**Plantago lanceolata L. (Plantaginaceae)**

**English Plantain, Ribwort Plantain**

**and Plantago major L., Common Plantain**

**Description**. Scapose, herbaceous perennials, from fibrous-rooted,
subterranean caudices, sometimes densely woolly at the apex (base of
leaves); scapes 1.5-6 dm tall, puberulent to strigose. Leaves in a basal
rosette, 5-40 cm long, 0.5-4 cm wide, petiolate, blades narrowly
elliptic to narrowly lanceolate, conspicuously 3-several veined,
glabrous to appressed villous, margins entire to denticulate, apices
acute, the bases tapered to the petiole. Inflorescences 0.5-8 cm long, a
cylindrical, terminal spike. Flowers radial, sessile, each subtended by
an ovate bract, bracts 2.5-4 mm long, ovate; calyx 2-3 mm long, lobes
four, ovate, outer 2 fused, inner ones distinct; corolla lobes 2-2.5 mm
long, spreading; stamens 4, exserted; ovary superior, locule 1, style 1,
stigma slightly 2-lobed. Fruit a capsule, 2-4 mm long, subglobose,
circumscissile; seeds 2, ca. 2 mm long, shiny, black. In California,
flowering from March to June (Chater and Cartier 1976, Cronquist 1984,
Dempster 1993, Gleason and Cronquist 1991, Holmgren 1984, McGregor and
Brooks 1986, Webb et al. 1988, Welsh et al. 1987).

The related *P. major* (common plantain) differs primarily by having
ovate to widely elliptic leaf blades with rounded to cordate bases and 6
to 30 seeds per fruit. It also lacks a distinct taproot (Chater and
Cartier 1976, Gleason and Cronquist 1991, Holmgren 1984, McGregor and
Brooks 1986).

**Note**: Both *P. lanceolata* and *P. major* show considerable ecotypic
variation and occupy a diversity of habitats throughout Europe (Kuiper
1992, Lotz 1990, Lotz et al. 1990, Sagar and Harper 1960, 1964, van Dijk
1989, 1992, van Dijk and Kuiper 1992, Wolff 1991).

**Geographic distribution**. Natives of Eurasia, both English and common
plantain have become naturalized throughout North America, South
America, Australia, New Zealand, Hawaii, southeast Asia, and eastern
Africa. (Arnold and de Wet 1993, Chapman 1991, Chua et al. 1994,
Cronquist 1984, Pereira and Romaniuc Neto 1993, Sagar and Harper 1964,
Wagner 1990, Webb et al. 1988). *Plantago lanceolata* is also known from
Japan and Taiwan (Chen et al. 1996, Ohwi 1965).

Both species were reported as widely distributed and naturalized by the
end of the 19th century (Brewer et al. 1876, Robbins 1940). Both species
have been reported from Santa Catalina and Santa Cruz islands; *P.
major* is also known from San Nicolas Island (Junak et al. 1997). Both
species are widely distributed throughout California (Dempster 1993,
Munz 1959).

**Reproductive and vegetative biology**: *Plantago lanceolata* is
gynodioecious, self-incompatible, and both insect- and wind-pollinated
(Allard 1965, Bos et al. 1986, Proctor et al. 1996, Ross 1973, van Damme
and van Delden 1982, 1984). Hoverflies (Syrphidae) are apparently the
most effective pollinators (Stelleman and Meeuse 1976, Stelleman 1978,
1981). *Plantago* *major*, in contrast, has bisexual flowers, is
self-compatible, and largely self-pollinating (Proctor et al. 1996,
Wolff et al. 1994).

*Plantago* species have mucilaginous seeds (Samuelsen et al. 1996),
which, when wetted, contribute to low dispersability and enhanced
establishment of the seedling phase (Harper 1977, Tonsor 1985). Seeds of
both species have a dormancy requirement that generally is broken by
either a brief light requirement or by a dry period (Pons 1992, Sagar
and Harper 1960, 1964). Seeds of *Plantago lanceolata* are less
sensitive to light than *P. major*, the former often germinating at
greater soil depths than the latter (Pons and van der Toorn. 1988, van
der Toorn and Pons 1988). The highest germination rates under field
conditions are generally observed in open sites and in the fall or early
winter in north temperate climates (Sagar and Harper 1964), although
Popay et al. (1995) reported peak emergence during the New Zealand
spring months. McRill and Sagar (1973) also reported increased
germinability of *P. lanceolata* seeds after ingestion by earthworms.
Palmblad (1968) reported relatively high survivorship among seed cohorts
of both species, with mortality experienced at later stages of plant
development and strongly related to plant density.

Under competitive conditions, *Plantago lanceolata* plants live from 1
to 3 years, but survivorship can exceed 4 years in undisturbed sites
(Harper 1977). *Plantago lanceolata* also also reproduces by lateral
crown shoots, whereas reproduction by *P. major* is restricted entirely
to seeds (Sagar and Harper 1964).

*Plantago lanceolata* produces iridoid glycosides, which are toxic or
unpalatable to most insects, except arctiid and nymphalid butterflies
and weevils (Bowers 1984, Bowers and Stamp 1992, Bowers et al. 1992b,
Pereyra and Bowers 1988, Dethier 1993).

**Ecological Distribution**. Both species are reported from waste
places, roadsides, gardens, lawns, and fallow fields (McGregor and
Brooks 1986, Munz 1959, Sagar and Harper 1964, Welsh et al. 1987).
*Plantago* *lanceolata* is one of the more common herbaceous perennials
reported from northern coastal prarie in California (Heady et al. 1988).

**Weed status.** *Plantago* lanceolata and *P*. *major* are considered
noxious weeds in agricultural or horticultural practice at a global
level (Holm et al. 1977), However, it is considered as a potentially
noxious weed by the State Dept. of Food and Agriculture (Anonymous
1996). Both are listed for the United States in Lorenzi and Jeffery
(1987).

**Microbial pathogens**. Fungal diseases reported for *Plantago major*
include *Erysiphe cichoracearum* and *Peronospora altal* (Drozdovskaia
and Nikolaeva. 1984). *Plantago lanceolata* has been reported as a host
to *Phomopsis subordinaria* (de Nooij 1988, de Nooij and van der Aa
1987, de Nooij and van Damme 1988a, 1988b, de Nooij et al. 1986),
*Diaporthe adunca* (Linders and van der Aa 1995), and *Sclerotina*
(Saito 1997).

**Insect pathogens**. *Plantago lanceolata* and *P. major* have been
reported as hosts to arctiid and nymphalid butterfly larvae (Dethier
1993, Bowers et al. 1992a, Ehrlich et al. 1975, Pereyra and Bowers 1988,
Stamp and Bowers 1992). Weevils (*Trichosirocalus troglodytes*) are
important herbivores of *Plantago lanceolata* and also may serve to
transmit the fungus *Diaporthe* (Linders et al. 1995, 1996)

**Herbicide control**. A number of herbicides, including dicamba,
clopyrid, chlorflurenol, trichlopyr, and 2,4-D, have been found
effective on *P. major* in turf grass (Neal 1990). Del Amor (1981) used
drip irrigation as a method for applying triazine herbicides to control
*Plantago* in citrus orchards.

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