Viper's bugloss Echium vulgare, copyright Emily Kloosterman.

Belongs within: Boraginaceae.

The Lithospermeae are a clade of plants in the Boraginaceae which have the style usually divided, and nutlets with a broad basal attachment scar (Långström & Chase 2002). This group includes the viper's buglosses of the genus Echium, some of which are widely grown as ornamentals though a few have become invasive species in some parts of the world. Lithospermum, the stoneseeds and gromwells, bear yellow (sometimes white) flowers in which the corolla tube is longer than the free petals (Hickman 1993).

Characters (from Långström & Chase 2002): Style usually divided at apex, with two to four stigmas; pollen colporate, heteropolar or prolate; nutlets with broad basal attachment scar, flat gynobase.

    |--Buglossoides purpureo-caeruleum LC02 [=Lithospermum purpureocaeruleum H91]
    `--+--Cystostemon heliocaris LC02
       |--Lithodora diffusa LC02
       |--Cerinthe [Cerintheae] LC02
       |    `--C. major LC02
       |--Lithospermum LC02
       |    |--L. arvense [=Buglossoides arvensis] H93
       |    |--L. californicum H93
       |    |--L. cobrense BT72
       |    |--L. incisum H93
       |    |--L. incrassatum C55b
       |    |--L. officinale C55b
       |    |--L. prostratum C55a
       |    `--L. ruderale H93
       `--Echieae LC02
            |--Lobostemon fruticosus LC02
            |--Megacaryon LC02
            |--Zwackhia LC02
            `--Echium LC02
                 |--E. arenarium PT98
                 |--E. candicans [incl. E. pininana, E. strictum] H93
                 |--E. italicum BR65
                 |--E. maritimum C55b
                 |--E. plantagineum BR65
                 |--E. pyramidatum C55a
                 |--E. violaceum BR65
                 `--E. vulgare LC02

Lithospermeae incertae sedis:
  Alkanna LC02
    |--A. graeca Boissier & Spruner 1844 PL04
    |    |--A. g. ssp. graeca PL04
    |    `--A. g. ssp. baeotica (de Candolle) Nyman 1881 PL04
    `--A. tinctoria C74
  Ancistrocarya LC02
  Arnebia LC02
    |--A. euchroma O88
    `--A. nepalensis O88
  Choriantha LC02
  Colsmannia LC02
  Halacsya LC02
  Lasiarrhenum LC02
  Macromeria LC02
  Macrotomia LC02
  Maharanga emodi LC02, O88
  Meratia LC02
  Onosmodium LC02
  Pentalophus LC02
  Stenosolenium LC02
  Onosma LC02
    |--O. bracteatum O88
    |--O. fruticosum D30
    `--O. hookeri O88

*Type species of generic name indicated


[BT72] Baker, E. W., & D. M. Tuttle. 1972. New species and further notes on the Tetranychoidea mostly from the southwestern United States (Acarina: Tetranychidae and Tenuipalpidae). Smithsonian Contributions to Zoology 116: 1–37.

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

[C55a] Candolle, A. de. 1855a. Géographie Botanique Raisonée: Ou exposition des faits principaux et des lois concernant la distribution géographique des plantes de l’époque actuelle vol. 1. Librairie de Victor Masson: Paris.

[C55b] Candolle, A. de. 1855b. 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.

[C74] Coineau, Y. 1974. Éléments pour une monographie morphologique, écologique et biologique des Caeculidae (Acariens). Mémoires du Muséum National d'Histoire Naturelle, nouvelle série, Série A, Zoologie 81: 1–299, 24 pls.

[D30] Druce, G. C. 1930. Account of a botanical tour in Cyprus. Proceedings of the Linnean Society of London 141: 50–52.

[H93] Hickman, J. C. (ed.) 1993. The Jepson Manual: Higher Plants of California. University of California Press: Berkeley (California).

[H91] Hubálek, Z. 1991. Biogeographic indication of natural foci of tick-borne infections. In: Dusbábek, F., & V. Bukva (eds) Modern Acarology: Proceedings of the VIII International Congress of Acarology, held in České Budĕjovice, Czechoslovakia, 6–11 August 1990 vol. 1 pp. 255–260. SPB Academic Publishing: The Hague.

[LC02] Långström, E., & M. W. Chase. 2002. Tribes of Boraginoideae (Boraginaceae) and placement of Antiphytum, Echiochilon, Ogastemma and Sericostoma: A phylogenetic analysis based on atpB plastid DNA sequence data. Plant Systematics and Evolution 234: 137–153.

[O88] Ohba, H. 1988. The alpine flora of the Nepal Himalayas: an introductory note. In: Ohba, H., & S. B. Malla (eds) The Himalayan Plants vol. 1. The University Museum, University of Tokyo, Bulletin 31: 19–46.

[PT98] Panitsa, M., & D. Tzanoudakis. 1998. Contribution to the study of the Greek flora: Flora and vegetation of the E Aegean islands Agathonisi and Pharmakonisi. Willdenowia 28: 95–116.

[PL04] Pohl, G., & I. Lenski. 2004. Zur Verbreitung und Vergesellschaftung von Pennisetum orientale Rich. in Nordeuböa (Griechenland) (Poaceae, Paniceae). Senckenbergiana Biologica 83 (2): 209–223.

Last updated: 16 October 2019.

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