Few-flowered bushy bugloss Echiochilon pauciflorum, copyright Ankush Dave.

Belongs within: Lamiidae.
Contains: Ehretioideae, Hydrophylloideae, Heliotropioideae, Cordioideae, Boragineae, Cynoglosseae, Lithospermeae.

The Boraginaceae as recognised herein is a clade supported by molecular analyses (Långström & Chase 2002). It combines older concepts of the Boraginaceae with the Hydrophylloideae, previously treated as a separate family distinguished by the possession of capsular fruits. Some members of the family are grown as herbs (such as borage Borago officinalis) or ornamentals (such as forget-me-nots, Myosotis, and heliotropes, Heliotropium). Subfamilies within the Boraginaceae other than the Hydrophylloideae are primarily separated by characters of the gynoecium. Boraginoideae, for instance, have a deeply four-lobed ovary maturing to four nutlets and a basifixed style. Within Boraginoideae, the Echiochileae are a clade found in southern North America, northern Africa and south-western Asia, characterised by a bifid style with two stigmas, a basal or submedial attachment scar, a flat to pyramidal gynobase, and more or less spherical or square pollen (Långström & Chase 2002).

Synapomorphies (from Plant more or less herbaceous; pyrrolizidine alkaloids; cork superficial to mid-cortical (pericyclic); sieve tubes with nuclear non-dispersive protein bodies; nodes 1:1; petiole bundle(s) arcuate; plant more or less roughly hairy, hairs with a basal cystolith or cystolith-like body, and/or walls calcified; leaves spiral; inflorescences terminal, cymose, cyme scorpioid; calyx free, corolla tube formation late; anther placentoid absent; nectary not vascularized; ovary with secondary septae, stigma dry; ovules 2/carpel, apical, pendulous; calyx persistent in fruit; micropylar endosperm haustoria present.

Boraginaceae [Codonaceae, Ehretiaceae, Lennoaceae] APG16
    |--+--Ehretioideae LC02
    |  |--Hydrophylloideae LC02
    |  `--+--Heliotropioideae LC02
    |     `--Cordioideae LC02
    `--Boraginoideae [Trigonotideae] LC02
         |  i. s.: Amphibologyne LC02
         |         Brachybotrys LC02
         |         Decalepidanthus LC02
         |         Mairetis LC02
         |         Mertensia LC02
         |           |  i. s.: M. cryptocarpa D03
         |           |         M. maritima C55
         |           |         M. paniculata PSM96
         |           |--M. sect. Mertensia H93
         |           |    |--M. ciliata [incl. M. ciliata var. stomatechoides] H93
         |           |    |--M. cusickii H93
         |           |    |--M. longiflora H93
         |           |    `--M. oblongifolia H93
         |           |         |--M. o. var. oblongifolia H93
         |           |         |--M. o. var. amoena H93
         |           |         `--M. o. var. nevadensis H93
         |           `--M. (sect. Neuranthia) bella H93
         |         Moltkiopsis LC02
         |         Moritzia LC02
         |         Neatostema Johnston 1953 LC02, BR65
         |           `--N. apulum (Linnaeus) Johnston 1953 (see below for synonymy) BR65
         |         Omphalotrigonotis LC02
         |         Sinojohnstonia LC02
         |         Thaumatocaryum LC02
         |         Trigonocaryum LC02
         |         Zoelleria LC02
         |         Embadium Black 1931 BR65
         |           `--E. stagnense Black 1931 BR65
         |--Boragineae LC02
         |--Cynoglosseae LC02
         `--+--Lithospermeae LC02
            `--Echiochileae LC02
                 |--Antiphytum hintoniorum LC02
                 |--Ogastemma pusillum LC02
                 `--Echiochilon [Echiochilinae] LC02
                      |--E. collenettei LC02
                      |--E. kotschyi [incl. Sericostoma pauciflorum var. qeshmensis Ghahreman & Attar 1996] LC02
                      `--E. pauciflorum (Stocks ex Wight) Långström & Chase 2002 (see below for synonymy) LC02

Boraginaceae incertae sedis:
  Chionocharis hookeri O88
  Lasiocaryum O88
    |--L. densiflorum O88
    |--L. diffusum O88
    `--L. munroi O88
  Microcaryum pygmaeum O88
  Wellstedia [Wellstediaceae, Wellstedioideae] T00
  Tournefortia J87
    |--T. argentea H09
    |--T. bicolor J87
    |--T. glabra J87
    |--T. gnaphalodes RJ11
    |--T. hirsutissima J87
    |--T. laurifolia C55
    `--T. selleana J87
  Messerschmidia argentea SJP60
  Codon T00
  Lennoa T00
  Pholisma T00
    |--P. arenarium H93
    `--P. sonorae [=Ammobroma sonorae] H93
  Argusia argentea CBH93, NC91
  Cienkowskia Regel & Rach 1859 KC01

Echiochilon pauciflorum (Stocks ex Wight) Långström & Chase 2002 [=Sericostoma pauciflorum Stocks ex Wight 1848] LC02

Neatostema apulum (Linnaeus) Johnston 1953 [=Myosotis apula Linnaeus 1753, Lithospermum apulum (Linnaeus) Vahl. 1938, Rhytispermum apulum (Linnaeus) Reichenbach 1858] BR65

*Type species of generic name indicated


[APG16] Angiosperm Phylogeny Group. 2016. An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG IV. Botanical Journal of the Linnean Society 181: 1–20.

[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. 1. Librairie de Victor Masson: Paris.

[CBH93] Collinson, M. E., M. C. Boulter & P. L. Holmes. 1993. Magnoliophyta (‘Angiospermae’). In: Benton, M. J. (ed.) The Fossil Record 2 pp. 809–841. Chapman & Hall: London.

[D03] Dusén, P. 1903. The vegetation of western Patagonia. In: Scott, W. B. (ed.) Reports of the Princeton University Expeditions to Patagonia, 1896–1899 vol. 8. Botany pp. 1–34. The University: Princeton (New Jersey).

[H09] Hedley, C. 1909. The Marine Fauna of Queensland: Address by the President of Section D. Australasian Association for the Advancement of Science: Brisbane.

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

[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.

[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.

[NC91] Nielsen, E. S., & I. F. B. Common. 1991. Lepidoptera (moths and butterflies). In: CSIRO. The Insects of Australia: A textbook for students and research workers 2nd ed. vol. 2 pp. 817–915. Melbourne University Press: Carlton (Victoria).

[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.

[RJ11] Rising, J. D., A. Jaramillo, J. L. Copete, P. G. Ryan & S. C. Madge. 2011. Family Emberizidae (buntings and New World sparrows). In: Hoyo, J. del, A. Elliott & D. A. Christie (eds) Handbook of the Birds of the World vol. 16. Tanagers to New World Blackbirds pp. 428–683. Lynx Edicions: Barcelona.

[SJP60] St. John, H., & W. R. Philipson. 1960. List of the flora of Oeno Atoll, Tuamotu Archipelago, south-central Pacific Ocean. Transactions of the Royal Society of New Zealand 88 (3): 401–403.

[T00] Thorne, R. F. 2000. The classification and geography of the flowering plants: dicotyledons of the class Angiospermae (subclasses Magnoliidae, Ranunculidae, Caryophyllidae, Dilleniidae, Rosidae, Asteridae, and Lamiidae). The Botanical Review 66: 441–647.

Last updated: 27 October 2019.

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