Subphases1.
Hydrophyllaceae.
2.
Wellstediaceae?.
3.
Cynoglossoideae.
4.
Boraginoideae.
5.
Lithospermoideae.
6.
Ehretiaceae, including
Cordiaceae,
Heliotropiaceae.
7.
Lennoaceae.
Plant theory 2Boraginales is maintained as an order in the Subclass
Lamiidae in
Phase 3. The theme of not being seen or heard is central in
Boraginales. This was first discovered in a case of Borago offcinalis. Later it got extended to all
Boraginaceae and Borginales. It was found to be a theme of
Phase 3.
Plant theory 1In the
Plant theory Boraginales is treated as an order in the Subclass
Lamiidae. Boraginaceea is split in Subfamilies, treated as families in
Boraginales.
Boraginales have strong
Boron qualities, which has a peculiar similarity in the name.
Boraginales is placed in
Phase 3. The confusion about their placement fits with
Phase 3.
TaxonomyIn the Cronquist calssification
Boraginales does not exist as an oder.
Boraginaceae as a family includes
Cordiaceae,
Ehretiaceae and
Heliotropiaceae and is placed together with
Lennoaceae in
Lamiales.
Hydrophyllaceae was placed in
Solanales.
In the Apg2 and the
Apg3 classification Boraginales is not recognised.
In the
Apg3 classification Boraginales is not recognised.
Boraginaceae as traditionally defined is paraphyletic and needs the inclusion of
Hydrophyllaceae and
Lennoaceae to become monophyletic. Apg3 includes
Boraginaceae in
Lamiidae, but otherwise unplaced, but as a probable sister to
Lamiales.
In the
Apg4 classification Boraginales is an order in
Lamiidae with only one family
Boraginaceae, including
Hydrophyllaceae,
Cordiaceae,
Ehretiaceae,Codonaceae, and
Heliotropiaceae.
In some recent publications
Boraginaceae, as defined in the
Apg3 classification, is split into several families:
Boraginaceae,
Cordiaceae,
Ehretiaceae,
Heliotropiaceae and
Lennoaceae; these families comprise the Order
Boraginales.
Codon was long regarded as an unusual member of
Hydrophyllaceae, but in 1998, a molecular phylogenetic study showed that it is closer to
Boraginaceae. Some authors have defined a resticted
Boraginaceae excluding Codon en Wellstedia. They were placed in the monogeneric Families Codonaceae and
Wellstediaceae.
Boraginaceae is hard to characterize morphologically if it includes the genera Codon and Wellstedia.
Lennoaceae consists of the achlorophyllous holoparasites Lennoa and Pholisma. Studies indicatye that
Lennoaceae form a clade within
Ehretiaceae.
Some studies have indicated that
Hydrophyllaceae is paraphyletic if the tribe Nameae is included within it, but further studies will be needed to resolve this issue.
The genus Hoplestigma in
Boraginales was occasionally doubted. In 2014 a study showed it to be a close relative of
Cordiaceae.
Hydrolea was thought to belong in
Hydrophyllaceae for more than a century. It is now shown to belong to the order
Solanales as sister to Sphenoclea.
Pteleocarpa was long placed in
Boraginales, but with considerable doubt. The molecular evidence strongly supports it as sister to
Gelsemiaceae.
BotanyBoraginales: inflorescence a scorpioid cyme!; two-parted gynoecium!; leaves mostly alternate, mostly with only evident midrib, if so leaves abundant, relatively narrow, thick indument; vestured pits present; flowers actinomorphic; corolla plicate; leaves hairy, rough, felty, trichomes; placentation parietal; ovary with secondary septae; pollen with pseudocolpi.
Content: pyrrolizidine alkaloids; iridoids lacking.
Hydrophyllaceae: vestured pits absent.
Boraginaceae includes ± 110 genera and ± 1600 species.
Boraginaceae: inflorescence a scorpioid cyme; a gynobasic style, and a two-part ovary that breaks into four nutlets; flowers 5 merous; bracteate inflorescences; faucal appendages, also called fornices, thickenings or inward evaginations of the corolla.
Heterostyly is common, scattered in at least nine genera among three tribes.
Codon: flowers 10 to 12-merous; fruit a capsule.
Wellstedia: flowers 4-merous; fruit a capsule.
Trichodesmeae includes Caccinia Savi and Trichodesma, is included in
Cynoglossoideae; pollen with 2 to 5 pores, isocolpate.
Cynoglossoideae: nutlets with non-basal attachment; pollen 6 to 8 pores, heterocolpat; bracteate inflorescences absent.
Eritricheae, including Eritrichium, Hackelia, Lappula, Lepechiniella, Rochelia, Suchtelenia, is included in
Cynoglossoideae.
Lithospermoideae: fornices absent; flowers zygoomorphic.
Variable: corolla shapes; pollen size, shape, pore number, pore position; ellipsoid pollen is ancestral.
LiteratureHilger, H & others; Molecular Systematics of
Boraginaceae Tribe Boragineae Based on ITS1 and trnL Sequences, with Special Reference to Anchusa s.l.; Annals of
Botany 94: 201–212; 2004.
Nazaire, Mare; Hufford, Larry; A Broad Phylogenetic
Analysis of
Boraginaceae: Implications for the Relationships of Mertensia; Systematic
Botany, Volume 37,Number 3, pp. 758-783; 2012.
Luebert, Federico, et al,
Boraginales Working Group; Familial classification of the
Boraginales; TAXON 65.3; p 502–522; 2016.
Nazaire, Mare & Hufford, Larry; A Broad Phylogenetic
Analysis of
Boraginaceae: Implications for the Relationships of Mertensia; Systematic
Botany, 37,3, pp. 758–783; 2012.