Belongs within: Lamiidae.
Contains: Ehretioideae, Hydrophylloideae, Heliotropioideae, Cordioideae, Boragineae, Cynoglosseae, Lithospermeae.
The Boraginaceae as recognised herein are a clade supported by molecular analyses (Långström & Chase 2002). They combine 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). Codon is a South African genus of spiny herbs of historically uncertain affinities, bearing broadly campanulate flowers.
Synapomorphies (www.mobot.org): 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
|--+--EhretioideaeLC02
| |--HydrophylloideaeLC02
| `--+--HeliotropioideaeLC02
| `--CordioideaeLC02
`--Boraginoideae [Trigonotideae]LC02
| i. s.: AmphibologyneLC02
| BrachybotrysLC02
| DecalepidanthusLC02
| MairetisLC02
| MertensiaLC02
| | i. s.: M. cryptocarpaD03
| | M. maritimaC55
| | M. paniculataPSM96
| |--M. sect. MertensiaH93
| | |--M. ciliata [incl. M. ciliata var. stomatechoides]H93
| | |--M. cusickiiH93
| | |--M. longifloraH93
| | `--M. oblongifoliaH93
| | |--M. o. var. oblongifoliaH93
| | |--M. o. var. amoenaH93
| | `--M. o. var. nevadensisH93
| `--M. (sect. Neuranthia) bellaH93
| MoltkiopsisLC02
| MoritziaLC02
| Neatostema Johnston 1953LC02, BR65
| `--N. apulum (Linnaeus) Johnston 1953 (see below for synonymy)BR65
| OmphalotrigonotisLC02
| SinojohnstoniaLC02
| ThaumatocaryumLC02
| TrigonocaryumLC02
| ZoelleriaLC02
| Embadium Black 1931BR65
| `--E. stagnense Black 1931BR65
|--BoragineaeLC02
|--CynoglosseaeLC02
`--+--LithospermeaeLC02
`--EchiochileaeLC02
|--Antiphytum hintoniorumLC02
|--Ogastemma pusillumLC02
`--Echiochilon [Echiochilinae]LC02
|--E. collenetteiLC02
|--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 hookeriO88
LasiocaryumO88
|--L. densiflorumO88
|--L. diffusumO88
`--L. munroiO88
Microcaryum pygmaeumO88
Wellstedia [Wellstediaceae, Wellstedioideae]T00
Tournefortia L. 1753L95
|--T. argenteaH09
|--T. bicolor Sw. 1788 [incl. T. laevigata Lam. 1791]L95
|--T. cirrhosa Vaupel 1916V16
|--T. coriacea Vaupel 1916V16
|--T. filiflora Griseb. 1861L95
|--T. glabraJ87
|--T. hirsutissima L. 1753L95
|--T. laurifolia Vent. 1803 [=Messerschmidtia laurifolia (Vent.) R. & S. 1819]L95
|--T. maculata Jacq. 1760 [incl. T. peruviana Poir. 1816, T. syringaefolia Vahl 1794]L95
|--T. microphylla Bertero ex Spreng. 1824 [=T. volubilis var. microphylla (Bertero) DC. 1845]L95
|--T. scabra Lam. 1791L95
|--T. selleanaJ87
`--T. volubilis L. 1753 [=Messerschmidtia volubilis (L.) R. & S. 1819]L95
Messerschmidia argenteaSJP60
CodonT00
LennoaT00
PholismaT00
|--P. arenariumH93
`--P. sonorae [=Ammobroma sonorae]H93
Argusia Amman ex Boehmer in Ludwig 1760L95
|--A. argenteaNC91
`--A. gnaphalodes (L.) Heine 1976 (see below for synonymy)L95
Cienkowskia Regel & Rach 1859KC01
Rotula Lour. 1790KC01
Bourreria Browne 1756 (nom. cons.)L95
|--B. domingensis (DC.) Griseb. 1861 [=Ehretia domingensis DC. 1845]L95
|--B. succulenta Jacq. 1760L95
| |--B. s. var. succulenta [incl. B. succulenta var. canescens Schulz in Urb. 1910]L95
| `--B. s. var. revoluta (HBK.) Schulz in Urb. 1911 [=B. revoluta HBK. 1818]L95
`--B. virgata (Sw.) Don 1838 [=Ehretia virgata Sw. 1788; incl. B. virgata vestita Schulz in Urb. 1911]L95
Rochefortia Sw. 1788L95
|--R. acanthophora (DC.) Griseb. 1862 [=Ehretia acanthophora DC. 1845]L95
`--R. spinosa (Jacq.) Urb. 1920 [=Ehretia spinosa Jacq. 1760; incl. R. cuneata Sw. 1788]L95
Megastoma pusillumD17
Argusia gnaphalodes (L.) Heine 1976 [=Heliotropium gnaphalodes L. 1759, Tournefortia gnaphalodes (L.) Br. ex R. & S. 1819]L95
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
References
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