Boraginaceae Juss.

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.

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

Morphology General Habit

Annual or perennial herbs, often woody at the base, subshrubs, trees or woody climbers, usually characteristically scabrid or hispid

Morphology Leaves

Leaves simple, alternate or less often opposite, petiolate or sessile, entire or variously toothed or crenate; stipules absent

Morphology Reproductive morphology 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

Morphology Reproductive morphology Flowers

Flowers hermaphrodite or unisexual (Cordia), sessile or pedicellate, mostly 5-merous, regular or somewhat irregular, occasionally heterostylous

Morphology Reproductive morphology Flowers Calyx

Calyx tubular or campanulate, persistent and sometimes accrescent in fruit, with (2–)4–7 (or more) imbricate or rarely valvate or open lobes

Morphology Reproductive morphology Flowers 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

Morphology Reproductive morphology Flowers Androecium

Stamens the same number and alternating with the corolla-lobes, exserted or included, epipetalous; anthers 2- thecous

Morphology Reproductive morphology Flowers Nectaries

Nectariferous disc usually present

Morphology Reproductive morphology Flowers 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

Morphology Reproductive morphology 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

Morphology Reproductive morphology Seeds

Seed with or without endosperm, erect, oblique or horizontal; testa membranous; embryo straight or curved; cotyledons flat or plicate

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

Morphology General Habit

Herbs, shrubs or rarely trees

Morphology Leaves

Leaves alternate, rarely subopposite or opposite, simple exstipulate

Morphology Reproductive morphology Inflorescences

Inflorescence usually cymose, composed of one or more helicoid or scorpioid cymes, sometimes paniculate or racemose or sometimes flowers solitary, terminal or axillary

Morphology Reproductive morphology Flowers

Flowers hermaphrodite or sometimes unisexual, actinomorphic or sometimes zygomorphic

Morphology Reproductive morphology Flowers Calyx

Calyx (4)5(6)-lobed or 3–5-toothed, imbricate or valvate

Morphology Reproductive morphology Flowers 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

Morphology Reproductive morphology Flowers 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

Morphology Reproductive morphology Flowers Nectaries

Disk present or absent

Morphology Reproductive morphology Flowers 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)

Morphology Reproductive morphology 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

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

Morphology General Habit

Trees

Morphology Leaves

Leaves alternate, entire; stipules absent

Morphology Reproductive morphology Flowers

Flowers hermaphrodite, in terminal cymes

Morphology Reproductive morphology Flowers Calyx

Calyx closed in bud, bursting irregularly

Morphology Reproductive morphology Flowers Corolla

Corolla shortly tubular, with about 3 series of overlapping corolla lobes

Morphology Reproductive morphology Flowers Androecium

Stamens numerous (about 21), in several rows inserted at the base of the corolla; anthers 4-celled, opening by longitudinal slits

Morphology Reproductive morphology Flowers 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

Morphology Reproductive morphology Fruits

Fruit with a deep groove up each side, the calyx forming a plate at the base, endocarp hard

Morphology Reproductive morphology Seeds

Seeds with scanty endosperm and long thick straight embryo

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

Morphology General Habit

Annual or perennial herbs

Morphology Leaves

Leaves radical or alternate, rarely opposite, entire to pinnately or palmately lobed

Morphology Reproductive morphology Flowers

Flowers usually cymose, hermaphrodite, actinomorphic

Morphology Reproductive morphology Flowers Calyx

Calyx-segments 5, imbricate

Morphology Reproductive morphology Flowers Corolla

Corolla gamopetalous, lobes imbricate or contorted

Morphology Reproductive morphology Flowers Androecium

Stamens the same number as the corolla-lobes and alternate with them, mostly inserted towards the base of the tube; anthers 2-celled

Morphology Reproductive morphology Flowers Gynoecium

Ovules numerous Ovary superior, 1-celled with 2 parietal placentas, or 2-3-celled; styles 1 or 2

Morphology Reproductive morphology Fruits

Fruit a loculicidal or rarely septicidal capsule

Morphology Reproductive morphology Seeds

Seeds with fleshy endosperm and small straight embryo

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

Morphology General Habit

Annual or perennial herbs, rarely subshrubs, often scabrid

Morphology Leaves

Leaves usually alternate or radical, exstipulate, entire to deeply pinnately or palmately lobed, the margins entire, serrate or lobulate

Morphology Reproductive morphology Flowers

Flowers regular, hermaphrodite, in scorpioid or trichotomous cymes, false racemes or clustered, less often solitary

Morphology Reproductive morphology Flowers Calyx

Calyx deeply divided into 5 (rarely 8–12) imbricate or open, equal or unequal lobes or 5 with appendages between

Morphology Reproductive morphology Flowers Corolla

Corolla often blue, variously shaped, the lobes usually 5 (rarely 8–12), imbricate or rarely contorted

Morphology Reproductive morphology Flowers 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

Morphology Reproductive morphology Flowers Nectaries

Disk inconspicuous or absent

Morphology Reproductive morphology Flowers 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

Morphology Reproductive morphology Fruits

Fruit usually a loculicidal or less often septicidal 2-valved or irregularly splitting capsule

Morphology Reproductive morphology Seeds

Seeds oblong, globose or angled, with tuberculate, reticulate or rugose testa; albumen copious and fleshy; embryo small; cotyledons entire

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

Morphology General Habit

Annual or perennial herbs, rarely subshrubs, often scabrid

Morphology Leaves

Leaves usually alternate or radical, exstipulate, entire to deeply pinnately or palmately lobed, with margins entire, serrate or lobulate

Morphology Reproductive morphology Flowers

Flowers regular, hermaphrodite, in scorpioid or trichotomous cymes, false racemes or clustered, less often solitary

Morphology Reproductive morphology Flowers Calyx

Calyx deeply divided into 5 rarely 8–12 imbricate or open equal or unequal lobes or 5 with appendages between

Morphology Reproductive morphology Flowers Corolla

Corolla often blue, variously shaped, the lobes usually 5, rarely 8–12, imbricate or rarely contorted

Morphology Reproductive morphology Flowers 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

Morphology Reproductive morphology Flowers Nectaries

Disk inconspicuous or absent

Morphology Reproductive morphology Flowers 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

Morphology Reproductive morphology Fruits

Fruit usually a loculicidal or less often septicidal bivalved or irregularly splitting capsule

Morphology Reproductive morphology Seeds

Seeds oblong, globose or angled with tuberculate reticulate or rugose testa; albumen copious and fleshy; embryo small; cotyledons entire