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-caeruleumLC02 [=Lithospermum purpureocaeruleumH91]
    `--+--Cystostemon heliocarisLC02
       |--Lithodora diffusaLC02
       |--Cerinthe [Cerintheae]LC02
       |    `--C. majorLC02
       |    |--L. arvense [=Buglossoides arvensis]H93
       |    |--L. californicumH93
       |    |--L. callosumD17
       |    |--L. cobrenseBT72
       |    |--L. incisumH93
       |    |--L. incrassatumC55b
       |    |--L. officinaleC55b
       |    |--L. prostratumC55a
       |    `--L. ruderaleH93
            |--Lobostemon fruticosusLC02
                 |--E. aculeatumMP05
                 |--E. arenariumPT98
                 |--E. candicans [incl. E. pininana, E. strictum]H93
                 |--E. confusumD17
                 |--E. humileD17
                 |--E. italicumBR65
                 |--E. maritimumC55b
                 |--E. plantagineumBR65
                 |--E. pyramidatumC55a
                 |--E. rauwolfiD17
                 |--E. violaceumBR65
                 `--E. vulgareLC02
Lithospermeae incertae sedis:
    |--A. graeca Boissier & Spruner 1844PL04
    |    |--A. g. ssp. graecaPL04
    |    `--A. g. ssp. baeotica (de Candolle) Nyman 1881PL04
    `--A. tinctoriaC74
    |--A. decumbensD17
    |--A. euchromaO88
    `--A. nepalensisO88
  Maharanga emodiLC02, O88
    |--O. bracteatumO88
    |--O. fruticosumD30
    `--O. hookeriO88

*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.

[D17] Diels, L. 1917. Beiträge zur Flora der Zentral-Sahara und ihrer Pflanzengeographie. Nach der Sammelausbeute des Freiherrn Hans Geyr von Schweppenburg. Botanische Jahrbücher für Systematik, Pflanzengeschichte und Pflanzengeographie 54 (Beiblatt 120): 51–155.

[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.

[MP05] Moraza, M. L., & M. A. Peña. 2005. Ácaros Mesostigmata (Acari, Mesostigmata) de hábitats seleccionados de la Gomera (Islas Canarias, España). Graellsia 61 (1): 109–114.

[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.

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