Common cocklebur Xanthium strumarium, photographed by J. V. Littell.

Belongs within: Asteroideae.

The Heliantheae are a group of composite-flowered plants distinguished by their scaled receptacles and (usually) stiff pappuses. Significant members include the sunflower Helianthus annuus and Jerusalem artichoke H. tuberosus. Molecular studies indicate the presence of two major clades within the Heliantheae; interestingly, the morphologically similar genera Montanoa and Rojasianthe, sharing such characters as chromosome number, opposite phyllotaxy, accrescent pales, blackish disc corollas and neutral ray flowers with white corollas, are each placed basally on oppposite clades (Panero & Funk 2002).

Characters (from Black & Robertson 1965): Receptacle with chaffy scales; involucral bracts without scarious margins; heads mostly heterogamous-radiate; pappus of scales, awns or stiff plumose bristles, rarely absent; style-branches flattish, not truncate; leaves often opposite.

    |  i. s.: Spilanthes paniculata PF02, BB07
    |         Galinsoga [Galinsoginae] BR65
    |           `--G. parviflora BR65
    |--+--Montanoa PF02
    |  |    |--M. bipinnatifida Z02
    |  |    |--M. hibiscifolia H06
    |  |    |--M. liebmannii PF02
    |  |    `--M. pteropoda PF02
    |  `--Ecliptinae PF02
    |       |--Salmea PF02
    |       `--Eclipta BR65
    |            |--E. alba PP07
    |            |--E. erecta C55
    |            |--E. platyglossa BR65
    |            `--E. prostrata VB02
    `--+--Rojasianthe Standley & Steyermark 1940 [Rojasianthinae] PF02
       `--+--Rudbeckiinae PF02
          |--Zaluzania YY22 [Zaluzaniinae PF02]
          |--Zinnia YY22 [Zinniinae PF02]
          |--Engelmaniinae PF02
          |--Helianthus BR65 [Helianthinae PF02]
          |    |--H. annuus Linnaeus 1753 CD07
          |    |--H. bolanderi D51
          |    |--H. paradoxus MAA95
          |    |--H. petiolaris D51
          |    `--H. tuberosus BR65
          |--Verbesininae PF02
          |    |--Siegesbeckia orientalis BR65
          |    |--Verbesina DH98
          |    |    |--V. encelioides DH98
          |    |    `--V. turbaiensis MM96
          |    `--Wedelia von Jacquin 1763 BR65
          |         |--W. biflora P88
          |         |--W. glauca (Orteg) Blake 1917 [=Pascalia glauca Orteg 1797] SS02
          |         |--W. serrata J87
          |         |--W. trilobata P88
          |         `--W. verbesinoides [incl. W. stirlingii] BR65
          `--Ambrosiinae [Ambrosieae] PF02
               |--Iva axillaris BR65
               |--Ambrosia BR65
               |    |--A. artemisiifolia BR65
               |    |--A. dumosa C90
               |    |--A. psilostachya BR65
               |    `--A. tenuifolia C55
               `--Xanthium BR65
                    |--X. californicum BR65
                    |--X. canadense MH98
                    |--X. indicum C55
                    |--X. macrocarpum C55
                    |--X. orientale BR65
                    |--X. pungens BR65
                    |--X. spinosum BR65
                    `--X. strumarium BR65

*Type species of generic name indicated


[BB07] Baishya, A. K. & P. J. Bora. 2007. Cross community ethno-medico botany of Dibru-Saikhowa Biosphere Reserve, Assam. Bulletin of the Botanical Survey of India 49 (1-4): 121-154.

[BR65] Black, J. M., & E. L. Robertson. 1965. Flora of South Australia. Part IV. Oleaceae-Compositae. W. L. Hawes, Government Printer: Adelaide.

[C55] Candolle, A. de. 1855. Géographie Botanique Raisonée: Ou exposition des faits principaux et des lois concernant la distribution géographique des plantes de l’époque actuelle vol. 2. Librairie de Victor Masson: Paris.

[CD07] Cantino, P. D., J. A. Doyle, S. W. Graham, W. S. Judd, R. G. Olmstead, D. E. Soltis, P. S. Soltis & M. J. Donoghue. 2007. Towards a phylogenetic nomenclature of Tracheophyta. Taxon 56 (3): E1-E44.

[C90] Crawford, C. S. 1990. Scorpiones, Solifugae, and associated desert taxa. In: Dindal, D. L. (ed.) Soil Biology Guide pp. 421-475. John Wiley & Sones: New York.

[DH98] Danin, A., & I. C. Hedge. 1998. Contributions to the flora of Jordan 2. A new species of Satureja (Labiatae) and some new records. Willdenowia 28: 135-142.

[D51] Dobzhansky, T. 1951. Genetics and the Origin of Species, 3rd ed. Columbia University Press: New York.

[H06] Henderson, L. 2006. Comparisons of invasive plants in southern Africa originating from southern temperate, northern temperate and tropical regions. Bothalia 36 (2): 201-222.

[J87] Judd, W. S. 1987. Floristic study of Morne La Visite and Pic Macaya National Parks, Haiti. Bulletin of the Florida State Museum – Biological Sciences 32 (1): 1-136.

[MAA95] McCarthy, E. M., M. A. Asmussen & W. W. Anderson. 1995. A theoretical assessment of recombinational speciation. Heredity 74: 502-509.

[MH98] Morikawa, H., A. Higaki, M. Nohno, M. Takahashi, M. Kamada, M. Nakata, G. Toyohara, Y. Okamura, K. Matsui, S. Kitani, K. Fujita, K. Irifune & N. Goshima. 1998. More than a 600-fold variation in nitrogen dioxide assimilation among 217 plant taxa. Plant, Cell and Environment 21: 180-190.

[MM96] Mound, L. A., & R. Marullo. 1996. The thrips of Central and South America: an introduction (Insecta: Thysanoptera). Memoirs on Entomology, International 6: 1-487.

[PP07] Pandey, R. P. & P. M. Padhye. 2007. Studies on phytodiversity of Arid Machia Safari Park-Kailana in Jodhpur (Rajasthan). Bulletin of the Botanical Survey of India 49 (1-4): 15-78.

[PF02] Panero, J. L., & V. A. Funk. 2002. Toward a phylogenetic subfamilial classification for the Compositae (Asteraceae). Proceedings of the Biological Society of Washington 115 (4): 909-922.

[P88] Polunin, I. 1988. Plants and Flowers of Malaysia. Times Editions: Singapore.

[SS02] Salunkhe, C. B., V. B. Shimpale, M. M. Sardesai & S. R. Yadav. 2002. Some new records of Asteraceae for the state of Maharashtra. Journal of the Bombay Natural History Society 99 (3): 555-557.

[VB02] Vijay, S. K., & T. N. Bhardwaja. 2002. Vegetation and phenodynamics of wetlands of central Rajasthan. Journal of the Bombay Natural History Society 99 (3): 573-581.

[YY22] Yampolsky, C., & H. Yampolsky. 1922. Distribution of sex forms in the phanerogamic flora. Bibliotheca Genetica 3: 1-62.

[Z02] Zhang, Z.-Q. 2002. Taxonomy of Tetranychus ludeni (Acari: Tetranychidae) in New Zealand and its ecology on Sechium edule. New Zealand Entomologist 25: 27-34.

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