BTK Santa Barbara County

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BTK In San Marcos

Brodiaea jolonensis?

Brodiaea elegans

Key To Brodiaeas

Index Of Brodiaea Pages

Coastal BTK In Santa Barbara County

Note: The Brodiaea, that I referred to as "Coastal BTK," is listed as a possible
undescribed taxon under B. jolonensis in the revised Jepson Manual II (2011)

Note: Although listed in many references for mainland & insular southern California, B. jolonensis appears to be endemic to the Coast Ranges of Monterey and possibly San Luis Obispo Counties.

Brodiaea terrestris ssp. kernensis (BTK), including coastal & montane forms, is a large and diverse complex that extends from Santa Barbara and Kern Counties to the Mexican border. PCA data thus far indicates that BTK is one variable species that does not warrant segregation. BTK intergrades into variable populations on the Santa Rosa Plateau of Riverside County and at Cuyamaca Lake and Camp Pendleton in San Diego County. Fertile clonal variants have hooded staminodes, staminodes that are flattened and strap-shaped or inrolled along upper margins, and narrow staminodes that are tapered toward the apex. In addition, the staminodes may be erect, leaning slightly outward or leaning inward. Whether the anther connectives of BTK have a dentate appendage or V-shaped notch is also quite variable, particularly in Kern County populations. This trait is of little value in separating populations of BTK, but is fairly consistent in populations of B. terrestris ssp. terrestris (BTT) in San Luis Obispo and Monterey Counties. In general, populations of BTK in southern California lack a well-defined dentate connective; however, this trait occurs in some Kern County populations. These traits are apparently not under selection pressure and hence exhibit a lot of variability. In addition to clonal variants, there are occasional sterile hybrids between BTK and other sympatric populations, such as B. filifolia and B. orcuttii.

Staminode Variation Of BTK On Santa Rosa Plateau
Staminode Variation Of BTK At Camp Pendleton
Staminode Variation Of BTK At Cuyamaca Lake
Dentate Anther Connective In BTK Populations

Overlook Trail Above Gaviota Pass

Trait	Kern County BTK	S. California Coastal BTK	S. California Montane BTK	Central Calif. BTT	Central Calif. BJ
Fls gen 22-36 mm	X	X	X	--	--
Fls gen 15-22 mm	--	X	--	X	X
Staminode inrolled	X	--	X	X	--
Staminode hooded	--	X	--	--	X
Staminode slightly hooded & inrolled	--	X	--	--	X
Ovary green	X	X	X	X	--
Ovary purple	--	--	--	--	X
Top of connective with dentate lobe	--	--	--	X	--
Connective with V-shaped notch	X	X	X	--	X
Dentate lobe in center of notch	X	X	--	--	--
Connate ridge 0.5-2 mm	--	X	--	X	X
Connate ridge < 0.5 mm or 0	X	--	X	--	--
Inner perianth 9 vasc strands	X	X	X	--	--
Inner perianth 3-4 strands	--	--	--	X	X
Sporophyte chromosome #	48	36?	?	12, 36	12 (Jolon)

1. Measured from attachment point at pedicel to tip of perianth segment.
2. Flower length of Coastal BTK has range of 17-30 mm.
3. Top of connective tissue extends into a pointed tooth resembling an inverted-V.
4. Top of connective has an indentation or notch resembling a V or a U.
5. Notch in top of connective contains a pointed toothlike lobe resembling the center of a W.
6. This is a narrow ridge of inner perianth tissue between the bases of adjacent staminodes and filaments. When viewed from above, it appears to fuse the bases of staminodes and filaments into a connate ring of tissue. In some species this ridge may be at least one mm in thickness. In others, it is less than 0.5 mm and barely discernable or lacking. It is barely discernable in some populations of Kern County BTK and southern California Montane BTK, but is conspicuous in Coastal BTK of southern California and Brodiara jolonensis of central California.

See Explanation for the Connate Ridge In Brodiaea Species
Connate Ridge of Brodiaea jolonensis in Monterey County
Connate Ridge of Coastal BTK in Santa Barbara County

7. Brodiaea jolonensis appears to be endemic to the Coast Ranges of central California (primarily Monterey County). Reports of BJ in southern California (Santa Barbara to the Mexican border) appear to be Coastal BTK. Reports of BJ on Santa Catalina and San Miguel Island are also Coastal BTK. The taxonomic status of Coastal BTK is enigmatic because it tends to intergrade with Montane BTK on the Santa Rosa Plateau of Riverside County.
8. There are other characteristics not listed in the above table. Some of these are quite variable and might be correlated with annual rainfall or other factors. In general, BJ has consistently the longest scapes of all brodiaea populations observed (up to 29 cm). BTT has the consistently shortest scapes, often only 2-6 cm long. Inland BTT in the La Panza Range resemble a diminutive version of Kern County BTK, except the staminodes are more strongly bent inward. Ovaries of BJ in Monterey County are consistently purple, a trait that is shared by no other species. Some Coastal and Montane BTK in southern California have green ovaries with faint streaks of purple, but these could hardly be called purple. Chromosome numbers need further study. Niehaus' numbers of 36 for BJ in southern California obviously refer to Coastal BTK.

Coastal BTK occur on distant grassy ridge in center of image.

Ridge high above Gaviota Pass dominated by naturalized annual grasses, including Avena, Lolium and Phalaris. The faint path follows the Overlook Trail (Fire Road). The Coastal BTK were completely obscured by the dense grasses. You literally had to search through the grasses to find them.

The Coastal BTK above Gaviota Pass have white staminodes. The staminodes are hooded at the apex. They are not inrolled along the margins as in southern California Montane BTK and Kern County BTK.

The Coastal BTK above Gaviota Pass have white staminodes.

BTK in San Marcos With White Staminodes
BTK on Santa Rosa Plateau With White Staminodes

Close-up view of V-shaped notch at the top of anther connective from the Gaviota Pass population. At the base of this notch is a slight proliferation of the connective tissue. This is more pronounced in other populations of Coastal BTK, such as Otay Mesa in San Diego County. In some populations of Coastal BTK with a more prominent lobe, the notch appears W-shaped rather than V-shaped. Unfortunately, this lobe at the base of the notch does not show up in all samples of Coastal BTK.

Close-up view of the anthers of Coastal BTK (previously labeled B. jolonensis) from the San Marcos vernal pool area showing details of the adaxial side of the anther connective (anther axis). A and B show a minute "dentate" lobe in the apical notch of the anther connective (red arrows). C has a reduced lobe and in D there is practically no visible lobe. The minute lobe at the base of the notch is formed by a proliferation (elongation) of the connective tissue. The presence of this "dentate" lobe and green ovary indicate that this species is Coastal BTK rather than B. jolonensis. Unfortunately, this "dentate" lobe does not show up in all samples of Coastal BTK. The term "dentate connective" described in the 1996 Jepson Manual is clearly evident in B. terrestris ssp. terrestris of central and northern California and in some ssp. kernensis from Kern County.

The ovary is green, typical of southern California BTK.

Coastal BTK showing connate ring (ridge) between bases of staminodes and filaments. The staminodes are hooded and slightly inrolled along their margins.

Coastal BTK showing connate ring (ridge) between bases of staminodes and filaments. The hooded staminodes are also slightly inrolled along their margins.

Reports of BJ in Santa Barbara County are undoubtedly Coastal BTK that occurs in populations the extend south to the Mexican border. It has been reported from vernal pool fields such as this one just north of UCSB. We have not been able to locate any Coastal BTK in these fields which are dominated by tall annual grasses (mostly Avena and Lolium). The vernal pool in foreground contains button celery (Eryngium aristulatum var. aristulatum).

Phacelia grandiflora in the chaparral of Gaviota Pass.

Vascular Pattern Of Coastal BTK Inner Perianth Segment

The inner perianth segments of all Brodiaea species are composed of three primary vascular bundles. The bundles are subdivided into separate vascular strands, each composed of vessels with spirally thickened secondary cell walls. According to Niehaus (1971), the bundles separate into strands, the number of which is characteristic for certain species. The bundles of Brodiaea jolonensis do not separate into multiple strands, and therefore, each bundle represents one strand. I have seen one or more bundles of BJ with two strands; and therefore, the total number of strands can be at least 4. In Coastal BTK, the bundles typically separate into two or three strands, and the total number of strands can be nine. The three primary bundles are relatively easy to determine at 100x magnification; however, the total number of strands can be difficult to see. The bottom line here is that the vascular bundles of Coastal BTK clearly separate into more vascular strands than BJ. The number may be as high as nine. This higher number number for Coastal BTK has been verified from populations throughout southern California and from Santa Catalina and San Miguel Islands. It could be explained by the fact that BTK typically has larger flowers than BJ.

Microscopic view of the inner perianth segment of a Coastal BTK flower from above Gaviota Pass, Santa Barbara County. There are three primary vascular bundles each containing three vascular strands. Each strand is composed of vessels with spirally thickened secondary cell walls. This is an exceptional view because it is usually difficult to show all three strands of each primary bundle in a single image. Under the microscope, it is often necessary to move along the midrib of the perianth segment to see where each primary bundle separates into multiple strands. In dried herbarium specimens, the strands often become broken or distorted, especially if the segment is wrinkled or creased. In Brodiaea jolonensis there is only one strand per bundle, a total of three strands. Sometimes a bundle separates into two strands, giving a total of four strands. Perianth segment cleared in 10% NaOH and stained with safranin. (Magnification 200 x).

Explanation of Vascular Strands in Brodiaeas

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