Italian bugloss Anchusa azurea, copyright Alberto Salguero Quiles.

Belongs within: Boraginaceae.

The Boragineae is a group of plants in the family Boraginaceae found almost entirely in Eurasia, characterised by flowers with a simple style and usually well-developed faucal appendages (Långström & Chase 2002). Borage Borago officinalis is a native of the Mediterranean region that is cultivated as a vegetable green.

Borage and comfrey and bugloss
Published 12 October 2010
Anchusa undulata ssp. granatensis, photographed by James Gaither.

The tribe Boragineae includes about 170 species of herbaceous flowering plants, mostly found in the Palaearctic region with only a couple of species extending into southern Africa. The group is well-distinguished by the presence of what are called fornices, the whitish lobes at the base of each petal that you can see in the photo above, as well as features of their seeds. Many Boragineae seeds have an elaiosome, a fatty plug at one end that attracts foraging ants (Hilger et al. 2004). The ants carry the seed back to their nest as food, but the plant produces enough seeds that at least some will not be eaten but will be able to germinate after being carried under the ground and away from anything else that might eat them.

Abraham-Isaac-Jacob, Trachystemon orientalis, a native of forests around the Black Sea and one of the more unusual species of Boragineae. Apparently the unusual name refers to the flowers changing colour as they age. Photo by Daniel Mosquin.

The species are divided between about fifteen genera (the exact number varies depending on whom you ask). The largest generally-recognised genus, Anchusa (the buglosses), was identified by Hilger et al. (2004) as para-/polyphyletic with a number of smaller genera also nested within the Anchusa clade, suggesting that the currently recognised constituent subgenera may need to be recognised as separate subgenera (or else the genera Lycopsis and Cynoglottis submerged into Anchusa). Other relationships within the tribe recognised by this and other studies include a close relationship between the genera Borago (borage) and Symphytum (comfrey), and between Nonea and Pulmonaria (lungwort). The basalmost member of the tribe is Pentaglottis sempervirens, which is also the only member of the tribe found in the Atlantic region of southwest Europe. The relict distribution of this species, as well as the concentration of diversity for the tribe overall, have been cited as supporting a Mediterranean origin for the Boragineae.

Green alkanet, Pentaglottis sempervirens, the sister species to all other Boragineae, photographed by Carl Farmer.

A number of members of the tribe have long been cultivated and many are even labelled by their botanical names as officinal (the Medieval Latin term ‘officinalis’ refers to a plant or substance that is kept in an apothecary; not surprisingly, many plants with supposed medicinal values are also eaten for their nutritional values). Borago officinalis, borage, is used as a salad or pot herb in Europe. Symphytum officinale, comfrey, has also been widely used medicinally, mainly for external uses such as soothing bruises (some of the properties attributed to comfrey verge on the ridiculous: a bath steeped in comfrey was supposedly able to restore a woman’s virginity). Pulmonaria officinalis, lungwort, received its name because of the supposed resemblance of its blotchy leaves to lung tissue. Under the unabashedly loopy herbalist principle known as the Doctrine of Signatures, this outward resemblance indicated its suitability in treating lung diseases such as tuberculosis (in fact, lungwort contains toxic alkaloids that make it dangerous to take internally).

Systematics of Boragineae

Characters (from Långström & Chase 2002): Flowers with usually well-developed faucal appendages; style simple, with one or two stigmas; pollen colporate; nutlets with basal scar often surrounded by annular rim, gynobase flat.

<==Boragineae [Anchusae, Borageae]
    |--Borago officinalis Linnaeus 1753LC02, CD07
    `--+--+--Nonea versicolorLC02
       |  `--Pentaglottis sempervirensLC02
            |--A. arvensis [=Lycopsis arvensis]H93
            |--A. azureaH91
            |--A. caespitosaSL06
            |--A. capensisBR65
            |--A. officinalis [incl. A. procera]H93
            `--A. ovataO88
Boragineae incertae sedis:
  Caryolopha sempervirensLC02, C55b [=Anchusa sempervirensC55b]
  Cynoglottis barrelieriLC02, GR98
    |--S. asperumH93
    |--S. caucasicumV09
    |--S. cordatumH09
    `--S. officinaleC55b
    |--L. orientalisC55b
    `--L. variegataC55a
  Pulmonaria L. 1753LC02, KC01
    |--P. mollisC55a
    |--P. officinalisC55b
    |--P. rubraH09
    `--P. virginicaC55b

*Type species of generic name indicated


[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.

[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.

[GR98] Greuter, W., & T. Raus (eds.) 1998. Med-Checklist Notulae, 17. Willdenowia 28: 163–174.

[H09] Heltmann, H. 2009. Der Königstein (Piatra Craiului), die Perle der Burzenländer Gebirge. Mauritiana 20 (3): 515–527.

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

Hilger, H. H., F. Selvi, A. Papini & M. Bigazzi. 2004. Molecular systematics of Boraginaceae tribe Boragineae based on ITS1 and trnL sequences, with special reference to Anchusa s.l. Annals of Botany 94 (2): 201–212.

[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.

[KC01] Kirk, P. M., P. F. Cannon, J. C. David & J. A. Stalpers. 2001. Ainsworth & Bisby’s Dictionary of the Fungi 9th ed. CAB International: Wallingford (UK).

[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.

[SL06] Schulz, H.-J., & P. Lymberakis. 2006. First contribution to the knowledge of the Collembola fauna of the White Mountains (Lefká Óri) in west Crete (Insecta, Collembola, Isotomidae). Senckenbergiana Biologica 86 (2): 229–234.

[T-W89] Tenison-Woods, J. E. 1889. On the vegetation of Malaysia. Proceedings of the Linnean Society of New South Wales, series 2, 4 (1): 9–106, pls 1–9.

[V09] Verdcourt, B. (ed.) 2009. Additions to the wild fauna and flora of the Royal Botanic Gardens, Kew. XXVI. Miscellaneous records. Kew Bulletin 64 (1): 183–194.

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