Plantaginaceae Juss.

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

Morphology General Habit

Annual, biennial or perennial herbs, shrubs or less frequently trees or climbers

Morphology Leaves

Inflorescences cymose or racemose or more frequently flowers solitary in the axils of the upper leaves Leaves simple, opposite, alternate or less frequently whorled; venation mainly pinnate or palmate; margin entire, serrate or sometimes deeply divided; without stipules

Morphology Reproductive morphology Flowers

Flowers mostly showy, naked or with calyx and corolla, bisexual or rarely unisexual; calyx with sepals fused or free, equal or unequal, 4-5- lobed; corolla with petals fused, zygomorphic, commonly bilabiate, sometimes actinomorphic (Plantago), generally 5- lobed; stamens (1-)2-4, sometimes with a staminode, generally didynamous (two pairs of unequal length), alternating with corolla lobes, anthers dithecal or monothecal, opening by longitudinal slits; gynoecium syncarpous of 2 carpels, rarely 1 (Hippuris), ovariy superior, rarely inferior (Hippuris), locules as many as carpels, style generally 1, ovules 1-many, placentation axile, rarely basal

Morphology Reproductive morphology Fruits

Fruit usually a septicidal, loculicidal, circumscissile or poricidal capsule

Morphology Reproductive morphology Seeds

Seeds numerous, minute, testa often ornamented, usually wind-dispersed.

Note

Plantaginaceae is a highly heteromorphic group, including both genera and species with naked flowers and flowers with a double perianth, which is unusual in the Lamiales. Previous studies (summarized by Olmstead et al., 2001 and Tank et al., 2006) revealed the need for recircumscribing this family in a broader sense to include previously recognized families such as Hippuridaceae, Callitrichaceae and a significant part of Scrophulariaceae. The evolutionary relationships (and even their limits) among the families  of Lamiales are still poorly known.  Ongoing phylogenetic studies suggest that some additional changes are expected, including the acceptance of Gratiolaceae as proposed by Rahmanzadeh (2005) and consequently a separate family for the members of Angelonieae. Number of genera: Plantaginaceae have a cosmopolitan distribution and include about 100 genera and 2,000 species.  Of these, 45 genera and 370-400 species are native in the Neotropics. The distributional limit of Collinsia Nutt. and Holmgrenanthe Elisens. is adjacent to the Neotropics in southern USA Additionally, Melosperma Benth. and Monttea C. Gay. are endemic to the Central and South Andes in Chile and Argentina, but not present in the Neotropics.

Distribution

Throughout the Neotropics, most common in savannas and open fields.  Achetaria Cham. & Schltdl. (including Otacanthus Lindl.): (8 species) from Venezuela and Guianas to southern Brazil, mainly in eastern Brazil. Ameroglossum Eb. Fisch., S. Vogel & A.V. Lopes: (1 species) NE Brazil. Anamaria V.C. Souza: (1 species) NE Brazil. Angelonia Bonpl.: (25-30 species) Mexico to Argentina, mainly in NE Brazil. Aragoa Kunth: (20 species) Colombia and Venezuela, endemic to the paramos. Bacopa Aubl. (including Benjaminia Mart. Ex Benj. and Maeviella Rossow: (40-60 species) pantropical, but nearly all species native in the Neotropics. Basistemon Turcz.: (4-8 species) South America, especially western extra-Andean Peru and Bolivia. Callitriche L.: (60-65 species, 10-15 in Neotropics) cosmopolitan, the Neotropical species scattered across a wide range of habitats. Conobea Aubl.: (3 species) Mexico to Argentina. Cymbalaria Hill: (9 species, 1 Neotropical) northern Mediterranean, southern Alps, naturalized elsewhere, including in the Neotropics. Dizygostemon (Benth.) Radlk. ex Wettst: (1 species) NE Brazil. Galvezia Dombey ex Juss.: (4 species) Ecuador and Peru, mainly near the coast: (2 species endemic to the Galapagos Islands). Gambelia Nutt.: (4 species, all Neotropical) southern USA and Mexico. Gratiola L.: (25-30 species, 5 Neotropical) cosmopolitan, but mainly in temperate North America. Hebe Comm. ex Juss.: (80-90 species, 1 species naturalized in the Neotropics) mainly in New Zealand; 2 species in Chilean Patagonia. Hippuris L.: (2-3 species) worldwide, mainly in temperate zones; 1 species native in Patagonia. Ildefonsea Gardner: (1 species) endemic to SE Brazil (Rio de Janeiro state). Keckiella Straw: (7 species, 3 in the Neotropics) USA to Mexico. Kickxia Dumort.: (45-50 species, 2 naturalized in the Neotropics) Africa, Europe and Asia. Limosella L.: (5-10 species, 2-3 species in the Neotropics) Africa and the Americas. Linaria Mill.: (150 species, 3 naturalized in the Neotropics, mainly in Mexico) Europe, Asia and Africa. Lophospermum D. Don (including Rhodochiton Zucc. Ex Otto): (20 species) Mexico and Guatemala. Mabrya Elisens: (5 species, 4 in the Neotropics) southern USA to Mexico. Maurandya Ortega (including Epixiphium (Engelm. Ex A. Gray) Munz and Maurandella (A. Gray) Rothm.: (4 species, all in the Neotropics) USA to Peru. Mecardonia Ruiz & Pav.: (5-10 species, all in the Neotropics) USA. to Argentina. Misopates Rafin.: (7 species, 1 naturalized in the Neotropics) Europe, Asia, Africa. Mohavea A. Gray: (2 species, 1 in the Neotropics) southern USA and NW Mexico. Monopera Barringer: (2 species) disjunct distribution (1 in Paraguayan Chaco and Brazilian Pantanal and 1 in NE Brazil). Neogaerrhinum Rothm.: (2 species, 1 in the Neotropics) southern USA and NW Mexico. Nuttallanthus D.A. Sutton: (4 species, 3 in the Neotropics) Canada, USA and western South America, naturalized elsewhere. Ourisia Comm. ex Juss.: (28 species, 5 in north-central Andes and 10 in extra-tropical Andes) New Zealand, Tasmania and Andes. Penstemon Schmidel: (270 species, 55-60 Neotropical) North America, Mexico, and Guatemala. Philcoxia P. Taylor & V.C. Souza: (3 species) endemic to the highlands of Central and NE Brazil. Plantago L. (including Bougueria Decne: (270 species, 40-50 Neotropical, including some naturalized) cosmopolitan. Pseudorontium (A. Gray) Rothm.: (1 species) southern USA and NW Mexico. Russelia Jacq.: (50-55 species) Mexico to Colombia. Sairocarpus D.A. Sutton: (13 species, 5 Neotropical) SW North America. Schistophragma Benth. ex Endl.: (3 species, all in Neotropics) southern USA to Colombia. Scoparia L.: (20 species) tropical America, 1 species a pantropical weed. Sibthorpia L.: (5-6 species, 2 Neotropical) Africa, and Mexico to Bolivia. Stemodia L.: (56 species, 32 Neotropical) Pantropical. Tetranema Benth.: (4 species) Mexico and Central America. Tetraulacium Turcz.: (1 species) Brazil, mainly NE. Uroskinnera Lindl.: (4 species) Mexico and Central America. Veronica L.: (450-500 species, c. 10 species naturalized in the Neotropics and at least 1 native) worldwide, but concentrated in temperate zones. Many genera of the Plantaginaceae are used as ornamentals worldwide, including coral-plant (Russelia), beardtongue (Penstemon), hebe (Hebe) and snapdragon (Antirrhinum L.). Some species of Plantaginaceae have a pantropical distribution, including some weeds, such as Scoparia dulcis L. and Bacopa monnieri (L.) Wettst.  Many species native to the northern Hemisphere are weeds in the Neotropics.  These include species of Linaria, Plantago and Veronica. Nearly half of the genera occurring in the Neotropics are endemic.

Diagnostic

Key differences from similar families: The high level of infrafamiliar morphological variation and the many characters shared with other families such as Verbenaceae, Bignoniaceae, Acanthaceae and Scrophulariaceae s.s. make it hard to summarize the morphological differences between Plantaginaceae and other families of the Lamiales. The characters listed above are constant in Plantaginaceae, but they are also present in almost all Lamiales families. Mostly herbs or shrubs. Leaves usually opposite. Corolla usually zygomorphic- Ovules usually numerous. Fruit usually a capsule. Ovary mainly superior. Carpels mainly 2. Distinguishing characters (always present): Simple leaves. Stipules absent. Corolla with fused petals (if present). Anamaria V.C. Souza is an endemic, monoespecific genus from the caatinga in semi-arid Brazil.  Despite occurring in a semi-arid region, Anamariaheterophylla (A.M. Giulietti & V.C. Souza) it is an aquatic plant with vegetative branches with floating leaves.  There is no reason to accept the synonimization of Anamaria under Stemodia, as proposed by some authors.  First of all, there is no phylogeny that shows Stemodia to be monophyletic, and any synonymization therefore ought to wait for the results of a study that samples all Stemodia and Anamaria species.  Secondly, there are many morphological characters that distinguish the two genera even though they share a connective splitting the anther thecae (a rare character also shared with Mecardonia). Ildefonsia Gardner is a monoespecific genus, known only from remnant forests in the city of Rio de Janeiro.  Some authors previously synonymised this genus under Bacopa, but since Ildefonsia does not share the main characters of Bacopa, especially calyx and anthers characters, it should be kept separate. Nuttallanthus D.A. Sutton is a recently described genus, which includes some New World species previously placed in Linaria.  One of the species (Nuttallanthussubandinus (Diels) D.A. Sutton) is native in the Andes and the other two (N. canadensis (L.) D.A.Sutton and N. texanus (Scheele) D.A. Sutton) probably only naturalized in the Neotropics. Philcoxia P. Taylor & V.C. Souza is a recently described genus, which includes three also recently described species, from the Brazilian highlands.  The aerial vegetative parts of these species include only the leaf lamina, which is only a few millimetre in diameter and looks similar to a sand grain.  For this reason the plants are virtually impossible to find in their natural habitat unless they are flowering.  Recent studies additionally showed that Philcoxiaminensis is carnivorous, trapping and digesting nematodes from the soil (Pereira et al., 2012). Veronica L. is present mainly in the northern Hemisphere, with some species naturalized worldwide.  Despite being the most speciose genus in Plantaginaceae, there is only one native species in the Neotropics (V. mexicana S. Watson). Interactive key to the Latin American Genera of Plantaginaceae - follow link http://www.kew.org/science/tropamerica/neotropikey/families/keys/plantaginaceae/index.htm Dichotomous key to genera of Neotropical Plantaginaceae Key based mainly in Neotropical species (since some genera has a much broader distribution). Morphological distinction between some genera is not yet enoughly clear, especially the ones restricted to Neotropics. A broad phylogenetic and morphological analysis is required. 1.  Flowers very reduced, without corolla (naked) — 2 1.  Flowers with corolla — 3 2.  Leaves opposite — Callitriche 2.  Leaves verticillate — Hippuris 3.  Plants with rhizome and petioles completely underground; leaves peltate — Philcoxia 3.  Vegetative shoots aerial; leaves not peltate — 4 4.  Plants with two kinds of branches arising from the base of the plant: vegetative with broad leaves and reproductive with more narrow leaves — Anamaria 4.  Plants without differentation between vegetative and reproductive branches — 5 5.  Flower with a spur — 6 5.  Flower without a spur (sometimes gibbous) — 9 6.  Anthers fused, forming a ring-like structure — Kickxia 6.  Anthers free or, if fused, not forming a ring-like structure — 7 7.  Leaves palmately veined— Cymbalaria 7.  Leaves pinnately veined — 8 8.  Lower corolla lip about twice as large as the upper lip — Nuttallanthus 8.  Lower corolla lip smaller or more or less equal to the upper lip — Linaria 9.  Flowers in spikes, not showy and wind-pollinated — Plantago 9.  Flowers not in spikes or, if so, then showy and insect- or bird-pollinated — 10 10.  Staminode almost as long as (or longer than) fertile stamens — 11 10.  Staminode smaller or absent — 13 11.  Flowers with a hypogynous disc— Keckiella 11.  Flowers without a hypogynous disc— 12 12.  Herbs to shrubs; capsulesepticidal; endemic to North America, Mexico, and Guatemala — Penstemon 12.  Shrubs; capsule loculicide; endemic to Central America — Uroskinnera 13.  Calyx with fused sepals (sometimes sepals connate near the base) — 14 13.  Calyx with free sepals — 31 14.  Stamens 2, plus 3 staminodes (2 of which with rudimentary anthers) — Mohavea 14.  Stamens 4, usually with 1 staminode (sometimes with a rudimentary anther) — 15 15.  Calyx longer or sometimes equal to the corolla tube — 16 15.  Calyx shorter than the corolla tube — 18 16.  Corolla almost actinomorphic— Limosella 16.  Corolla strongly zygomorphic— 17 17.  Lower corolla lip larger than upper lip; medium lobe of lower lip larger than the others — Sairocarpus 17.  Lower corolla lip more or less equal to upper lip; medium lobe of lower lip much narrower than the others — Misopates 18.  Leaves all alternate, palmately veined— 19 18.  Leaves opposite or verticillate, at least at the base of the plant, usually pinnately veined — 23 19.  Pedicels cirrhous, elongate, more than 8 times longer than flowers — Neogaerrhinum 19.  Pedicels not cirrhous or, if cirrhous, then much shorter than flowers — 20 20.  Climbing plants — 21 20.  Erect or creeping plants — 22 21.  Calyx lobes unequal; seeds not compressed or winged— Maurandya 21.  Calyx lobes subequal; seeds compressed, with 2 wings — Lophospermum 22.  Seeds not compressed or winged— Mabrya 22.  Seeds compressed, with 2 wings — Lophospermum 23.  Flowers solitary — 24 23.  Flowers in inflorescences — 29 24.  Corolla personate, with the lower lip closing the corolla mouth — 25 24.  Corolla not personate; the corolla mouth open — 27 25.  Shrubs; corolla usually red, rarely white — Gambelia 25.  Herbs; corolla usually violet, purple to blue, rarely white — 26 26.  Lower corolla lip larger the than upper lip — Sairocarpus 26.  Lower corolla lip more or less equal to the upper lip — Pseudorontium 27.  Creeping herbs — Ourisia 27.  Erect shrubs — 28 28.  Capsule wall thin, opening irregularly; staminode with rudimentary anther— Galvezia 28.  Capsule wall thick, opening in regular places; staminode without rudimentary anther— Gambelia 29.  Leaves usually in a basal rosette; inflorescence umbelliform — Tetranema 29.  Leaves not in a basal rosette; inflorescence not umbelliform — 30 30.  Stamens exserted; lower lip of the corolla slightly 3-partite; seeds not covered in trichomes — Ameroglossum 30.  Stamens included; lower lip of the corolla distinctly 3-lobed; seeds covered in long trichomes — Russelia 31.  Fertile stamens 2 — 32 31.  Fertile stamens 4 or 5(-8) — 36 32.  Staminodes 2 or 3 — 33 32.  Staminodes absent — 34 33.  Anthers 1-thecate — Achetaria 33.  Anthers 2-thecate — Gratiola 34.  Calyx with unequal sepals — Bacopa 34.  Calyx with equal sepals — 35 35.  Shrubs — Hebe 35.  Herbs — Veronica 36.  Anthers with well-developed connective, splitting the thecae — 37 36.  Anthers parallel or divergent, connective not well-developed — 38 37.  Calyx with unequal sepals; corolla yellow — Mecardonia 37.  Calyx with equal sepals; corolla blue, lilac, purple, sometimes with small, yellow parts — Stemodia 38.  Anthers divergent — 39 38.  Anthers with one theca withered or both fertile and parallel — 42 39.  Shrubs to trees; ovary locules with 1-12 ovules — 40 39.  Herbs to shrubs; ovary locules with many more ovules — 41 40.  Leaves fleshy, overlapping each other; petals 4 — Aragoa 40.  Leaves not fleshy, nor nor overlapping each other; petals 5 — Basistemon 41.  Corolla bigibbous — Angelonia 41.  Corolla unigibbous — Monopera 42.  Leaves alternate — Sibthorpia 42.  Leaves opposite or verticillate — 43 43.  Calyx with unequal sepals — 44 43.  Calyx with equal sepals — 46 44.  Anthers with both thecae fertile — Bacopa 44.  Anthers unithecal or with one theca withered — 45 45.  All anthers with one theca withered — Tetraulacium 45.  Dorsal anthers with one theca withered, ventral anthers unithecal — Dizygostemon 46.  Corolla 4-lobed, wheel-shaped — Scoparia 46.  Corola 5-lobed, tubular — 47 47.  Corolla red — Galvezia 47.  Corola white, blue, purple or lilac — 48 48.  Leaves deeply pinnatifid to pinnatisect — Schistophragma 48.  Leaves entire — 49 49. Staminode present — Ildefonsia 49.  Staminode absent — Conobea

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

Morphology General Habit

Herbs

Morphology Leaves

Leaves all radical or alternate or opposite, simple, sometimes much reduced

Morphology Reproductive morphology Flowers

Flowers usually hermaphrodite, spicate, actinomorphic

Morphology Reproductive morphology Flowers Calyx

Calyx herbaceous

Morphology Reproductive morphology Flowers Corolla

Corolla gamopetalous, scarious, 3-4-lobed, lobes imbricate

Morphology Reproductive morphology Flowers Androecium

Stamens usually 4, inserted on the corolla-tube and alternate with the lobes or hypogynous; anthers 2-celled, opening lengthwise

Morphology Reproductive morphology Flowers Gynoecium

Ovules 1 or more in each cell, axile or basal Ovary superior, 1-4-celled; style simple

Morphology Reproductive morphology Fruits

Fruit a circumscissile capsule or nut

Morphology Reproductive morphology Seeds

Seeds peltately attached, with fleshy endosperm and straight or curved embryo

Plantaginaceae, B. Verdcourt. Flora of Tropical East Africa. 1971

Morphology General Habit

Animal or perennial, terrestrial or, in one small genus, aquatic herbs

Morphology Leaves

Leaves mostly spirally arranged and all radical, less often cauline, alternate or opposite, simple, often sheathing at the base, sometimes reduced, exstipulate

Morphology Reproductive morphology Flowers

Flowers small, hermaphrodite, or in one small genus unisexual, regular, sessile in the bract-axils, forming axillary spikes or capitate racemes

Morphology Reproductive morphology Flowers Calyx

Calyx herbaceous, 4-lobed, the lobes ± free or sometimes the lower pair ± united

Morphology Reproductive morphology Flowers Corolla

Corolla gamopetalous, scarious, 3-4-lobed, the lobes imbricate

Morphology Reproductive morphology Flowers Androecium

Stamens (l-2-)4, inserted in the corolla-tube and alternating with the lobes, rarely hypogynous; filaments usually long; anthers versatile, 2-thecous, the thecae opening lengthwise

Morphology Reproductive morphology Flowers Gynoecium

Ovary superior, 1-4-locular; style long, simple, often exserted; ovules l-? in each locule, axile or basal

Morphology Reproductive morphology Fruits

Fruit a membranous circumscissile capsule or a 1-seeded bony nut

Morphology Reproductive morphology Seeds

Seeds peltately attached, often sticky when wet; embryo straight, rarely curved, placed in the middle of the fleshy endosperm

Plantaginaceae, G. Lehmann. Flora Zambesiaca 9:1. 1988

Morphology General Habit

Annual or perennial, terrestrial or aquatic (not in Flora Zambesiaca area) herbs

Morphology Leaves

Leaves mostly spirally arranged and all radical, less often cauline and spiral, alternate or opposite, simple, often sheathing at the base, sometimes reduced, exstipulate

Morphology Reproductive morphology Inflorescences

Inflorescence usually a spike, rarely capitate racemes, terminating a scape. Each flower subtended by a bract

Morphology Reproductive morphology Flowers

Flowers small, hermaphrodite or unisexual (not in Flora Zambesiaca area), usually 4-merous, actinomorphic or rarely so, sessile, bracteate

Morphology Reproductive morphology Flowers Calyx

Calyx herbaceous, persistent, 4-lobed, the lobes more or less free or sometimes the lower pair more or less united

Morphology Reproductive morphology Flowers Corolla

Corolla gamopetalous, scarious, 3–4 lobed; tube ampulliform or cylindric, lobes imbricate, equal, spreading

Morphology Reproductive morphology Flowers Androecium

Stamens (1–2) 4, inserted in the corolla tube and alternating with the leaves, rarely hypogynous, usually exserted; filaments usually long, filiform; anthers versatile, 2-thecous, the thecae dehiscing longitudinally

Morphology Reproductive morphology Flowers Gynoecium

Ovary superior, 1–4-locular, with 1-many ovules in each locule, axile or basal; style single, long, usually filiform, simple, often exserted

Morphology Reproductive morphology Fruits

Fruit a capsule, membranous, circumscissile or 1-seeded, hard, indehiscent

Morphology Reproductive morphology Seeds

Seeds peltately attached, often mucilaginous when wet, 1-many; embryo straight, rarely curved, enclosed by the fleshy endosperm

Gemma Bramley, Anna Trias-Blasi & Richard Wilford (2023). The Kew Temperate Plant Families Identification Handbook. Kew Publishing Royal Botanic Gardens, Kew

Recognition

Characters of similar families: Scrophulariaceae s.s.: primarily southern hemisphere; (sub)actinomorphy more common. Lamiaceae: up to 4 nutlets, or a drupe with 1–4 seeds. Orobanchaceae: hemi- or holoparasites.

Morphology General Habit

Herbs, sometimes shrubs; terrestrial, rarely aquatic; autotrophic; hairs simple, often glandular, with gland head not vertically divided; stipules absent

Morphology Leaves

Leaves alternate and spiral, opposite or basally opposite, apically alternate, usually simple, cauline or basal; margins entire to subentire, toothed or lobed

Morphology Reproductive morphology Inflorescences

Inflorescence variable; sometimes flowers solitary

Morphology Reproductive morphology Flowers

Flowers bisexual, occasionally monoecious or dioecious; zygomorphic, less often actinomorphic, rarely reduced; calyx lobes fused, 4–5-merous; corolla usually 5–(8)-merous, can appear 4-merous due to the fusion of the upper lobes, often 2-lipped, tubular to various degrees, sometimes with basal nectar spur or personate; stamens (2)4, didynamous to equal, adnate to corolla, 0–1(–3) staminode(s) present; anthers 2-locular, sacs divergent, opening by 2 longitudinal slits, or a single inverted slit if apical portion of anther sacs adnate; ovary superior, 2-carpellate, carpels fused, placentation axile, ovules (1)–numerous, anatropous or hemitropous; stigma bilobed or capitate

Morphology Reproductive morphology Fruits

Fruit a capsule, usually septicidal or loculicidal, sometimes poricidal or circumsessile

Morphology Reproductive morphology Seeds

Seeds (1)–numerous, winged or angular.

Distribution

At least 90 genera and 1,900 species; mostly in the northern hemisphere. Plantaginaceae expanded significantly following the disintegration of the large and polyphyletic Scrophulariaceae.

Note

Leaves alternate and spiral or opposite; usually simple. Flowers bisexual, zygomorphic, 2-lipped, occasionally actinomorphic; stamens (2)–4; ovary superior, 2 fused carpels, ovules numerous, placentation axile. Fruit a capsule.

Description Author

Gemma Bramley

Callitrichaceae, Inga Hedberg & Olov Hedberg. Flora of Tropical East Africa. 2003

Morphology General Habit

Creeping or submerged, monoecious, aquatic herbs with filiform stems and opposite, entire exstipulate leaves

Morphology Stem

Stems with axillary glandular scales; both stem and leaves with peltate hairs

Morphology Reproductive morphology Flowers

Flowers axillary, either solitary or with one male and one female flower in the same axil; perianth absent; bracteoles 2, crescent-shaped or absent; male flower consisting of one stamen with a slender filament and a reniform anther; female flower consisting of a single carpel with a 4-celled ovary containing a single anatropous ovule in each locule; styles 2, filiform

Morphology Reproductive morphology Fruits

Fruit schizocarpic, separating into 4 mericarps

Morphology Reproductive morphology Seeds

Seeds with a fleshy endosperm

Use

Veronica, Russelia, Angelonia and Penstemon are popular ornamentals; Plantago is unusual with ±parallel venation and wind-pollinated flowers; Callitriche are aquatics; Antirrhinum is a model organism for floral development; and Digitalis species are used medicinally. Several species are noxious weeds.