Caulanthus

Annuals, biennials, or perennials; (caudex woody); not scapose; (usually glaucous), glabrous or pubescent, trichomes usually simple, rarely mixed with fewer, stalked, 2-rayed ones. Stems usually erect or ascending, rarely subdecumbent, unbranched or branched distally, (sometimes inflated). Leaves basal and cauline; petiolate, subsessile, or sessile; basal rosulate or not, petiolate, blade margins entire, dentate, sinuate, lobed, or pinnatifid; cauline petiolate, subsessile, or sessile, blade (base sometimes sagittate or amplexicaul, rarely subauriculate), margins entire, dentate, or lobed. Racemes (corymbose, sometimes with a terminal cluster of sterile flowers from grouping of dark purple flower buds, rarely proximalmost flowers bracteate), elongated in fruit. Fruiting pedicels erect, ascending to divaricate, or reflexed, slender or stout. Flowers: sepals usually erect or ascending, rarely spreading, oblong or ovate to lanceolate, (usually forming urceolate calyx, sometimes keeled), lateral pair saccate or not basally; petals yellow, brown, purple, lavender, white, yellowish green, or, rarely, pinkish, linear to oblanceolate, (often channeled and crisped, rarely neither, equal or abaxial (lower) pair slightly shorter than adaxial (upper) pair, longer than sepals), claw often well-differentiated from blade (sometimes longer and wider than blade, entire); stamens tetradynamous or 3 unequal pairs (rarely subequal); filaments not dilated basally, (usually distinct, sometimes adaxial or both median pairs connate); anthers oblong, (equal or adaxial pair smaller, apex obtuse or apiculate); nectar glands usually confluent, subtending bases of stamens. Fruits sessile, linear, usually smooth, rarely torulose, usually terete, latiseptate, or angustiseptate, rarely 4-angled; valves each with distinct or obscure midvein, glabrous or sparsely pubescent; replum rounded; septum complete, (veinless); ovules 14–60[–210] per ovary; style obsolete or distinct; stigma capitate or not, entire, subentire, or 2-lobed (lobes opposite valves or replum). Seeds uniseriate, usually flattened (plump in C. californicus), usually not winged, rarely margined, oblong, ovoid, or, rarely, subglobose; seed coat not mucilaginous when wetted; cotyledons incumbent or obliquely so, (entire or, rarely, 3-fid). x = 14.

Distribution

w United States, nw Mexico.

Discussion

Species 17 (17 in the flora).

The limits of Caulanthus have not changed for about 60 years following the excellent account by E. B. Payson (1923). R. C. Rollins (1993), however, resurrected Stanfordia mainly based on angustiseptate instead of terete fruits and on the presence of 3-fid cotyledons. The latter character is also present in C. coulteri, a species that Rollins maintained in Caulanthus. As emphasized by Payson and by I. A. Al-Shehbaz (1973), Stanfordia does not merit recognition, a position taken also by O. Appel and Al-Shehbaz (2003). R. E. Buck (1993, 1995) excluded three species of Caulanthus (C. anceps, C. flavescens, C. lasiophyllus) and placed them in the long-abandoned Guillenia. His primary reasoning was that they do not have the urceolate calyx or the crisped and channeled petals. However, C. flavescens has crisped and channeled petals, and sometimes has somewhat urceolate calyces, a feature not found in all species of Caulanthus. Extensive discussion on the maintenance of these three species in Caulanthus was given by Al-Shehbaz and needs not be repeated here. In my opinion, the recognition of Guillenia as distinct from Caulanthus is based on over-emphasis of a few petal characters and, without thorough molecular studies on the entire streptanthoid genera, it is more practical to maintain a more inclusive Caulanthus.

Selected References

Buck, R. E. 1995. Systematics of Caulanthus (Brassicaceae). Ph.D. dissertation. University California, Berkeley.

Lower Taxa

Caulanthus amplexicaulis, Caulanthus anceps, Caulanthus barnebyi, Caulanthus californicus, Caulanthus cooperi, Caulanthus coulteri, Caulanthus crassicaulis, Caulanthus flavescens, Caulanthus glaucus, Caulanthus hallii, Caulanthus heterophyllus, Caulanthus inflatus, Caulanthus lasiophyllus, Caulanthus lemmonii, Caulanthus major, Caulanthus pilosus, Caulanthus simulans

Key

1	Median and/or distal cauline leaves sessile, blade bases sagittate or amplexicaul	> 2
1	Median and/or distal cauline leaves usually petiolate or subsessile (sessile in C. hallii), blade bases not auriculate or sagittate	> 9
2	Stems inflated (to 4 cm diam. at widest point).	Caulanthus inflatus
2	Stems not inflated	> 3
3	Fruits angustiseptate; seeds subglobose, plump, cotyledons 3-fid.	Caulanthus californicus
3	Fruits terete, latiseptate, or 4-angled; seeds ovoid to oblong, flattened, cotyledons usually entire (3-fid in C. coulteri)	> 4
4	Racemes with a terminal cluster of sterile flowers; filaments in 3 unequal pairs, median pairs often connate	> 5
4	Racemes without a terminal cluster of sterile flowers; filaments tetradynamous (except in C. amplexicaulis), median pairs distinct	> 6
5	Fruits usually reflexed, rarely divaricate; basal leaf blade surfaces with simple and forked trichomes; cotyledons 3-fid; stigma lobes 0.5-1.5 mm.	Caulanthus coulteri
5	Fruits erect or ascending; basal leaf blade surfaces with simple trichomes; cotyledons entire; stigma lobes 1-4 mm.	Caulanthus lemmonii
6	Plants glabrous throughout; filaments in 3 unequal pairs; fruits divaricate to ascending.	Caulanthus amplexicaulis
6	Plants hispid or puberulent at least basally; filaments tetradynamous; fruits usually reflexed, rarely divaricate	> 7
7	Plants puberulent, trichomes appressed, 2-rayed; stems erect to ascending (often flexuous, weak).	Caulanthus cooperi
7	Plants hispid, trichomes spreading, simple; stems erect	> 8
8	Fruits latiseptate or 4-angled; cauline leaf blades linear-lanceolate.	Caulanthus heterophyllus
8	Fruits terete; cauline leaf blades ovate to oblong.	Caulanthus simulans
9	Perennials, with woody caudices; plants glabrous or sparsely pubescent; anthers 3-6.5 mm	> 10
9	Annuals, biennials, or, rarely, perennials (short-lived), without caudices; plants moderately to densely hispid, hirsute, or pilose (at least basally); anthers 0.4-3 mm	> 13
10	Stems fusiform, strongly inflated (to 3 cm diam. at widest point).	Caulanthus crassicaulis
10	Stems cylindrical, not or slightly inflated	> 11
11	Basal leaves rosulate; fruiting pedicels 1-6 mm; stigma lobes opposite valves.	Caulanthus major
11	Basal leaves not rosulate; fruiting pedicels 5-35 mm; stigma lobes opposite replum	> 12
12	Petals prominently dark-veined; stigma lobes to 0.4 mm in fruit.	Caulanthus barnebyi
12	Petals not or faintly veined; stigma lobes 0.9-1.5 mm in fruit.	Caulanthus glaucus
13	Sepals widely spreading; filaments subequal.	Caulanthus anceps
13	Sepals usually erect, rarely ascending to spreading; filaments tetradynamous or in 3 unequal pairs	> 14
14	Petals 2.5-5(-6.5) mm, not crisped or channeled, claw undifferentiated from blade; calyces not urceolate.	Caulanthus lasiophyllus
14	Petals 6-13 mm, crisped and channeled (except C. hallii), claw well-differentiated from blade; calyces urceolate (except C. flavescens)	> 15
15	Petals purple; ovules 152-198 per ovary.	Caulanthus pilosus
15	Petals usually creamy white, rarely pink; ovules 44-96 per ovary	> 16
16	Filaments tetradynamous; fruits 3.8-7.7 cm; ovules 44-58 per ovary.	Caulanthus flavescens
16	Filaments in 3 unequal pairs; fruits 6.5-12.5 cm; ovules 78-96 per ovary.	Caulanthus hallii