1. Boraginaceae Juss.

    1. This family is accepted.

[FWTA]

Boraginaceae, H. Heine. Flora of West Tropical Africa 2. 1963

Habit
Trees, shrubs or herbs, often roughly scabrid or hispid
Leaves
Leaves alternate or rarely opposite, simple; stipules absent
Flowers
Flowers often in scorpoid cymes, actinomorphic or rarely oblique, mostly hermaphrodite
Calyx
Calyx-lobes imbricate or rarely valvate
Corolla
Corolla with contorted or imbricate lobes
Androecium
Anthers 2-celled, opening lengthwise Stamens the same number as the corolla-lobes and alternate with them, inserted on the corolla
Nectaries
Disk present or absent
Gynoecium
Ovules paired, erect or spreading from the central axis Ovary superior, 2-celled or 4-celled by spurious septa, entire or deeply 4-lobed; style terminal or gynobasic
Fruits
Fruit a drupe or of 4: nutlets
Seeds
Seeds with or without endosperm
[NTK]

Stapf, M.N.S. (2009). Neotropical Boraginaceae.

Morphology
Description

Herbs, subshrubs, shrubs, trees, rarely lianas; rarely obligate root parasites with chlorophyll absent in Lennoa Lex. Hairs usually present, occasionally containing cystoliths, simple or glandular . Leaves usually alternate , rarely subopposite (Tournefortia L.), sometimes basal (Plagiobothrys Fisch. & C.A.Mey.), simple , rarely compound (Phacelia Juss.), reduced to scales in Lennoa, exstipulate ; lamina entire , usually rough to hispid , margins mostly entire , or crenate , or serrate , or dentate with venation pinnate domatia occurring in Cordia L. and Ehretia P.Browne. Inflorescence terminal , axillary , or intermodal, cymose, the branches often scorpioid, helicoid, or reduced and capitate to glomerate (as Varronia P.Browne), rarely reduced to one solitary flower (some Heliotropium L.). Flowers actinomorphic , sometimes distylous (Varronia and Cordia), usually 5- merous , hermaphrodite or unisexual; calyx usually 5- lobed , distinct or connate . Corolla sympetalous, usually 5- lobed ; stamens isomerous, alternate with corolla lobes; filaments adnate to  corolla tube; anthers 2-locular, dehiscing longitudinally;  ovary superior , 2-carpellate, but usually becoming falsely 4-locular, with axile placentation or often parietal with 2 placentae in Phacelia; ovules usually 4, anatropous to hemitropous; the style terminal or gynobasic, simple , or bifid (Ehretioideae, Phacelia), or twice 2- cleft (Cordia and Varronia); stigmas 1-4;  nectar disk usually present around base of ovary . Fruits fleshy or dry drupaceous , or mericarps 2 (2-seeded) or 4 ( 4-seeded), or a capsule ; seeds 4 per fruit , often less by abortion; endosperm present or absent; cotyledons entire of flat, or rarely plicate (Cordia and Varronia).  

Diagnostic
Notable genera and distinguishing features
  • Cordia is characterized by the habit (trees or shrubs); inflorescence cymose, the branches often scorpioid, helicoid; terminalstyle twice bifid forming four stigmatic lobes; fruitdrupaceous. 
  • Varronia is characterized by the habit (multi-stemmed shrubs or rarely several-stemmed treelets); inflorescence reduced, capitate, glomerate, spicate, or if cymose, these less than 4 cm broad; terminalstyle twice bifid forming four stigmatic lobes; fruitdrupaceous. 
  • Heliotropium is characterized by a terminalstyle and dry fruits, which divide into four or two mericarps.
  • Tournefortia is characterized by a terminalstyle and drupaceous fruits, which never divide into mericarps.
  • Ehretia is characterized by the habit (trees or shrubs), and bifid terminalstyle.
Distinguishing characters (always present)
  • Leaves usually alternate, exstipulate.
  • Inflorescence cymose, the branches often scorpioid, helicoid, or reduced and capitate to glomerate.
  • Corolla gamopetalous.
  • Stamens isomerous inserted in the corolla tube.
  • Ovarysuperior, 2-carpellate, but usually becoming falsely 4-locular.
Other important characters
  • Cystoliths commonly present.
Useful tips for generic identification
  • Habit
  • Inflorescence
  • Style
  • Fruit
General Description
Status
  • Many species have been introduced as ornamental plants or weeds, e.g. the genera Anchusa L., Borago L., Cynoglossum L., Echium L., Myosotis L. and Symphytum L..
  • The genus Varronia is endemic.
Notes on delimitation
  • The family is circumscribed broadly, including Hydrophyllaceae (except Hydrolea L.) and Lennoaceae.
  • It is considered monophyletic on the basis of inflorescence type and DNA sequences data (Ferguson 1999; Gottschling et al. 2001).
Number of genera

The Boraginaceae comprise approximately 205 genera and 2500 species worldwide, of which 23 genera and approximately 500 species are neotropical.

 

Neotropical genera:

  • Amsinckia Lehm.
  • Antiphytum DC. ex Meisn.
  • Bourreria P.Browne
  • Cordia (including Auxemma Miers, Patagonula L.and Saccellium Humb. & Bonpl.)
  • Cryptantha Lehm. ex Fisch. & C.A.Mey.
  • Ehretia
  • Hackelia Opiz
  • Heliotropium
  • Lasiarrhenum I.M. Johnst.
  • Lennoa
  • Lepidocordia Ducke
  • Lithospermum L.
  • Macromeria D.Don
  • Moritzia DC. ex Meisn.
  • Nama L.
  • Pectocarya DC. ex Meisn.
  • Phacelia
  • Plagiobothrys
  • Rochefortia Sw.
  • Tiquilia Pers.
  • Tournefortia
  • Wigandia Kunth
  • Varronia P.Browne
Literature
Important literature

Candolle, A.P. de. 1845. Borragineae. Pp. 467-501. In De Candolle, A. (ed.), Prodromus Systematis Naturalis Regni Vegetabilis, 9. Treuttel & Würtz, Paris.

Diane, N., H. Förther & H.H. Hilger. 2002. A systematic analysis of Heliotropium, Tournefortia, and allied taxa of the Heliotropiaceae (Boraginales) based on ITS1 sequences and morphological data. American Journal of Botany 89(2): 287-295.

Ferguson, D. M. 1999. Phylogenetic analysis and relationships in Hydrophyllaceae s. str. based on ndhF sequence data. Syst. Bot. 23: 253-268.

Gottschling, M. & J.S. Miller. 2006. Clarification of the taxonomic position of Auxemma, Patagonula, and Saccellium (Cordiaceae, Boraginales). Syst. Bot. 31(2): 361-367.

Gottschling, M., H.H. Hilger, M. Wolf & N. Diane. 2001. Secondary structure of the ITS1 transcript and its application in a reconstruction of the phylogeny of Boraginales. Plant Biology 3: 629-636.

Gottschling, M., M. Weigend & H.H. Hilger. 2005. Congruence of a phylogeny of Cordiaceae (Boraginales) inferred from ITS1 sequence data with morphology, ecology, and biogeography. Ann. Missouri Bot. Gard. 92: 425-437.

Miller, J.S. & M. Gottschling. 2007. Generic classification in the Cordiaceae (Boraginales): Resurrection of the genus Varronia P. Br. (Cordiaceae). Taxon 56(1): 163-169.

Yatskievych, G. and C.T. Mason. 1986. A revision of the Lennoaceae. Syst. Bot. 11: 531-548.

Smith, R.A., D.M. Ferguson, T.J. Barkman & C.W. Pamphilis. 2000. Molecular phylogenetic evidence for the origin of Lennoaceae: A case of adelphoparasitism in the angiosperms? Amer. J. Bot. [Suppl.] 87: 158.

Stapf, M.N.S. 2007.  Avaliação da classificação infragenérica de Cordia L. (Cordiaceae) e revisão taxonómica de Cordia sect. Pilicordia DC. Para o Brasil.  Tese Doutorado Botánica, Feira de Santana: Universidade Estadual de Feira, Brasil.

[FTEA]

Boraginaceae, B. Verdcourt. Flora of Tropical East Africa. 1991

Habit
Annual or perennial herbs, often woody at the base, subshrubs, trees or woody climbers, usually characteristically scabrid or hispid
Leaves
Leaves simple, alternate or less often opposite, petiolate or sessile, entire or variously toothed or crenate; stipules absent
Inflorescences
Inflorescences terminal or axillary, bifurcate or in raceme-like or panicled groups, basically dichotomous cymes, the branches frequently scorpioid and spirally inrolled when young; bracts present or absent
Flowers
Flowers hermaphrodite or unisexual (Cordia), sessile or pedicellate, mostly 5-merous, regular or somewhat irregular, occasionally heterostylous
Calyx
Calyx tubular or campanulate, persistent and sometimes accrescent in fruit, with (2–)4–7 (or more) imbricate or rarely valvate or open lobes
Corolla
Corolla tubular, campanulate, funnel-shaped or rotate with tube very reduced; lobes 3–16, imbricate or contorted (>i>Myosotis); throat often with scales, thickenings, folds or crests
Androecium
Stamens the same number and alternating with the corolla-lobes, exserted or included, epipetalous; anthers 2- thecous
Nectaries
Nectariferous disc usually present
Gynoecium
Style usually 1, 2 in one genus or sometimes 4 in one species of >i>Cordia, terminal or gynobasic, entire or 2–4-fid or twice 2-fid; stigmas clavate, capitate or peltate or linear to narrowly foliaceous Ovules axile, anatropous, erect, horizontal, basal or rarely pendulous Ovary superior, entire or deeply 4-lobed, 2-locular with 2 ovules in each or with 4, 1-ovuled locules due to the development of false partitions
Fruits
Fruit drupaceous with 1, 4-celled endocarp or separating in 2 bilocular or 4 unilocular 1-seeded pyrenes or with 4 quite distinct nutlets
Seeds
Seed with or without endosperm, erect, oblique or horizontal; testa membranous; embryo straight or curved; cotyledons flat or plicate
[FZ]

Boraginaceae, E. S. Martins (Trichodesma by R. K. Brummit). Flora Zambesiaca 7:4. 1990

Habit
Herbs, shrubs or rarely trees
Leaves
Leaves alternate, rarely subopposite or opposite, simple exstipulate
Inflorescences
Inflorescence usually cymose, composed of one or more helicoid or scorpioid cymes, sometimes paniculate or racemose or sometimes flowers solitary, terminal or axillary
Flowers
Flowers hermaphrodite or sometimes unisexual, actinomorphic or sometimes zygomorphic
Calyx
Calyx (4)5(6)-lobed or 3–5-toothed, imbricate or valvate
Corolla
Corolla (4)5(6)-lobed, imbricate or contorted in bud, tubular, funnel-shaped, campanulate, salver-shaped or rotate; tube sometimes with folds, scales or hairs in the throat
Androecium
Stamens (or staminodes) as many as corolla lobes and alternate with them, inserted on the corolla tube; anthers 2-thecous, dehiscing longitudinally, introrse, dorsifixed
Nectaries
Disk present or absent
Gynoecium
Ovary superior, entire or deeply 4-lobed, bi-locular or spuriously 4-locular, placentation axial, ovules 4, erect or spreading; style 1, terminal or gynobasic, entire or cleft or twice cleft, or styles 2, terminal (Coldenia)
Fruits
Fruit drupaceous with a 1–4-seeded stone or splitting into 4 pyrenes, or of one to four-seeded nutlets; seeds generally without endosperm
[FWTA]

Hoplestigmataceae, F.N. Hepper. Flora of West Tropical Africa 2. 1963

Habit
Trees
Leaves
Leaves alternate, entire; stipules absent
Flowers
Flowers hermaphrodite, in terminal cymes
Calyx
Calyx closed in bud, bursting irregularly
Corolla
Corolla shortly tubular, with about 3 series of overlapping corolla lobes
Androecium
Stamens numerous (about 21), in several rows inserted at the base of the corolla; anthers 4-celled, opening by longitudinal slits
Gynoecium
Ovules pendulous, one on each side of the bilobed placentas Ovary superior, 1-celled with 2 intrusive parietal placentas; style bilobed nearly to the base, branches sharply bent; stigmas capitate
Fruits
Fruit with a deep groove up each side, the calyx forming a plate at the base, endocarp hard
Seeds
Seeds with scanty endosperm and long thick straight embryo
[FWTA]

Hydrophyllaceae, F.N. Hepper. Flora of West Tropical Africa 2. 1963

Habit
Annual or perennial herbs
Leaves
Leaves radical or alternate, rarely opposite, entire to pinnately or palmately lobed
Flowers
Flowers usually cymose, hermaphrodite, actinomorphic
Calyx
Calyx-segments 5, imbricate
Corolla
Corolla gamopetalous, lobes imbricate or contorted
Androecium
Stamens the same number as the corolla-lobes and alternate with them, mostly inserted towards the base of the tube; anthers 2-celled
Gynoecium
Ovules numerous Ovary superior, 1-celled with 2 parietal placentas, or 2-3-celled; styles 1 or 2
Fruits
Fruit a loculicidal or rarely septicidal capsule
Seeds
Seeds with fleshy endosperm and small straight embryo
[FTEA]

Hydrophyllaceae, B. Verdcourt. Flora of Tropical East Africa. 1989

Habit
Annual or perennial herbs, rarely subshrubs, often scabrid
Leaves
Leaves usually alternate or radical, exstipulate, entire to deeply pinnately or palmately lobed, the margins entire, serrate or lobulate
Flowers
Flowers regular, hermaphrodite, in scorpioid or trichotomous cymes, false racemes or clustered, less often solitary
Calyx
Calyx deeply divided into 5 (rarely 8–12) imbricate or open, equal or unequal lobes or 5 with appendages between
Corolla
Corolla often blue, variously shaped, the lobes usually 5 (rarely 8–12), imbricate or rarely contorted
Androecium
Stamens as many as the corolla-lobes, inserted at the sinuses or low in the tube, filaments filiform or dilated at the base; anthers 2-thecous, versatile, opening by longitudinal slits, included or exserted
Nectaries
Disk inconspicuous or absent
Gynoecium
Ovary superior (very unusually inferior), 1-locular with 2 parietal fleshy placentas or 2(–3)-locular; ovules 2 or more on each placenta; style filiform, usually bifid or styles 2; stigmas simple or capitate
Fruits
Fruit usually a loculicidal or less often septicidal 2-valved or irregularly splitting capsule
Seeds
Seeds oblong, globose or angled, with tuberculate, reticulate or rugose testa; albumen copious and fleshy; embryo small; cotyledons entire
[FZ]

Hydrophyllaceae, B. Verdcourt. Flora Zambesiaca 7:4. 1990

Habit
Annual or perennial herbs, rarely subshrubs, often scabrid
Leaves
Leaves usually alternate or radical, exstipulate, entire to deeply pinnately or palmately lobed, with margins entire, serrate or lobulate
Flowers
Flowers regular, hermaphrodite, in scorpioid or trichotomous cymes, false racemes or clustered, less often solitary
Calyx
Calyx deeply divided into 5 rarely 8–12 imbricate or open equal or unequal lobes or 5 with appendages between
Corolla
Corolla often blue, variously shaped, the lobes usually 5, rarely 8–12, imbricate or rarely contorted
Androecium
Stamens as many as the corolla lobes, inserted at the sinuses or low in the tube; filaments filiform or dilated at the base; anthers bilocular, versatile, opening by longitudinal slits, included or exserted
Nectaries
Disk inconspicuous or absent
Gynoecium
Ovary superior (very unusually inferior) unilocular with 2 parietal fleshy placentas or 2(3)-locular Ovules 2 or more on each placenta; style filiform, usually bifid or styles 2; stigmas simple or capitate
Fruits
Fruit usually a loculicidal or less often septicidal bivalved or irregularly splitting capsule
Seeds
Seeds oblong, globose or angled with tuberculate reticulate or rugose testa; albumen copious and fleshy; embryo small; cotyledons entire

Images

Boraginaceae Juss. appears in other Kew resources:

First published in Gen. Pl. [Jussieu] 128. 1789 [4 Aug 1789] (1789)

Accepted by

  • APG IV (2016) http://dx.doi.org/10.1111/boj.12385

Sources

Flora Zambesiaca
Flora Zambesiaca
http://creativecommons.org/licenses/by-nc-sa/3.0

Flora of Tropical East Africa
Flora of Tropical East Africa
http://creativecommons.org/licenses/by-nc-sa/3.0

Flora of West Tropical Africa
Flora of West Tropical Africa
http://creativecommons.org/licenses/by-nc-sa/3.0

Kew Names and Taxonomic Backbone
The International Plant Names Index and World Checklist of Selected Plant Families 2018. Published on the Internet at http://www.ipni.org and http://apps.kew.org/wcsp/
© Copyright 2017 International Plant Names Index and World Checklist of Selected Plant Families. http://creativecommons.org/licenses/by/3.0

Neotropikey
Milliken, W., Klitgard, B. and Baracat, A. (2009 onwards), Neotropikey - Interactive key and information resources for flowering plants of the Neotropics.
http://creativecommons.org/licenses/by/3.0