SCROPHULARIACEAE (Figwort Family)

Plants biennial or perennial herbs or shrubs (annuals elsewhere). Leaves basal and/or opposite or alternate along the stems, sessile to long-petiolate. Stipules absent or inconspicuous, then herbaceous, attached along a line or slender ridge, the adjacent stipules of the leaves at each node fused into a single triangular to depressed-ovate structure. Leaf blades unlobed or less commonly pinnately lobed, variously shaped, the margins usually scalloped or toothed. Inflorescences racemes (sometimes spikelike) or panicles of usually numerous flowers, at least the lower nodes subtended by bracts, inconspicuous bractlets sometimes also present below each flower. Flowers perfect, hypogynous. Calyces actinomorphic or nearly so (zygomorphic elsewhere), 4- or 5-lobed, persistent at fruiting. Corollas actinomorphic or zygomorphic and sometimes bilabiate, variously colored, 4- or 5-lobed, the tube variously short to long. Stamens 4 or 5, alternating with the corolla lobes, the filaments attached in the corolla tube (sometimes near its base), sometimes unequal, the anthers exserted or not, attached near their midpoints or sometimes fused to the filament tips for 1/2 or more of the length, often appearing U-shaped or 1-locular, yellow or orange. Staminodes absent or (sometimes in flowers with 4 fertile stamens) present, then well differentiated from the fertile stamens. Pistil 1 per flower, of 2 fused carpels. Ovary 2-locular (sometimes appearing 4-locular elsewhere), with numerous ovules, the placentation axile. Style 1, often persistent at fruiting (but sometimes shriveled or shed toward the tip), the stigma 1, variously shaped, entire or 2-lobed. Fruits capsules (drupes or schizocarps elsewhere), dehiscent longitudinally from the tip. Seeds numerous, minute. About 52 genera, about 1,680 species, worldwide.

Perhaps no other family in the traditional Englerian classification has undergone as profound a reevaluation as the Scrophulariaceae. In the traditional broad sense, the family once included nearly 200 genera and about 4,000 total species. However, a number of botanists had already noted the extreme morphological diversity within the group and the difficulties involved in using morphological characters to separate the Scrophulariaceae from some related families, such as the Acanthaceae, Bignoniaceae, Globulariaceae, and Orobanchaceae (Cronquist, 1981). The application of molecular data from selected DNA marker sequences has had a profound effect on the classification of families and orders across the angiosperms and has greatly altered the classification and circumscription of the groups associated with the Scrophulariaceae, Lamiaceae, and related families. As presently circumscribed, a very large order Lamiales includes the families formerly included in that order, as well as those previously segregated into Bignoniales and Scrophulariales by some botanists (see Judd et al. [2008] for a brief review). As researchers began to sample a larger suite of species within the various groups contained in the Lamiales, it soon became apparent that the Scrophulariaceae, as traditionally treated, was an unnatural group (as reviewed by Tank et al. [2006]). This has resulted in the break-up of the Scrophulariaceae into seven to eleven families, but the number of families and the generic limits of each have not yet become fully stabilized (Angiosperm Phylogeny Group, 1998, 2003, 2009). Additionally, a few groups formerly classified as distinct families have been included within some of these recently recircumscribed families. The introduced genus Buddleja was not known to be a member of the state’s flora when Steyermark (1963) completed his volume. Many botanists have treated it in the segregate family Buddlejaceae (Leeuwenberg and Leenhouts, 1980; Cronquist 1981, 1991; G. K. Rogers, 1986; Norman, 2000), but it currently is considered by most botanists to belong in the Scrophulariaceae.

A summary of the current familial placement of the various genera included by Steyermark in the Scrophulariaceae is presented below, along with other genera that are now classified into this group of families. Members of the Acanthaceae, Bignoniaceae, Martyniaceae, and Oleaceae, families in the Lamiales whose circumscriptions for the genera present in Missouri have not changed since Steyermark’s (1963) time, are not included in the list.

Linderniaceae: Lindernia (Albach et al., 2005; Oxelman et al., 2005; Rahmanzadeh et al., 2005; Tank et al., 2006)

Orobanchaceae: Agalinis (Gerardia), Aureolaria (Gerardia), Buchnera, Castilleja, Dasistoma, Epifagus (formerly Orobanchaceae), Orobanche (formerly Orobanchaceae), Pedicularis

Paulowniaceae: Paulownia (Olmstead and Reeves, 1995)

Phrymaceae: Mazus, Mimulus, Phryma (formerly Phrymaceae) (Oxelman et al., 1999; Beardsley and Olmstead, 2002; Tank et al., 2006; Albach et al., 2009)

Plantaginaceae: Callitriche (formerly Callitrichaceae), Plantago (formerly Plantaginaceae), Antirrhinum, Bacopa, Chaenorhinum, Chelone, Collinsia, Cymbalaria, Gratiola, Kickxia, Leucospora (formerly Conobea), Linaria, Mecardonia, Nuttallanthus (formerly Linaria), Penstemon, Veronica, Veronicastrum (Albach and Chase, 2004; Albach et al., 2005; Oxelman et al., 2005)

Scrophulariaceae: Buddleja (see note above), Scrophularia, Verbascum (Olmstead et al., 2001; Oxelman et al., 2005)

Tetrachondraceae: Polypremum (formerly Loganiaceae) (Oxelman et al., 2005)

In its diminished current form, the family Scrophulariaceae has relatively limited economic importance. Members of several genera are cultivated as ornamentals, including Buddleja and Verbascum. Conversely, in some parts of the United States, species in these genera are considered troublesome weeds. Many species in the family are considered at least mildly toxic to livestock and a few genera contain plants that have been used medicinally.

In addition to the taxa treated below, Steyermark (1963) also included a report of Limosella aquatica L. (common mudwort; Pl. 559 a, b) from Missouri. He noted that the inclusion of this species was based on a single historical voucher specimen collected by B. F. Bush in Jackson County, but did not cite a collection number or herbarium where the specimen was accessioned. Palmer and Steyermark (1935) had stated that the plant occurred on sand bars of the Missouri River (as L. subulata E. Ives), but gave no other clues to the location of a voucher. Pennell (1935), in his study of the eastern temperate North American Scrophulariaceae, did not cite Missouri in the range of any species of Limosella, and Missouri has not been cited as part of the species range in other publications. During the present research, despite diligent searches, no specimens were discovered to support the inclusion of Limosella in the Missouri flora, and it is thus excluded for the present. Limosella aquatica is widespread but of sporadic occurrence in Europe and Asia, and in the New World from Canada south to South America. In the United States it is known from most western states eastward to Minnesota and Nebraska. It is a submerged or emergent aquatic that likely is spread by mud on the feet and feathers of migratory waterfowl. The species is inconspicuous; plants are rarely 15 cm tall, although the stems are stoloniferous and creeping. The leaves occur in dense tufts at the rooted stem nodes and consist of a long, slender petiole terminating in a short, lanceolate to elliptic blade. The inconspicuous flowers are solitary at the tip of a stalk that is shorter than the leaves. Corollas are 3–4 mm long, more or less actinomorphic, with a slender tube and 5 short lobes, white or pale pinkish- to purplish-tinged. Botanists who do field work in wetlands that are stopover points for ducks and other migratory waterfowl should be alert for new records of this diminutive member of the Scrophulariaceae.