Research Article
Research Article
Lantana camara L. (sensu lato): an enigmatic complex
expand article infoNeha Goyal, Gyan Prakash Sharma
‡ University of Delhi, Delhi, India
Open Access


Lantana camara L., considered among the world’s worst invaders is in identity crisis and contentiously referred as Lantana camara L. (sensu lato). Taxonomic ambiguity in L. camara L. (sensu lato), a species complex is one of the grim caveats behind incompetence of its management efforts. Recognizing the extent of variability within the complex, we aim to highlight the need to circumscribe its composition to bring effective management and control efforts into practice. There is a need for clear terminology to examine weedy, naturalized and/or invasive complex constituents that have been placed under the contentious umbrella of ‘L. camara L. (sensu lato)’. The time is ripe for invasion ecologists, cytogeneticists and conservationists to collaboratively focus on disentangling the complex and integrate their knowledge and expertise into management and control programs.


Control, genetics, invasive, species complex, taxonomy


Lantana camara L. is one of the world’s worst ten invaders (Lowe et al. 2000). The copious invader has disseminated rapidly at temporal as well as spatial scale. This widespread invader inhabits a wide range of habitats adversely impacting native plant species diversity and ecosystem functioning (Gentle and Duggin 1997, Sharma et al. 2005, Love et al. 2009a, Sharma and Raghubanshi 2009). Despite profound environmental and economic threats posed by L. camara infestations in non-native range, considerable success has not been achieved to effectively control its spread (Cilliers 1983, Bhagwat et al. 2012). Nevertheless, species invasiveness is too complex to be simply predicted and managed (Williamson 1996). However, we anticipate that the prime reason behind the failure to control invasive Lantana could be its existence as an unresolved species complex, L. camara L. (sensu lato). Inability to correctly identify the plant of interest in the complex has incapacitated control measures (Thomas and Ellison 1999, Day et al. 2003a, Urban et al. 2011).

Historically, taxonomy of the genus Lantana has been very complicated (Sanders 2006, Love et al. 2009b). Further, recognition of all weedy and/or invasive genets created due to endless episodes of horticultural improvement within the genus is extremely challenging. Over the last two to three decades, a few articles highlighted the seminal concept of addressing different weedy and/or invasive genets [cultivars, variants, sub-species, hybrids, varieties, forms or even allies (if they exist); hereafter, referred as complex constituents] as L. camara L. (sensu lato) (Stirton 1977, Stirton 1979, Sanders 2006, Sanders 2012; for details, see Urban et al. 2011). However, the importance of this concept has been realized in invasion ecology after decades of consistently delimiting all of the invasive complex constituents with the epithet L. camara.

Importantly, little information is available on taxonomic identity and genetic makeup of existent complex constituents and their populations. This further makes it difficult to surmise what facilitates their success as invaders. Considerably, complex constituents’ invasive success might be ascribed to phenotypic and/or genotypic novelty created by rigorous hybridization events within the complex, their variable response to selection or their differential adaptive plasticity. However, in order to ascertain the key attribute(s) responsible for their invasive success, identification and differentiation of all possible invasive genets in the complex is fundamental. Detailed study of the complex constituents will offer opportunities to answer key questions about plant invasions. Moreover, it may be useful in making informed choices about monitoring their spread throughout the invaded range. The present synthesis will generate an impetus for comprehensive research efforts to study the remarkably diverse species complex in order to appraise control efforts.

Historical events leading to the species complex

Although specific origin of L. camara is unknown, some authors have suggested the species to be a native of South America or Mexico (Howard 1969, Spies and du Plessis 1987), while others suggested West Indies as the place of origin (Moldenke 1973, Palmer and Pullen 1995). Studies report that Lantana (including L. camara) was imported from America to Europe in mid-16th and 17th centuries for its horticultural value (Stirton 1977, Swarbrick et al. 1995). In Europe, the species underwent substantial horticultural breeding, creating hundreds of cultivars of mixed parentage from the introduced stock (Howard 1969). Subsequently, these cultivars traversed to America, Australia, India and Africa in the mid-19th century (Howard 1969, Stirton 1977, Swarbrick 1985, Morton 1994). With time, many cultivars escaped cultivation, spread beyond the ornamental confines of the garden and became weeds (Spies 1984, Swarbrick 1985, Palmer and Pullen 1995). Studies have identified hybridization to contribute substantially to invasiveness, weediness and/or range expansion (Brown and Marshall 1981, Ellstrand and Schierenbeck 2000, Hovick and Whitney 2014). Likewise, innumerable intentional as well as unintentional hybridization events in Lantana led to remarkable increase in its complexity. Anthropogenically-induced genetic diversity in the species complex indeed facilitated the species to invade heterogeneous habitats (Cilliers 1983, Bhagwat et al. 2012, Goncalves et al. 2014). The highly invasive species now exists in 60 countries and island groups of Asia, Africa and Australia (Cronk and Fuller 1995, Day et al. 2003b).

Enigma of the species complex

Extensive hybridization followed by polyploidy or polyploidization followed by hybridization events within and between wild, naturalized and cultivated taxa further enhanced complexity leading to the evolution of L. camara L. (sensu lato) (Sanders 1987, Sanders 2006). Wild complex constituents have also been reported to hybridize and genetically assimilate with the rare native counterparts, threatening the existence of rare genets (see Maschinski et al. 2010). These evolutionary processes have led to enormous phenotypic as well as genotypic variability, which complicates species delimitation in the complex. The ones growing in wild potentially differ morphologically, karyologically, physiologically and ecologically from those prized for their horticultural value, multicolored flowers, and ease of propagation (Spies 1984, Sanders 2006). Therefore, weedy, naturalized and/or invasive complex constituents, broadly referred as L. camara L. (sensu lato) merit a deliberate taxonomic delineation (Sanders 2006).

Complex constituents can be distinguished morphologically (flower size, shape and color; leaf size, hairiness and color; stem thorniness; height and branch architecture), physiologically (growth rates, toxicity to livestock) and, by their chromosome number, nuclear DNA content (Stirton 1979, Gujral and Vasudevan 1983, Scott et al. 1997) and ploidy level (Stirton 1977, Palmer and Pullen 1995). Studies have also reported leaf anatomical characteristics (Passos et al. 2009) and detailed chemical profiling of foliar chemical constituents (Love et al. 2009b, Sena et al. 2012) as useful markers for supporting species delimitation. However, obscure limits of natural variation hamper workers in the field to effectively classify and disentangle complex constituents. The disputed limits of L. camara also complicate identification of the genotypes that have naturalized and are proliferating in the non-native range.

Understanding the species complex

Unlocking diversity in the complex is considered a formidable taxonomic problem (Khoshoo and Mahal 1961, Howard 1969, Moldenke 1971, Spies 1984, Sanders 2006). The complex with a broad spectrum of variability has no record of parental species after 1492 (Stirton 1977). It is highly difficult to deduce putative parents of each constituent in wild as they interbreed freely leading to immense variation in the gene pool (Binggeli 2003, Spies 1984, Urban et al. 2011). Further, ongoing hybridization events in the cryptic complex have blurred taxonomic distinctions of complex constituents and reduced classification accuracy (Sanders 2006, Maschinski et al. 2010). Although the taxonomically uncertain complex has not been subjected to considerable progress till date; a few studies have attempted to reasonably explore the composition and nomenclature to classify the genus Lantana through well-devised keys (Munir 1996, Rajendran and Daniel 2002, Méndez Santos 2002, Sanders 2006, Sanders 2012).

Cytological studies on different populations of Lantana have reported basic chromosome numbers to be 11 & 12 (Henderson 1969, Sinha and Sharma 1984) and 8 & 11 (Moldenke 1983). They have also pointed out the existence of polyploid series in the genus based on reported base numbers. Highest chromosome number was reported as 2n = 72 (Natarajan and Ahuja 1957), and 2n = 66 (Bir and Chatha 1983), while 2n = 22 was recorded to be the lowest (Sen and Sahni 1955, Sanders 1987). Basic chromosome numbers of 11 & 12 have been recorded, with ploidy levels ranging from diploid to hexaploid in L. camara (Tjio 1948, Spies 1984, Ojha and Dayal 1992, Munir 1996, Brandao et al. 2007). Frequent hybridizations between different ploidy levels have also been reported (Spies 1984). Existence of multiple polyploidization pathways has been considered to contribute towards enormous complexity in L. camara (Czarnecki II and Deng 2009).

Rapid adaptive evolution and genetic change have been proposed to contribute significantly to the success of invasive species in the introduced range (Prentis et al. 2008, Prentis and Pavasovic 2013). Studies have revealed significant information regarding genetic variation, population differentiation, and introduction history of a few invasive species using molecular markers (Chun et al. 2010, Thompson et al. 2012, Vardien et al. 2013). Few studies have also attempted to explore range expansion of L. camara in different countries (Vardien et al. 2012, Ray and Quader 2014). Microsatellite markers developed for L. camara have also been successfully used in assessment of genetic variation and population structure in India, broadening understanding on dynamics of its introduction, range expansion and gene flow (Ray and Quader 2014, Ray and Ray 2014). Ray and Quader (2014) identified that the present diversity of L. camara in India is an output of multiple introduction episodes followed by gradual spatial expansion with the recurrent gene flow. Recently, Ray and Ray (2014) studied genetic variation in L. camara in India and synthesized that the species consists of two genetic clusters, representing emerging ecotypes across space that could be differentially adapted to local habitat conditions. Broadly, these studies have revealed high genetic diversity in L. camara and have tried to elucidate past dispersal patterns (Vardien et al. 2013, Ray and Quader 2014, Ray and Ray 2014). Further, these markers can be useful in addressing several questions about breeding system, pollination and dispersal of the species (Ray et al. 2012). However, to improve our understanding of the range expansion of the complex, there is a need for further research at a global scale that examines genetic and genomic attributes of the complex constituents. It is highly essential to understand the genetic system of a taxon as it affects the nature and extent of variability, evolutionary processes and pathways which may further affect invasiveness.

Unresolved species complex: an impediment to management efforts

Interestingly, a large proportion of invasive alien plants are those that were introduced as ornamentals (Mack and Lonsdale 2001, Foxcroft et al. 2008). In general, ornamental plants selected for introduction pose a high invasion risk as they possess traits such as high fruit/seed production, high growth rate, and tolerance to a wide range of environmental conditions (Anderson et al. 2006).

Highly variable L. camara is one amongst those introduced ornamental species that has a wide scale of distribution. In spite of longer residence time, there is a dearth of scientific studies integrating its history, spread, ecological impact, evolution, and management. Thus, attempts to control the invader using mechanical, chemical and biological means have met with limited success (Morton 1994, Thomas and Ellison 2000, Day et al. 2003a). Additionally, this can also be attributed to a gamut of complex constituents that differ in their distribution, habitat preferences, weediness, morphology, chemical constituents, toxicity to livestock, susceptibility to herbicide treatment, and susceptibility to bio-control agents (Smith and Smith 1982, Cilliers and Neser 1991). Further, success of bio-control agents employed in controlling L. camara (including all weedy and/or invasive complex constituents) may vary from one location to another owing to differential feeding habits of the bio-control agents, their host-specificity, climatic suitability, and plant-insect interactions (Broughton 2000, Zalucki et al. 2007). Biological control measures are principally constrained by our confounding understanding of the broad spectrum of phenotypic and genotypic variability present within the complex. However, genetic analysis of the complex can aid in identification of potential control agents to be specifically targeted (Scott et al. 2002).

Future directions

Under the current scenario of genetic diversity and associated taxonomic ambiguity, distinction of genets in L. camara L. (sensu lato) is highly dubious. However, a few studies erroneously address invasive Lantana with ambiguous identity as Lantana camara L. or Lantana camara L. (sensu stricto), which is extremely misleading with the current understanding of the complex. Merely considering L. camara, representative of all troublesome weedy genets in the whole complex, will neither ensure understanding all of the myriad invasive traits, nor would it serve to appropriately answer management questions. Hence, studies pertaining to invasive Lantana should address the individuals in the wild with the epithet ‘L. camara L. (sensu lato)’ (Stirton 1977, Neser and Cilliers 1990, Munir 1996, Baars and Neser 1999, Day and Zalucki 2009). Referral of the invasive complex constituents broadly as L. camara L. (sensu lato) is considered correct under both the International Code of Botanical Nomenclature and the International Code of Nomenclature for Cultivated Plants (see Urban et al. 2011).

To circumscribe the complex constituents, documentation of appropriate suite of distinguishing characters of complex constituents might facilitate delineation of the considerable variation existing in the complex. A consistent terminology based on morphology, cytology, and genetic attributes using advanced molecular techniques such as DNA-based molecular marker techniques, viz. random amplified polymorphism DNA (RAPD), inter simple sequence repeat (ISSR), amplified fragment length polymorphism (AFLP), quantitative trait locus (QTL) mapping, etc. can be devised to explore the genetic diversity of the complex constituents. Integrating the knowledge of plant morphology and chromosome number can also aid in species delimitation, as complex constituents have been reported to behold varying chromosome complements. Attempts to investigate total spectrum of variation using DNA C-values by flow cytometry can be extremely helpful to unravel the diversity in the complex (Suda et al. 2014). Though highly desirable, yet the extremely difficult task of ascertaining absolute taxonomic status to each of the complex constituents can be resolved by estimation of their genome sizes. Variation in chromosome numbers in genus Lantana encourages the use of genome size as a species-specific marker. Genome size has been well-applied to resolve notable species complexes such as Reynoutria (Mandák et al. 2003); Knautia arvensis (Kolář et al. 2009); Dryopteris carthusiana (Ekrt et al. 2010); Callitriche (Prančl et al. 2014) and, identification of invasive alien taxa (see Suda et al. 2010). Further, documentation of population cytotype structure of the invasive genets and their geographical distribution is central to monitor complex constituents’ invasion potential.

High diversity in the complex and continuing hybridization events may potentially broaden its ecological tolerance in climatically suitable as well as unsuitable areas (Goncalves et al. 2014). Furthermore, realizing remarkable spread and better performance of the invasive genets in warmer areas, it is highly probable that invasive Lantana will increase its expanse noticeably in future climate change scenarios (Zhang et al. 2014). Control of the invader would be quite challenging in future scenarios of global change. Lack of knowledge about actual genetic diversity in the complex will further undermine all efforts to regulate species’ invasion dynamics.

Concluding remarks

The study warrants that there is an urgent need to resolve the species complex to ensure concerted management and timely control over its proliferation. In a nutshell, future of efforts to control invasive Lantana lies with resolution of the species complex. There is an urgent need to disentangle the complex to decipher the niche adaptation and range expansive modifications that distinct complex constituents underwent over the evolutionary timeframe. Using insights, we can build-up and enhance our understanding of different facets of L. camara L. (sensu lato) invasion in entirety.


NG acknowledges Senior Research Fellowship (SRF) support from University Grants Commission, India. GPS acknowledges funding support from University of Delhi, India and Department of Science and Technology, India. Authors are grateful to the anonymous referee for the constructive criticism.


  • Anderson NO, Galatowitsch SM, Gomez N (2006) Selection strategies to reduce invasive potential in introduced plants. Euphytica 148: 203–216. doi: 10.1007/s10681-006-5951-7
  • Baars JR, Neser S (1999) Past and present initiatives on the biological control of Lantana camara (Verbenaceae) in South Africa. In: Olckers T, Hill MP (Eds) Biological Control of Weeds in South Africa (1990–1998). African Entomology Memoir 1: 21–33.
  • Bhagwat SA, Breman E, Thekaekara T, Thornton TF, Willis KJ (2012) A battle lost? Report on two centuries of invasion and management of Lantana camara L. in Australia, India and South Africa. PLoS ONE 7(3): e32407. doi: 10.1371/journal.pone.0032407
  • Binggeli P (2003) Verbenaceae, Lantana camara, fankatavinakoho, fotatra, mandadrieko, rajejeka, radredreka, ramity. In: Goodman SM, Benstead JP (Eds) The Natural History of Madagascar. University of Chicago Press, Chicago, 415–417.
  • Bir SS, Chatha GS (1983) SOCGI plant chromosome number reports-1. Journal of Cytology & Genetics 18: 56–58.
  • Brandao AD, Viccini LF, Salimena FRG, Vanzela ALL, Recco-Pimentel SM (2007) Cytogenetic characterization of Lippia alba and Lantana camara (Verbenaceae) from Brazil. Journal of Plant Research 120: 317–321. doi: 10.1007/s10265-006-0041-4
  • Broughton S (2000) Review and evaluation of lantana biocontrol programs. Biological Control 17(3): 272–286. doi: 10.1006/bcon.1999.0793
  • Brown AHD, Marshall DS (1981) Evolutionary changes accompanying colonization in plants. Evolution Today. In: Scudder GGE, Reveal JL (Eds) Proceedings of Second International Congress of Systematics and Evolutionary Biology. Carnegie-Mellon University, Pittsburg, 351–363.
  • Chun YJ, Fumanal B, Laitung B, Bretagnolle F (2010) Gene flow and population admixture as the primary post-invasion processes in common ragweed (Ambrosia artemisiifolia) populations in France. New Phytologist 185: 1100–1107. doi: 10.1111/j.1469-8137.2009.03129.x
  • Cilliers CJ (1983) The weed, Lantana camara L., and the insect natural enemies imported for its biological control into South Africa. Journal of the Entomological Society of Southern Africa 46: 131–138.
  • Cilliers CJ, Neser S (1991) Biological control of Lantana camara (Verbenaceae) in South Africa. Agriculture, Ecosystems & Environment 37: 57–75. doi: 10.1016/0167-8809(91)90139-O
  • Cronk QCB, Fuller JL (1995) Plant invaders. Chapman & Hall, London.
  • Czarnecki II DM, Deng Z (2009) Occurrence of unreduced female gametes leads to sexual polyploidization in Lantana. Journal of the American Society for Horticultural Science 134: 560–566.
  • Day MD, Broughton S, Hannan-Jones MA (2003a) Current distribution and status of Lantana camara and its biological control agents in Australia, with recommendations for further biocontrol introductions into other countries. BioControl 24: 63–76.
  • Day MD, Wiley CJ, Playford J, Zalucki MP (2003b) Lantana: current management status and future prospects. Australian Centre for International Agricultural Research Monograph 102, Canberra.
  • Day MD, Zalucki MP (2009) Lantana camara Linn. (Verbenaceae). In: Muniappan R, Reddy GVP, Raman A (Eds) Biological Control of Tropical Weeds using Arthropods. Cambridge University Press, Cambridge (UK), 211–246. doi: 10.1017/CBO9780511576348.012
  • Ekrt L, Holubová R, Trávníček P, Suda J (2010) Species boundaries and frequency of hybridization in the Dryopteris carthusiana (Dryopteridaceae) complex: a taxonomic puzzle resolved using genome size data. American Journal of Botany 97: 1208–1219. doi: 10.3732/ajb.0900206
  • Ellstrand NC, Schierenbeck KA (2000) Hybridization as a stimulus for the evolution of invasiveness in plants? Proceedings of the National Academy of Sciences of the United States of America 97: 7043–7050. doi: 10.1073/pnas.97.13.7043
  • Foxcroft LC, Richardson DM, Wilson JRU (2008) Ornamental plants as invasive aliens: problems and solutions in Kruger National Park, South Africa. Environmental Management 41: 32–51. doi: 10.1007/s00267-007-9027-9
  • Gentle CB, Duggin JA (1997) Lantana camara L. invasions in dry rainforest–open forest ecotones: the role of disturbances associated with fire and cattle grazing. Australian Journal of Ecology 22: 298–306. doi: 10.1111/j.1442-9993.1997.tb00675.x
  • Goncalves E, Herrera I, Duarte M, Bustamante RO, Lampo M, Grisel V, Sharma GP, García-Rangel S (2014) Global invasion of Lantana camara: has the climatic niche been conserved across continents? PLoS ONE 9: e111468. doi: 10.1371/journal.pone.0111468
  • Gujral GS, Vasudevan P (1983) Lantana camara L., a problem weed. Journal of Scientific & Industrial Research 42: 281–286.
  • Henderson RJF (1969) A cytological study of Lantana montevidensis (Spreng.) Briq. in Queensland. Contributions from the Queensland Herbarium 3: 1–4.
  • Hovick SM, Whitney KD (2014) Hybridisation is associated with increased fecundity and size in invasive taxa: meta-analytic support for the hybridization-invasion hypothesis. Ecology Letters 17: 1464–1477. doi: 10.1111/ele.12355
  • Howard RA (1969) A check list of cultivar names used in the genus Lantana. Arnoldia 29: 73–109.
  • Khoshoo TN, Mahal C (1961) Versatile reproduction in Lantana camara. Current Science 36: 201–203.
  • Kolář F, Štech M, Trávníček P, Rauchová J, Urfus T, Vít P, Kubešová M, Suda J (2009) Towards resolving the Knautia arvensis agg. (Dipsacaceae) puzzle: primary and secondary contact zones and ploidy segregation at landscape and microgeographic scales. Annals of Botany 103: 963–974. doi: 10.1093/aob/mcp016
  • Love A, Babu S, Babu CR (2009a) Management of Lantana, an invasive alien weed, in forest ecosystems of India. Current Science 97: 1421–1429.
  • Love A, Naik D, Basak SK, Babu S, Pathak N, Babu CR (2009b) Variability in foliar essential oils among different morphotypes of Lantana species complexes, and its taxonomic and ecological significance. Chemistry & Biodiversity 6: 2263–2274. doi: 10.1002/cbdv.200800284
  • Lowe S, Browne M, Boudjelas S, De Poorter M (2000) 100 of the world’s worst invasive alien species. a selection from the Global Invasive Species Database. The Invasive Species Specialist Group (ISSG) a specialist group of the Species Survival Commission (SSC) of the World Conservation Union (IUCN), Auckland.
  • Mandák B, Pyšek P, Lysák M, Suda J, Krahulcová A, Bímová K (2003) Variation in DNA-ploidy levels of Reynoutria taxa in the Czech Republic. Annals of Botany 92: 265–272. doi: 10.1093/aob/mcg141
  • Maschinski J, Sirkin E, Fant J (2010) Using genetic and morphological analysis to distinguish endangered taxa from their hybrids with the cultivated exotic pest plant Lantana strigocamara (syn: Lantana camara). Conservation Genetics 11: 1607–1621. doi: 10.1007/s10592-009-0035-6
  • Méndez Santos IE (2002) A taxonomic revision of Lantana sect. Lantana (Verbenaceae) in the Greater Antilles. Willdenowia 32: 285–301.
  • Moldenke HN (1971) “A Fifth Summary of the Verbenaceae, Avicenniaceae, Stilbaceae, Dicrastylidaceae, Symphoremaceae, Nyctanthaceae and Eriocaulaceae of the world... ” etc. 1 & 2. Wayne, New Jersey.
  • Moldenke HN (1973) Verbenaceae. In: Woodson RE, Schery RW (Eds) Flora of Panama. Part 9. Annals of the Missouri Botanical Garden 60: 4–148.
  • Moldenke HN (1983) Verbenaceae. In: Dassanayake MD, Fosberg FA (Eds) A Revised Handbook to the Flora of Ceylon. Amerind Publishing Co., New Delhi, 196–487.
  • Morton JF (1994) Lantana, or red sage (Lantana camara L. (Verbenaceae)), notorious weed and popular garden flower; some cases of poisoning in Florida. Economic Botany 48: 259–270. doi: 10.1007/BF02862327
  • Munir AA (1996) A taxonomic review of Lantana camara L. and L. montevidensis (Spreng.) Briq. (Verbenaceae) in Australia. Journal of the Adelaide Botanical Gardens 17: 1–27.
  • Natarajan AT, Ahuja MR (1957) Cytotaxonomical studies in the genus Lantana. Journal of the Indian Botanical Society 36: 35–45.
  • Neser S, Cilliers CJ (1990) Work towards biological control of Lantana camara: perspectives. In: Delfosse ES (Ed.) Proceedings of the Seventh International Symposium on Biological Control of Weeds. Istituto Sperimentale per la Patologia Vegetale, Ministero dell’ Agricoltura e delle Foreste, Rome, Italy, 363–369.
  • Ojha BM, Dayal N (1992) Cytological investigations in the genus Lantana in India. Cytologia 57: 9–13. doi: 10.1508/cytologia.57.9
  • Palmer WA, Pullen KR (1995) The phytophagous arthropods associated with Lantana camara, L. hirsuta, L. urticifolia, and L. urticoides (Verbenaceae) in North America. Biological Control 5: 54–72. doi: 10.1006/bcon.1995.1007
  • Passos JL, Meira RMSA, Barbosa LCA (2009) Foliar anatomy of the species Lantana camara and L. radula (Verbenaceae). Planta Daninha 27: 689–700. doi: 10.1590/S0100-83582009000400007
  • Prančl J, Kaplan Z, Trávníček P, Jarolímová V (2014) Genome size as a key to evolutionary complex aquatic plants: polyploidy and hybridization in Callitriche (Plantaginaceae). PLoS ONE 9: e105997. doi: 10.1371/journal.pone.0105997
  • Prentis PJ, Pavasovic A (2013) Understanding the genetic basis of invasiveness. Molecular Ecology 22: 2366–2368. doi: 10.1111/mec.12277
  • Prentis PJ, Wilson JRU, Dormontt EE, Richardson DM, Lowe AJ (2008) Adaptive evolution in invasive species. Trends in Plant Science 13: 288–294. doi: 10.1016/j.tplants.2008.03.004
  • Rajendran A, Daniel P (2002) The Indian Verbenaceae (A Taxonomic Revision). Bishen Singh & Mahendra Pal Singh, Dehra Dun, 179–195.
  • Ray A, Quader S (2014) Genetic diversity and population structure of Lantana camara in India indicates multiple introductions and gene flow. Plant Biology 16: 651–658. doi: 10.1111/plb.12087
  • Ray A, Ray R (2014) Rapid divergence of ecotypes of an invasive plant. AoB PLANTS: plu052. doi: 10.1093/aobpla/plu052
  • Ray A, Sumangala RC, Ravikanth G, Uma Shaanker R, Quader S (2012) Isolation and characterization of polymorphic microsatellite loci from the invasive plant Lantana camara L. Conservation Genetics Resources 4: 171–173. doi: 10.1007/s12686-011-9501-9
  • Sanders RW (1987) Taxonomic significance of chromosome observations in Caribbean species of Lantana (Verbenaceae). American Journal of Botany 6: 914–920. doi: 10.2307/2443872
  • Sanders RW (2006) Taxonomy of Lantana sect. Lantana (Verbenaceae): I. Correct application of Lantana camara and associated names. SIDA 22(1): 381–421.
  • Sanders RW (2012) Taxonomy of Lantana Sect. Lantana (Verbenaceae): II. Taxonomic Revision. Botanical Research Institute of Texas 6: 403–441.
  • Scott LJ, Graham GC, Hannan-Jones MA, Yeates DK (1997) DNA Profiling resolves the limited importance of flower colour in defining varieties of Lantana camara. Electrophoresis 18: 1560–1563. doi: 10.1002/elps.1150180912
  • Scott LJ, Hannan-Jones MA, Graham GC (2002) Affinities of Lantana camara in the Australia–Pacific region. In: Spafford-Jacob H, Dodd J, Moore JH (Eds) Proceedings of the Thirteenth Australian Weeds Conference. Perth, Australia, 471–474.
  • Sen NK, Sahni VM (1955) Triploid, tetraploid and pentaploid Lantana camara. Science & Culture 20: 558–559.
  • Sena FJG, Xavier HS, Barbosa FJM, Duringer JM (2012) A chemical marker proposal for the Lantana genus: composition of the essential oils from the leaves of Lantana radula and L. canescens. Natural Product Communications 5: 635–40.
  • Sharma GP, Raghubanshi AS (2009) Lantana invasion alters soil nitrogen pools and processes in the tropical dry deciduous forest of India. Applied Soil Ecology 42: 134–140. doi: 10.1016/j.apsoil.2009.03.002
  • Sinha S, Sharma A (1984) Lantana camara L.-a review. Feddes Repertorium 95: 621–633.
  • Smith LS, Smith DA (1982) The naturalised Lantana camara complex in Eastern Australia. Queensland Botanical Bulletin 1: 1–26.
  • Spies JJ (1984) A cytotaxonomic study of Lantana camara (Verbenaceae) from South Africa. South African Journal of Botany 3: 231–250.
  • Spies JJ, du Plessis H (1987) Sterile Lantana camara: fact or theory. South African Journal of Plant & Soil 4: 171–174.
  • Stirton CH (1977) Some thoughts on the polyploid complex Lantana camara L. (Verbenaceae). In: Annecke DP (Ed.) Proceedings of the Second National Weeds Conference of South Africa. Balkema, South Africa, 321–340.
  • Stirton CH (1979) Taxonomic problems associated with invasive alien trees and shrubs in South Africa. In: Proceedings of the 9th Plenary Meeting AETFAT (Association pour l’étude taxonomique de la flore d’Afrique tropicale). Brussells, Belgium, 218–219.
  • Suda J, Meyerson LA, Leitch IJ, Pyšek P (2014) The hidden side of plant invasions: the role of genome size. New Phytologist. doi: 10.1111/nph.13107
  • Suda J, Trávníček P, Mandák B, Berchová-Bímová K (2010) Genome size as a marker for identifying the invasive alien taxa in Fallopia section Reynoutria. Preslia 82: 97–106.
  • Swarbrick JT (1985) History of the lantanas in Australia and origins of the weedy biotypes. Plant Protection Quarterly 1: 115–121.
  • Swarbrick JT, Willson BW, Hannan-Jones MA (1995) The biology of Australian Weeds 25. Lantana camara L. Plant Protection Quarterly 10: 82–95.
  • Thomas SE, Ellison CA (1999) The susceptibility of selected biotypes of Lantana camara and Lantana montevidensis from South Africa to the rust Prospodium tuberculatum. Research Report to Agricultural Research Council-Plant Protection Research Institute by CABI Bioscience, Berkshire (UK).
  • Thomas SE, Ellison CA (2000) A century of classical biological control of Lantana camara: can pathogens make a significant difference? In: Spencer NR (Ed.) Proceedings of the Tenth International Symposium on Biological Control of Weeds. Montana State University, USA, 97–104.
  • Thompson GD, Bellstedt DU, Byrne M, Millar MA, Richardson DM, Wilson JRU, Le Roux JJ (2012) Cultivation shapes genetic novelty in a globally important invader. Molecular Ecology 21: 3187–3199. doi: 10.1111/j.1365-294X.2012.05601.x
  • Urban AJ, Simelane DO, Retief E, Heystek F, Williams HE, Madire LG (2011) The invasive ‘Lantana camara L.’ hybrid complex (Verbenaceae): a review of research into its identity and biological control in South Africa. African Entomology 19: 315–348. doi: 10.4001/003.019.0225
  • Vardien W, Richardson DM, Foxcroft LC, Wilson JRU, Le Roux JJ (2013) Management history determines gene flow in a prominent invader. Ecography 36: 1–10. doi: 10.1111/j.1600-0587.2012.00120.x
  • Vardien W, Richardson DM, Foxcroft LC, Wilson JRU, Le Roux JJ (2012) Invasion dynamics of Lantana camara L. (sensu lato) in South Africa. South African Journal of Botany 81: 81–94. doi: 10.1016/j.sajb.2012.06.002
  • Williamson M (1996) Biological Invasions. Chapman & Hall, London.
  • Zalucki MP, Day MD, Playford J (2007) Will biological control of Lantana camara ever succeed? Patterns, processes & prospects. Biological Control 42: 251–261. doi: 10.1016/j.biocontrol.2007.06.002
  • Zhang Q, Zhang Y, Peng S, Zobel K (2014) Climate warming may facilitate invasion of the exotic shrub Lantana camara. PLoS ONE 9: e105500. doi: 10.1371/journal.pone.0105500