Plantaginaceae Juss.

Souza, V.C. (2012). Neotropical Plantaginaceae.

Morphology

Description

Annual, biennial or perennial herbs, shrubs or less frequently trees or climbers .  Leaves simple , opposite, alternate or less frequently whorled ; venation mainly pinnate or palmate ; margin entire , serrate or sometimes deeply divided ; without stipules.  Inflorescences cymose or racemose or more frequently flowers solitary in the axils of the upper leaves.  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 .  Fruit usually a septicidal , loculicidal, circumscissile or poricidal capsule .  Seeds numerous, minute, testa often ornamented, usually wind-dispersed.

General Description

Notes on delimitation

• 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.

Status

• 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.

General notes

• 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

Distribution in the Neotropics

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.

Diagnostic

Other important characters

• 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).

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.

Useful tips for generic identification

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

Notable genera and distinguishing features

• 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).

Literature

Important literature

Barringer, K. 1981. A revision of Angelonia (Scrophulariaceae). Ph.D. thesis. Univ. of ConecticutBarringer, K. 1983. Monopera, a new genus of Scrophulariaceae from South America Brittonia 35(2): 111-114.

Barringer, K. 1985. Revision of the genus Basistemon (Scrophulariaceae) Syst. Bot. 10(2): 125-133

Barringer, K. & Burger, W. 2000. Flora Costaricensis, Family 193 Scrophulariaceae. Fieldiana Botany 41: 1-69.

Barroso, G.M. 1952. Scrophulariaceae indígenas e exóticas do Brasil. Rodriguésia 15(27): 9-64.

Chodat, R. 1908. Étude critique des genres Scoparia L. et Hasslerella Chod. Bull. Herb. Boissier ser.2. 8: 1-16; 85-89.

D'Arcy, W.G. 1979. Flora of Panama: Scrophulariaceae. Annals of the Missouri Botanical Garden 66(2): 173-272.

Descole, H.R. & Borsini, O.E. 1954. Scrophulariaceae: Antirrhinoideae.  In: Descole, H.R. (ed.). Genera et species plantarum argentinarum - opus quod in ordinen redegit et direxit Horatius R. Descole, adiuvante personali Institutionis Michaelis Lillo Universitati nationali tucumanensi annexae, et cooperantibus botanicis argentinis et extraneis in capite totius familiae citatis vol. 5(1), pp. 3-164.  Guillermo Kraft ltda., Buenos Aires.

Díaz-Miranda, D. 1988. Genera of Scrophulariaceae in the paramos of the Venezuelan Andes. Brittonia 40(2): 235-239.

Elisens, W.J. 1985. Monograph of the Maurandyinae (Scrophulariaceae--Antirrhineae). Syst. Bot. Monogr. Vol. 5, pp. 1-97.  The American Society of Plant Taxonomists, Ann Arbor.

Fassett, N.C. 1951. Callitriche in the New World. Rhodora 53(630): 137-155.

Fassett, N.C. 1951. Callitriche in the New World [cont. 2]. Rhodora 53(631): 161-182.

Fassett, N.C. 1951. Callitriche in the New World [cont. 3]. Rhodora 53(632): 185-194.

Fassett, N.C. 1951. Callitriche in the New World [cont. 4]. Rhodora 53(633): 209-221.

Fernandez-Alonso, J.L. 1993. Novedades taxonómicas en Aragoa H.B.K. (Scrophulariaceae) y sinopsis del género. Anales Jard. Bot. Madrid 51: 73-96.

Fischer, E. 2004. Scrophulariaceae. In: K. Kubitzki (ed.) The Families and Genera of Vascular Plants vol.7: 333-432. Springer-Verlag, Berlin, Heidelberg.

Fries, R.E. 1906. Systematische Übersicht der Gattung Scoparia. Ark. Bot. 6(9): 1-31.

Fries, R.E. 1908. Einige weitere Bermerkungen über die Gattung Scoparia. Bull. Herb. Boissier ser. 2. 8: 934-940.

Holmgren, N.H. & Molau, U. 1984. 177. Scrophulariaceae vol. 21: 1-189. In: G.W. Harling & B.B. Sparre (eds.). Fl. Ecuador. University of Göteborg and Swedish Museum of Natural History, Göteborg and Stockholm.

Jørgensen, P.M. 1999. Scrophulariaceae. In: Jørgensen, P.M. & S. León-Yañes (eds.) Catalogue of the vascular plants of Ecuador. Monogr. Syst. Bot. Missouri Bot. Gard vol. 75, pp. 891-899.  Missouri Botanical Garden, St. Louis.

Kuhlmann, J.G. & Brade, A.C. 1943.  Contribuição para o conhecimento do Gênero Otacanthus. Fam. Scrophulariaceae. Arq. Serv. Florestal Rio de Janeiro 2(1): 17-20.

Larios, I.M. & Villaseñor, J.L. 1995. Revisión taxonômica del género Tetranema (Scrophulariaceae). Acta Bot. Mexicana 32: 53-68

Marticorena, C. & Quezada, M. 1985. Catálogo de la Flora Vascular de Chile. Gayana, Bot. 42: 1-157.

Méndez-Larios, I. & Villaseñor-Rios, J.L. 2001. La família Scrophulariaceae in México: diversidad y distribución. Bol. Soc. Bot. México 69: 101-121.

Meudt, H.M. 2006. Monograph of Ourisia (Plantaginaceae). Syst. Bot. Monogr. Vol. 77, 188 pp.  The American Society of Plant Taxonomists, Ann Arbor.

Minod, M. 1918. Contributions à l'etude du genre Stemodia et du groupe des Stemodiees en Amerique. Bull. Soc. Bot. Genève, ser. 2. 10: 155-252.

Olmstead, R.G. & Reeves, P.A. 1995. Evidence for the polyphyly of the Scrophulariaceae based on chloroplast rbcL and ndhF sequences. Annals of the Missouri Botanical Garden 82: 176-193.

Olmstead, R.G., Depamphilis, C.W., Wolfe, A.D., Young, N.D., Elisons, W.J. & Reeves, P.A. 2001. Disintegration of the Scrophulariaceae. Amer. J. Bot. 88: 348-361.

Oxelman, B., Kornhall, P., Olmstead, R.G. & Bremer, B. 2005. Further disintegration of Scrophulariaceae. Taxon 54(2): 411-425.

Pennell, F.W. 1920. Scrophulariaceae of Colombia. Proc. Acad. Nat. Sci. Philadelphia 72: 136-188.

Pennell, F.W. 1938. Taxonomy and distribution of Aragoa and its bearing on the geological history of the Northern Andes. Proc. Acad. Nat. Sci. Philadelphia 89: 425-432.

Pennell, F.W. 1946. Reconsideration of the Bacopa-Herpestis problem of the Scrophulariaceae. Proc. Acad. Nat. Sci. Philadelphia 98(2): 83-98.

Pennell, F.W. 1952. The Genus Achetaria (Scrophulariaceae) of Lowland South America. Not. Nat. 244: 1-4.

Pereira, C.G., Almenara, D.P., Winter, C.E., Fritsch, P.W., Lambers, H. & Oliveira, R.S. 2012. Underground leaves of Philcoxia trap and digest nematodes. Proc. Nat. Acad. Sc. United States of America 109(4): 1154-1158.

Rahmanzadeh, R., Müller, K., Fischer, E., Bartels, D. & Borsch, T. 2005. The Linderniaceae and Gratiolaceae are further lineages distinct from the Scrophulariaceae (Lamiales). Plant. Biol. 7(1): 67-78.

Rahn, K. 1981. Plantago ser. Sericeae, a taxonomic revision. Nordic J. Bot. 1(3): 297-323.

Rios, E.S. 1981. Contribuição ao conhecimento das espécies brasileiras do gênero Achetaria Cham. & Schlecht. Tese de Mestrado. UFRJ. Rio de Janeiro.

Ronse, A. 2001. A Revision of Otacanthus Lindl. (Scrophulariaceae). Brittonia 53(1): 137-153.

Rossow, R.A. 1985. Maeviella, nuevo género de Scrophulariaceae de Brasil. Arq.. Jard. Bot. Rio de Janeiro 27: 171-175.

Rossow R.A. 1986. Sinopsis de las especies austroamericanas del genero Ourisia (Scrophulariaceae). Parodiana 4(2): 239-265.

Rossow, R.A. 1987. Revisión del género Mecardonia (Scrophulariaceae). Candollea 42(2): 431-474. Souza, V.C. 2003. Scrophulariaceae. In: Wanderley, M.G.L., G.J. Shepherd, T.S. Melhen, A.M. Giulietti, M. Kirizawa (eds.), Flora Fanerogâmica do Estado de São Paulo vol. 3: 297-321. São Paulo. FAPESP, Rima.

Souza, V.C. 1997. Considerações sobre a delimitação de Mecardonia procumbens (Mill.) Small (Scrophulariaceae). Acta Botanica Brasilica: 11(2): 181-189.

Souza, V.C. 2001. Anamaria: um novo gênero de Scrophulariaceae do Brasil. Bol. Bot. Univ. São Paulo 19: 43-44.

Souza, V.C. & Giulietti, A.M. 1990. Scrophulariaceae de Pernambuco. Bol. Bot. Univ. São Paulo 12(1): 185-209.

Souza, V.C. & Giulietti, A.M. 2003. Flora da Serra do Cipó, Minas Gerais: Scrophulariaceae. Bol. Bot. Univ. São Paulo 21(2): 283-297.

Souza, V.C. & Giulietti, A. M. 2009. Levantamento das espécies de Scrophulariaceae s.l. nativas do Brasil. Pesq. Bot. 60: 1-288

Souza, V.C. & Souza, J.P. 1997. Flora de Goiás e Tocantins - Coleção Rizzo: Scrophulariaceae. Goiânia. Universidade Federal de Goiás.

Standley, P.C. & Williams, L.O. 1973. Scrophulariaceae. In: Standley, P.C. & Williams, L.O. (eds.), Flora of Guatemala Part IX, Number 4. Fieldiana, Bot. 24(9/4): 319-416.

Sutton D.A. 1988. A revision of the tribe Antirrhineae. British Museum (Natural History) & Oxford University Press: London & Oxford. 575 pp.

Tank, D.C., Beardsley, P.M., Kelchner, S.A. & Olmstead, R.G. 2006. L.A.S. Johnson Review No. 7. Review of the systematics of Scrophulariaceae s.l. and their current disposition. Australian Syst. Bot. 19: 289-307.

Taylor, P.; Souza, V.C.; Giulietti, A.M.; Harley, R.M. 2000. Philcoxia: A new genus of Scrophulariaceae with three new species from Eastern Brazil. Kew Bulletin 55: 155-163.

Thieret, J.W. 1967. Supraspecific classification in the Scrophulariaceae: a review. Sida 3(2): 87-106.

Turner, B.L. & Cowan, C.C. 1993. Taxonomic overview of Stemodia (Scrophulariaceae) for South America. Phytologia 75(4): 281-324.