Scrophulariaceae

Disclaimer

Content is from Kirkbride et al. 2006, without modification. 
Updates are forthcoming.

Taxonomy

Scrophulariaceae A.L. Jussieu

Common name: Figwort Family.

Number of genera: 276 genera (including Globularia & Poskea).

Number of species (Mabberley 1997): 4650 species.

Disseminule

Fruit (dehisced), or fruit (intact or entire), or seed.

Description

Fruits: Pistil(s) simple, or compound; 1; 1-pistillate; with carpels united. Fruit pericarpium; simple, or schizocarp; capsule, or berry (not Spjut); achenarium (Globularia, Poskea; unknown in Lagotis); denticidal capsule, or fissuricidal capsule, or foraminicidal capsule (Spjut Fig. 32B), or loculicidal capsule, or septicidal capsule, or poricidal capsule (3 families: Loganiaceae, Saxifragaceae, Scrophulariaceae & in this family opening by valves, teeth, irregular ruptures of operculum, forming 1–2 pores); capsule not inflated, or inflated; capsule without operculum; berry indehiscent; berry without central placental mass; without persistent central column; valves not diverging at top of central column; with styles(s); at apex; within accessory organ(s); within calyx; accrescent; persistent; soft calyx; many-seeded; many; with 2-carpellate; with carpels united; with carpels remaining united at maturity; with carpels not radiating at maturity; with carpels remaining connected at style, or separating at style; without sterile carpels; not sulcate; in transection terete; apex not beaked; wall membranaceous (at least); dehiscent, or indehiscent. Dehiscent unit seed(s). Dehiscent passively, or actively; and shedding seeds; without replum. Epicarp brown (all shades); dull; durable; membranous (Globularia); glabrous, or not glabrous (with hairs); hairs short; hairs dense, or scattered; hairs white; hairs not glandular; without armature; smooth; without wing(s); without apical respiratory hole. Mesocarp absent, or present. Endocarp present, or absent; not separating from exocarp; thin; not splitting into 1-seeded pyrenes; smooth; without wing; without operculum; without secretory cavities; without mechanism for seedling escape; without grooves; without longitudinal ridges. Funiculus short; short without seed bearing hooks (retinacula); not persisting in fruit after seed shed.

Seeds: Aril absent. Seed minute, or larger than minute; less than 1 mm long to 1 to less than 5 mm long; 0.5–3.5 mm long; irregular, or oval, or obovoid, or circular, or linear; in transection terete, or compressed, or flattened; bowl shaped, or not bowl shaped; not nutlike; without winglike beak; without caudate appendage(s); at maturity with food reserves, or without food reserves (rarely), or without apparent food reserves; with endosperm; without canavanine. Sarcotesta absent. Testa present, or absent; adnate to epicarp; without embryo surrounded and capped by viscid tissue; with markedly different marginal tissue, or without markedly different marginal tissue; marginal tissue winglike; without fleshy or leathery layer over hard layer; tight; surface unsmooth, or smooth; surface with depressed features, or discreet raised features, or merged raised features; surface punctate; surface granular, or papillate, or tuberculate; surface reticulate, or rugose, or striate; with crease or line separating cotyledons from hypocotyl-radicle, or without crease or line separating cotyledons from hypocotyl-radicle; with notch along margin where cotyledons from hypocotyl-radicle tip approaching each other, or without notch along margin where cotyledons from hypocotyl-radicle tip approach each other; without glands; without bristles; glabrous; without wings, or with wing(s); 1-winged; with wing encompassing seed; with wing(s) solid; with solid wing(s) similar to testa; without collar; without operculum; colored; monochrome; brown (all shades), or black; crustaceous; not becoming mucilaginous when wetted; surrounding endosperm, or surrounding embryo. Endosperm development cellular, or nuclear; copious; fleshy, or cartilaginous; smooth; with oils; without fatty acid containing cyclopropene; without apical lobes; without chlorophyll; without isodiametric faceted surface; without odor. Embryo differentiated from food reserve; well developed; 1 per seed; completely filling testa (no food reserve), or nearly filling testa (trace or scanty food reserve), or partially filling testa (with food reserve); chamber central to wings; 0.4–1 times the length of food reserve; at one end of seed not extending into a depression or cup; axile and centric; foliate, or linear, or miniature; with spatulate cotyledons; straight, or C-shaped, or annular (rarely); parallel to seed length; embedded in endosperm; with cotyledons abruptly connected to hypocotyl-radicle, or gradually connected to hypocotyl-radicle; without coleorhiza; without simmondsin; without stomata; not green (but may be violet); with 2 or more cotyledons, or acotyledonous (Lathraea). Cotyledons 2; tiny, or moderately developed, or well developed; 0.1–0.7 times length of embryo; as wide as hypocotyl-radicle, or somewhat to significantly wider than hypocotyl-radicle; 1–3 times wider than hypocotyl-radicle; partially concealing hypocotyl-radicle; thin; flat; smooth; with apices entire; with margins separate; basally entire; equal in size; not punctate dotted. Hypocotyl-radicle moderately developed, or well developed; straight, or curved; not thickened.

Habitat and crop association

Noxious weeds: 1 or more USA Federal noxious weeds in this family and 1 or more USA state noxious weeds in this family.

USA Federal Noxious Weeds: 
Alectra spp.: USA Federal Noxious Weed●; USA state noxious weed: FL●. 
Alectra vogelii Benth.: USA Federal Noxious Weed●. 
Limnophila sessiliflora (Vahl) Blume: USA Federal Noxious Weedª●; USA state noxious weed: ALª, FLª, NCª, SCª. 
Striga angustifolia (D. Don) C. J. Saldanha: USA Federal Noxious Weed●. 
Striga asiatica (L.) Kuntze: USA Federal Noxious Weed●; USA state noxious weed: CA●, NC°, SC°. 
Striga densiflora (Benth.) Benth.: USA Federal Noxious Weed●. 
Striga forbesii Benth.: USA Federal Noxious Weed●. 
Striga gesnerioides (Willd.) Vatke: USA Federal Noxious Weed●. 
Striga hermonthica (Delile) Benth.: USA Federal Noxious Weed●. 
Striga masuria (Buch.-Ham. ex Benth.) Benth.: USA Federal Noxious Weed●. 
Striga spp.: USA Federal Noxious Weed●; USA state noxious weed: AR●, AZ●, HI●°. 

Symbols: ªaquatic weed; ●terrestrial weed; °weed in seed. 
Last updated February 2006.

USA states and territories with listed noxious weeds: Alabama (AL), Alaska (AK), Arizona (AZ), Arkansas (AR), California (CA), Colorado (CO), Florida (FL), Hawaii (HI), Idaho (ID), Montana (MT), Nevada (NV), New Mexico (NM), North Carolina (NC), North Dakota (ND), Oklahoma (OK), Oregon (OR), Puerto Rico (PR), South Carolina (SC), South Dakota (SD), Washington (WA), and Wyoming (WY).

USA state and territory noxious weeds: 
Alectra spp.: USA Federal Noxious Weed●; USA state noxious weed: FL●. 
Bacopa monnieri (L.) Pennell: USA state noxious weed: CAª. 
Bacopa spp.: USA state noxious weed: PRª. 
Chaenorhinum minus (L.) Lange: USA state noxious weed: WA●. 
Limnophila indica (L.) Druce: USA state noxious weed: CAª. 
Limnophila sessiliflora (Vahl) Blume: USA Federal Noxious Weedª●; USA state noxious weed: ALª, FLª, NCª, SCª. 
Limnophila spp.: USA state noxious weed: OKª. 
Linaria dalmatica (L.) Mill.: USA state noxious weed: CO●°, ID°, MT●°, NV●°, OR●°, SD●, WA●°, WY●°. 
Linaria genistifolia (L.) Mill.: USA state noxious weed: CO●. 
Linaria genistifolia subsp. dalmatica (L.) Maire & Petitm. (=Linaria dalmatica (L.) Mill.): USA state noxious weed: CA●, ID●°, ND●, NM●. 
Linaria vulgaris Mill.: USA state noxious weed: AK°, CO●°, ID●°, MT°, NM●, NV●, OR●, SD●, WA●°, WY●°. 
Micranthemum umbrosum (J. F. Gmel.) S. F. Blake: USA state noxious weed: PRª. 
Striga asiatica (L.) Kuntze: USA Federal Noxious Weed●; USA state noxious weed: CA●, NC°, SC°. 
Striga spp.: USA Federal Noxious Weed●; USA state noxious weed: AR●, AZ●, HI●°. 
Verbascum blattaria L.: USA state noxious weed: CO●. 
Verbascum thapsus L.: USA state noxious weed: CO●, HI●°, SD●. 

Symbols: ªaquatic 1weed; ●terrestrial weed; °weed in seed. 

Last updated February 2006.
 

Distribution

General distribution: Cosmopolitan. New World and Old World.

Detailed distribution: North America to Oceania.

Notes

Now including Globulariaceae. Cronquist: Fruit rarely (Lagotis) a schizocarp - but which kind? Read Sutton (1988).

References

Literature specific to this family: Sutton, D.A. 1988. A revision of the tribe Antirrhineae. Oxford University Press, London; Musselman, L.J. & W.F. Mann, Jr. 1976. A survey of surface characteristics of seeds of Scrophulariaceae and Orobanchaceae using scanning electron microscopy. Phytomorphology 26:370–379; Elisens, W.J. & A.S. Thomb. 1983. Seed morphology in New World Antirrhinae (Scrophulariaceae): Systematic and phylogenetic implications. Pl. Syst. Evol. 142:23–47; Canne, J.M. 1980. Seed surface features in Aureolaria, Brachystigma, Tomanthera, and certain South American Agalinis (Scrophulariaceae). Syst. Bot. 5:241–252; Falcão Ichaso, C.L. 1978. Tipos de sementes encontradas nas Scrophulariaceae. Rodriguésia 30:335–344.

General references: Corner, E.J.H. 1976. The seeds of Dicots, esp. vol. 2. Cambridge University Press, New York, Cronquist, A. 1981. An integrated system of classification of flowering plants, 1,262 p. Columbia University Press, New York, Gaertner, J. 1788–1805. De fructibus et seminibus plantarum. The Author, Stuttgart, Goldberg, A. 1986 (dicots) & 1989 (monocots). Classification, evolution, and phylogeny of the familes of Dicotyledons. Smithsonian Contr. Bot. 58 for dicots (314 pp.) & 71 for monocots (74 pp.). [Goldberg's illustrations are reproduced from older publications and these should be consulted], Gunn, C.R. & C.A. Ritchie. 1988. Identification of disseminules listed in the Federal Noxious Weed Act. Techn. Bull. U.S.D.A. 1719:1–313, Gunn, C.R., J.H. Wiersema, C.A. Ritchie, & J.H. Kirkbride, Jr. 1992 & amendments. Families and genera of Spermatophytes recognized by the Agricultural Research Service. Techn. Bull. U.S.D.A. 1796:1–500, LeMaout, E. & J. Decaisne. 1876. A general system of botany, 1,065 p. Longmans, Green, & Co., London, Mabberley, D.J. 1987. The plant-book, 706 p. Cambridge University Press, Cambridge, Martin, A.C. 1946. The comparative internal morphology of seeds. Amer. Midl. Naturalist 36:513–660, Schopmeyer, C.S. 1974. Seeds of Woody plants in the United States. Agric. Handb. 450:1–883, Spjut, R.W. 1994. A systematic treatment of fruit types. Mem. New York Bot. Gard. 70:1–182, and Wood, C.E., Jr. 1974. A student's atlas of flowering plants: Some dicotyledons of eastern North America, 120 pp. Harper & Row, New York.

 Fruit:  Agalinis auriculata ; Photo by R. Gibbons, USDA APHIS PPQ, Kirkbride et al. (2006)
Fruit: Agalinis auriculata; Photo by R. Gibbons, USDA APHIS PPQ, Kirkbride et al. (2006)
 Seeds:  Agalinis auriculata ; Photo by R. Gibbons, USDA APHIS PPQ, Kirkbride et al. (2006)
Seeds: Agalinis auriculata; Photo by R. Gibbons, USDA APHIS PPQ, Kirkbride et al. (2006)
 Seeds:  Agalinis auriculata ; Photo by R. Gibbons, USDA APHIS PPQ, Kirkbride et al. (2006)
Seeds: Agalinis auriculata; Photo by R. Gibbons, USDA APHIS PPQ, Kirkbride et al. (2006)
 Fruit:  Alectra stricta ; Photo by R. Gibbons, USDA APHIS PPQ, Kirkbride et al. (2006)
Fruit: Alectra stricta; Photo by R. Gibbons, USDA APHIS PPQ, Kirkbride et al. (2006)
 Seeds:  Alectra stricta ; Photo by R. Gibbons, USDA APHIS PPQ, Kirkbride et al. (2006)
Seeds: Alectra stricta; Photo by R. Gibbons, USDA APHIS PPQ, Kirkbride et al. (2006)
 Fruit:  Buddleja davidii ; Photo by R. Gibbons, USDA APHIS PPQ, Kirkbride et al. (2006)
Fruit: Buddleja davidii; Photo by R. Gibbons, USDA APHIS PPQ, Kirkbride et al. (2006)
 Seeds:  Buddleja davidii ; Photo by R. Gibbons, USDA APHIS PPQ, Kirkbride et al. (2006)
Seeds: Buddleja davidii; Photo by R. Gibbons, USDA APHIS PPQ, Kirkbride et al. (2006)
 Fruit:  Castilleja pilosa ; Photo by R. Gibbons, USDA APHIS PPQ, Kirkbride et al. (2006)
Fruit: Castilleja pilosa; Photo by R. Gibbons, USDA APHIS PPQ, Kirkbride et al. (2006)
 Seeds:  Castilleja pilosa ; Photo by R. Gibbons, USDA APHIS PPQ, Kirkbride et al. (2006)
Seeds: Castilleja pilosa; Photo by R. Gibbons, USDA APHIS PPQ, Kirkbride et al. (2006)
 Seeds:  Castilleja pilosa ; Photo by R. Gibbons, USDA APHIS PPQ, Kirkbride et al. (2006)
Seeds: Castilleja pilosa; Photo by R. Gibbons, USDA APHIS PPQ, Kirkbride et al. (2006)
 Seeds:  Euphrasia subexserta ; Photo by R. Gibbons, USDA APHIS PPQ, Kirkbride et al. (2006)
Seeds: Euphrasia subexserta; Photo by R. Gibbons, USDA APHIS PPQ, Kirkbride et al. (2006)
 Seeds:  Euphrasia subexserta ; Photo by R. Gibbons, USDA APHIS PPQ, Kirkbride et al. (2006)
Seeds: Euphrasia subexserta; Photo by R. Gibbons, USDA APHIS PPQ, Kirkbride et al. (2006)
 Seeds:  Euphrasia subexserta ; Photo by R. Gibbons, USDA APHIS PPQ, Kirkbride et al. (2006)
Seeds: Euphrasia subexserta; Photo by R. Gibbons, USDA APHIS PPQ, Kirkbride et al. (2006)
 Fruit:  Kickxia elatine ; Photo by R. Gibbons, USDA APHIS PPQ, Kirkbride et al. (2006)
Fruit: Kickxia elatine; Photo by R. Gibbons, USDA APHIS PPQ, Kirkbride et al. (2006)
 Seeds:  Kickxia elatine ; Photo by R. Gibbons, USDA APHIS PPQ, Kirkbride et al. (2006)
Seeds: Kickxia elatine; Photo by R. Gibbons, USDA APHIS PPQ, Kirkbride et al. (2006)
 Fruit:  Linaria vulgaris ; Photo by R. Gibbons, USDA APHIS PPQ, Kirkbride et al. (2006)
Fruit: Linaria vulgaris; Photo by R. Gibbons, USDA APHIS PPQ, Kirkbride et al. (2006)
 Seeds:  Linaria vulgaris ; Photo by R. Gibbons, USDA APHIS PPQ, Kirkbride et al. (2006)
Seeds: Linaria vulgaris; Photo by R. Gibbons, USDA APHIS PPQ, Kirkbride et al. (2006)
 Fruit:  Penstemon palmeri ; Photo by R. Gibbons, USDA APHIS PPQ, Kirkbride et al. (2006)
Fruit: Penstemon palmeri; Photo by R. Gibbons, USDA APHIS PPQ, Kirkbride et al. (2006)
 Seeds:  Penstemon palmeri ; Photo by R. Gibbons, USDA APHIS PPQ, Kirkbride et al. (2006)
Seeds: Penstemon palmeri; Photo by R. Gibbons, USDA APHIS PPQ, Kirkbride et al. (2006)
 Seeds:  Striga  sp.; Photo by R. Gibbons, USDA APHIS PPQ, Kirkbride et al. (2006)
Seeds: Striga sp.; Photo by R. Gibbons, USDA APHIS PPQ, Kirkbride et al. (2006)
 Seeds:  Striga gesnerioides ; Photo by R. Gibbons, USDA APHIS PPQ, Kirkbride et al. (2006)
Seeds: Striga gesnerioides; Photo by R. Gibbons, USDA APHIS PPQ, Kirkbride et al. (2006)
 Fruit:  Verbascum thapsus ; Photo by R. Gibbons, USDA APHIS PPQ, Kirkbride et al. (2006)
Fruit: Verbascum thapsus; Photo by R. Gibbons, USDA APHIS PPQ, Kirkbride et al. (2006)
 Seeds:  Verbascum thapsus ; Photo by R. Gibbons, USDA APHIS PPQ, Kirkbride et al. (2006)
Seeds: Verbascum thapsus; Photo by R. Gibbons, USDA APHIS PPQ, Kirkbride et al. (2006)
 Seeds:  Veronica fruticulosa ; Photo by R. Gibbons, USDA APHIS PPQ, Kirkbride et al. (2006)
Seeds: Veronica fruticulosa; Photo by R. Gibbons, USDA APHIS PPQ, Kirkbride et al. (2006)
 Embryo:  Alectra stricta ; Illustration by K. Parker, Kirkbride et al. (2006)
Embryo: Alectra stricta; Illustration by K. Parker, Kirkbride et al. (2006)
 Embryo:  Bacopa decumbens ; Illustration by K. Parker, Kirkbride et al. (2006)
Embryo: Bacopa decumbens; Illustration by K. Parker, Kirkbride et al. (2006)
 Embryo:  Dopatrium junceum ; Illustration by K. Parker, Kirkbride et al. (2006)
Embryo: Dopatrium junceum; Illustration by K. Parker, Kirkbride et al. (2006)
 Embryo:  Limnophila heterophylla ; Illustration by K. Parker, Kirkbride et al. (2006)
Embryo: Limnophila heterophylla; Illustration by K. Parker, Kirkbride et al. (2006)
 Embryo:  Limnophila sessiliflora ; Illustration by K. Parker, Kirkbride et al. (2006)
Embryo: Limnophila sessiliflora; Illustration by K. Parker, Kirkbride et al. (2006)
 Embryo:  Linaria triphylla ; Illustration by K. Parker, Kirkbride et al. (2006)
Embryo: Linaria triphylla; Illustration by K. Parker, Kirkbride et al. (2006)
 Embryo:  Lindernia ciliata ; Illustration by K. Parker, Kirkbride et al. (2006)
Embryo: Lindernia ciliata; Illustration by K. Parker, Kirkbride et al. (2006)
 Embryo:  Lindernia crustacea ; Illustration by K. Parker, Kirkbride et al. (2006)
Embryo: Lindernia crustacea; Illustration by K. Parker, Kirkbride et al. (2006)
 Embryo:  Mazus pumilus ; Illustration by K. Parker, Kirkbride et al. (2006)
Embryo: Mazus pumilus; Illustration by K. Parker, Kirkbride et al. (2006)
 Embryo:  Mecardonia dianthera ; Illustration by K. Parker, Kirkbride et al. (2006)
Embryo: Mecardonia dianthera; Illustration by K. Parker, Kirkbride et al. (2006)
 Embryo:  Misopates orontium ; Illustration by K. Parker, Kirkbride et al. (2006)
Embryo: Misopates orontium; Illustration by K. Parker, Kirkbride et al. (2006)
 Embryo:  Striga asiatica ; Illustration by K. Parker, Kirkbride et al. (2006)
Embryo: Striga asiatica; Illustration by K. Parker, Kirkbride et al. (2006)