contribution in total basal area of all species. Thus, 
although the total basal area of plants present in invaded 
localities was 77.27 ± 4.54% of non-invaded localities, 
this was mainly due to L. camara. The contribution of 
other plants was very low, that is, 37.42 ± 0.8% of non-
invaded localities. Similarly, L. camara also had 
maximum (29.81 ± 3.1%) share in total IVI for invaded 
localities.  

After incorporating data on the exotic and native status 
of each species found during the present study, L. 
camara was found to have some relation with exotic and 
native status of species. There were 27% exotics among 
the flora of Nayar irrespective of invaded or non-invaded 
localities. However, the percentage of exotics was found 
to vary when plants in this region were divided into 
different subsets on the basis of their presence and 
absence in invaded and non-invaded localities. It was 30, 
19.35 and 50% among species common to invaded and 
non-invaded localities, species specific to non-invaded 
localities and species specific to L. camara invaded 
localities respectively.  

During this survey the overall number of exotics was 

found to be 43.8 ± 8.1% of the native species (Figure 2). 

 
 
 

 

The exotic to native species ratio was particularly high 
(60.3 ± 7.2%) in the invaded localities, compared to (39 ± 
9.6%) for non-invaded localities. The share of exotics 
among those species which were present exclusively in 
non- invaded localities was 26 ± 4.4% of native species. 
This was very low in comparison to overall per cent of 
exotics against natives. However, among those species 
which were restricted exclusively to L. camara invaded 
localities there were extremely high percentage (155.5 ± 
126.2) of exotics in relation to native species. The exotic 
to native ratio were significantly different in case of 
species those were exclusive to non- invaded localities 
and those were restricted to invaded localities. Exotic 
species had 63.78% share in total basal area cover of 
invaded localities. It was extremely high in comparison to 
3.55% for non-invaded localities. The percent share of 
exotics in overall frequency of occurrence of plant 
species in invaded localities (33.82%) was also higher 
compared to 26.83% for non-invaded localities (Figure 3). 
Exotics also constituted a higher percentage of total 
density of plants in invaded localities, although the trend 
was reversed in the case of abundance. The percentage 
of exotics in total IVI of invaded localities (44.38), that 
was nearly equal to per cent share (52.02) of non-
invaded localities, also indicated that exotics were 



9 

 

   
 
 
 

 
P

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r 

c
e

n
t 

s
h

a
re

 

v
a
lu

e
 

m
ea

n 

of
in

to
ta

l 

A
n

n
u

a
l 

 

 
 
 

 
 

100 
* 

   
Exotic Native Obsecure 

 

90 
   

 

         
 

80         
*  

         
 

70       
* 

  
 

60 
        

 

         
 

50          
 

40          
 

30         
* 

 

20 
        

 

         
 

10 
* 

        
 

0 
        

 

         
 

I. N.I. I. N.I. I. N.I. I. N.I. I. N.I. 
 

Basal area Frequency Density Abundance I.V.I. 
 

Parameters under study  

in Lantana camara invaded (I.) and non-invaded (N.I.) localities 
 

 
Figure 3. Percentage of species with exotic, native and obscure origin, in terms of basal area, frequency, density, abundance 

and IVI. Mean values for invaded localities with asterix (*) above bar are significantly different from corresponding non-invaded 

localities at P < 0.05, as estimated by Tukey-test, ± is standard deviation. 
 
 
 
 

occupying very important places in L. camara invaded 
localities (Figure 3).  

The species richness and diversity of invaded localities 
was very low in comparison to non- invaded localities. 
The difference was extremely high and significant in 
terms of species richness (Table 4). In terms of diversity, 
however, there was not a very large difference between 
the two compared localities. Despite this, the diversity of 
invaded localities was significantly lower than that of non-
invaded localities in terms of Fisher's diversity index ( ), 

Shannon’s index (H'), and Hills number (N1). The 
difference was neither very large and nor statistically 
significant in terms of Simpson's index ( ) and Hills 

number (N2). Despite having lower values of diversity, the 
invaded localities had nearly equally even distribution of 

diversity (as shown by evenness E1 and E2) to that of 
non-invaded localities (Table 4). 
 

 

DISCUSSION 

 

L. camara has serious deleterious effect on some of 

endemic animal (Morton, 1994; Sharma et al., 2007) and 
plant species. It is also known to displace natural scrub 
communities as well as prevent natural regeneration of 
some tree species (Ambika et al., 2003; Sharma and 

Raghubanshi, 2006; Dobhal, 2010). In Shivalik hills in 

 
 
 

 

northwestern Himalayan range of India, it was observed 
that the diversity, evenness and richness of the native 
species were drastically reduced in the forest invaded by 
the exotics (Dogra et al., 2009a,b). Similar patterns were 
observed in the case of riparian forests along the Nayar 
River, where a decrease of nearly 30% in species 
richness was recorded in L. camara invaded localities. 
Compared to non-invaded localities there were 
significantly fewer herb and tree species in L. camara 
invaded localities. Specially, there were no tree species 
among species found exclusively in invaded localities. 
Plant species with specific root and shoot distribution 
habits in relation to L. camara may come across different 
level of resource competition and allelopathic inhibition 
offered by this invasive (Dobhal et al., 2010). This could 
also be a plausible reason for observed variation in 
impact of L. camara invasion on different life forms.  

Anthropogenic and natural disturbances act together to 
facilitate the introduction and spread of exotic species in 
riparian zones. Further, the availability of moisture and 
the dispersal of propagules by water may increase 
invasibility of riparian zones (Hood and Naiman, 2000; 
Pysek and Prach, 1994) . In riparian zones between hill 
slopes, such as in Nayar region where more than 25% of 
total species were exotics, there can be concentrated 
energy and material flow during flooding and other 
geomorphic disturbances and riparian forest communities 



10 

 

 
 
 

 
Table 4. Values of different parameters associated with richness and diversity in L. camara invaded and non-invaded localities.  

 
Locality type 

Parameters associated with species richness and diversity and distribution  
Non-invaded L. camara invaded  

 
 

Richness 
 
 
 
 

 

Diversity 
 
 
 

 
Distribution of diversity (Evenness or equitability) 

  
 

Number of species (N0) 31.50 ± 6.24** 22.00 ± 4.07 

Peet index (P) 842.35 ± 291.59** 451.47 ± 165.01 

1. Margalef index (R1) 3.50 ± 0.53** 1.11 ± 0.14 

2. Menhinick index (R2) 4.68 ± 1.22* 0.78 ± 0.16 

Fisher's diversity index ( ) 6.87 ± 1.30** 5.05 ± 0.84 

Shannon’s index (H') 2.61 ± 0.32** 2.36 ± 0.32 

Simpson's index ( ) 0.12 ± 0.05 0.14 ± 0.06 

Hills number N1 14.29 ± 4.42** 11.07 ± 3.56 

Hills number N2 9.91 ± 3.71 8.11 ± 3.19 

Evenness E1 0.76 ± 0.07 0.77 ± 0.07 

Evenness E2 0.45 ± 0.09 0.50 ± 0.10   
*Significantly different at level P < 0.05** and 0.1* from associated L. camara invaded localities, as estimated by t-test, ± is standard deviation. 

 
 

 

use material trapped by these disturbances (Junk et al., 
1982; Nakamura and Inahara, 2007). In this region 
exotics had a higher percentage share in total density of 
plants in invaded localities, but the trend was reversed in 
the case of abundance. Higher percentage values of 
exotics in total abundance of non-invaded localities 
indicated that in non-invaded localities, exotics were 
mostly restricted to small abundant patches, whereas 
they were distributed densely throughout in the area 
invaded by L. camara. Further, there was a significantly 
higher proportion of exotics among species found 
exclusively with L. camara against species present 
exclusively in non-invaded localities, indicating that L. 
camara facilitates the invasion of other exotics. It appears 
that L. camara invasion sweeps over native flora and 
promotes the establishment of other exotic species. Thus, 
there was a possibility that in near future exotics may 
cover larger areas of the region in comparison to 
endemics, as L. camara infestation continues increasing 
without any barrier. The increase in the ration of exotic to 
endemic species in Nayar region becomes crucial as this 
region is a part of Garhwal-Kumaon Himalayas 
(Uttarakhand), one of 25 micro-endemic centers in India 
(Subramani et al., 2005).  

Basal areas and density of plants of a species, 
respectively, provide simple indicators of growth and 
survival rates of a species. In invaded areas there was 
lower basal area of plants, of which L. camara had most 

of the share; besides this there was decrease in the 
overall number of species, the number of individuals per 
species, and overall density. There were also lower 
chances of frequent occurrence of any species in invaded 
localities compared to associated non-invaded ones. In 
total there was significantly lower richness and diversity 
and overall health of plants in invaded localities 

 
 
 

 

compared to non-invaded ones. This may be attributed to 
thick monoculture of L. camara that alter the 
microenvironment (light, pH and temperature) beneath 
the L. camara thickets, and inhibit either germination or 
growth of other species (Sharma and Raghubanshi, 
2007). 

Although riparian zones can act as a buffer against fire 
and act as a refuge for fire-sensitive species, under some 
circumstances, like dry climate and the accumulation of 
dry fuel, riparian areas become corridors for fire 
movement. Fires in riparian zones create canopy gaps 
and drier conditions, which allow establishment of fire 
adapted species (Pettit and Naiman, 2007). Increased 
dominance of a fire loving species like L. camara in such 
an important buffer zone may have far reaching 
consequences on native riparian ecosystem. There may 
be a positive feedback between fires and invasion by L. 
camara, leading to a fire L. camara cycle that can have 
deleterious compositional and functional consequences 
for forest ecosystems (Hiremath and Sundaram, 2005, 
Dobhal et al., 2009). Being a complex process it is 
difficult to evaluate the effect of an invader on fire 
regimes (Strayer et al. 2006). However, reports of 
significant increase in L. camara seed germination in 
smoke (Raizada and Raghubanshi, 2010) and enhanced 
prolification after fire (Gentle and Duggin, 1997; Dobhal et 
al., 2009) supports the view given above. 
 

 

Conclusions 

 

L. camara poses a threat to local flora. The number, 

density and frequency and overall health of species were 

remarkably poor in invaded areas. Though it is premature 
to say, it seems that L. camara favours exotics over 



11 

 

 
 
 

 

endemic species. However, more studies are required to 
further investigate this relationship. Further, due to its 
inflammable nature and dominance in regional flora, L. 
camara is a potential threat to riparian zones that can act 

as a buffer against fire and act as a refuge for fire-
sensitive species. Consequently, beside individual 
species and life forms such as trees and herbs, the entire 
forest ecosystem itself is under threat due to L. camara 
invasion. It requires the immediate committed attention of 
ecologists, conservationists and policy makers. 
 

 

ACKNOWLEDGEMENTS 

 

Authors are thankful to University Grant Commission 
(India), for providing financial assistance. We also admire 
the help provided by Botanical Survey of India 
(Dehradun) and Panjab University (Chandigarh) in 
identification of various plant species. Thanks are also 
due to the anonymous reviewers, for their valuable 
suggestion for improving overall presentation, particularly 
the language of this manuscript. 
 

 
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