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Does Brodiaea elegans Occur in San Diego County?

Until a satisfactory name is applied to these populations, I will refer to southern California populations
previously called "Brodiaea jolonensis" as "Coastal BTK" and mountain populations previously called
"Brodiaea terrestris ssp. kernensis" as "Montane BTK."
The flower color for Brodiaea species on this page is blue-purple to violet. In the following images
I have attempted to match their true color;  however, they may appear different on your monitor.

After many years of botanizing in San Diego County, I was convinced that rumors of the occurrence of Brodiaea elegans were probably misidentified populations of Coastal BTK, previously identified as B. jolonensis; however, after seeing images from Mike Simpson of San Diego State University and revisiting his collection site, I am now convinced that B. elegans does in fact occur in San Diego County.

Brodiaea elegans ssp. elegans at Mission Trails Regional Park. The three white staminodes are clearly "standing away from stamens" and the "perianth tube is funnel-shaped." The staminodes in photo have an acute apex, but they can also be 3-toothed.

Brodiaea elegans ssp. elegans photographed by Mike Simpson at Mission Trails Regional Park. The three white staminodes (red arrow) are about the same height as stamens and are 3-toothed at the apex. Photo images courtesy of Mike Simpson, San Diego State University.


A variant brodiaea photographed at Mission Trails Regional Park in May 2005. The flower has 3-toothed or cuspidate staminodes similar to the 3-toothed staminodes of B. elegans. Unlike B. elegans, the staminodes are much longer than the stamens. The flowers appeared to be intermediate between nearby B. elegans ssp. elegans and Coastal BTK; however they had 65% pollen viability which is not typical of a true interspecific brodiaea hybrid. According to T. F. Niehaus (personal communication, 2004), hybrids are encountered where the distribution of two species overlap. The hybrids show a range of intermediate characteristics between the parents. Under controlled greenhouse conditions, Niehaus was able to hybridize every combination between all species and obtain seed. In a few crosses a relatively high percentage of seed set was obtained, although seed set was low in most interspecific crosses. According to Niehaus (1971), the majority of hybrids had sterile pollen. Although pollen viability is a good indicator of hybridization in brodiaeas, there may be other factors causing hybrid sterility. Hybrid clones can be propagated by growing cormlets.

Pollen Of The B. elegans Hybrid

Until a satisfactory name is applied to these populations I will refer to southern California populations of
B. jolonensis as "Coastal BTK" & mountain populations of B. terrestris ssp. kernensis as "Montane BTK."

Abstract: Coastal populations of Brodiaea in San Diego County with prominent hooded staminodes (with incurved apex) have been previously called B. jolonensis. Using the Jepson Manual (1996) which is based on A Biosystematic Study of the Genus Brodiaea (Amaryllidaceae) by T.F. Niehaus (1971), they key out to B. terrestris ssp. kernensis. This species has a green ovary, unlike the purple ovary of B. jolonensis. The term "dentate connective" used in the Jepson key ideally applies to B. terrestris ssp. terrestris of central and northern California. In this subspecies, the apex of the anther connective extends into a prominent dentate lobe. In southern California populations of Coastal BTK and Montane BTK, the connective apex has a U-shaped or V-shaped notch. At the base of this notch there may be a minute, dentate lobe, although it is absent in most southern California populations. With a dentate appendage at the base, the notch appears W-shaped. B. jolonensis also has a V-shaped notch without a dentate appendage. These subtle differences are difficult to observe in pressed herbarium specimens. Floral dimensions of Coastal BTK and Montane BTK are significantly larger than populations of B. jolonensis in Monterey County, and the scapes are generally shorter. In addition, vascular strand patterns of the inner perianth match those of B. terrestris kernensis.

Populations of Coastal BTK throughout San Diego County generally have staminodes that are incurved at the apex (hooded), while populations of Montane BTK generally have erect staminodes that are inrolled along the upper margins, but not hooded or slightly hooded. Both Munz (A Flora of Southern California, 1974) and Beauchamp (A Flora of San Diego County, California, 1986) recognized a difference between the coastal and mountain populations. These authors called the mountain populations B. terrestris ssp. kernensis and the coastal populations "B. jolonensis." Brodiaeas resembling both the coastal and montane forms of BTK grow together on the Santa Rosa Plateau of Riverside County. There is also staminode variation at Cuyamaca Lake in San Diego County. Perhaps this is a single variable species in southern California. This is an ongoing research project with Tom Chester to understand the Brodiaea populations in southern California. W.P. Armstrong, May 2005.

Coastal BTK at Mission Trails Regional Park. The staminodes are notched at the apex and erect. The apex is also curved inward toward the stamens (hooded). This hooded staminode is consistent with all populations of Coastal BTK in San Diego County.

Brodiaea elegans ssp. elegans at Mission Trails Regional Park.

Brodiaea elegans ssp. elegans at Mission Trails Regional Park. When photographed on 17 May 2005 the flowers in this population contained small flower beetles which probably serve as pollinators.

Minute flower beetles were common in the Mission Trails population of Brodiaea elegans when photographed in May 2005. These beetles were also observed in populations of Coastal BTK on Otay Mountain the previous year. By the way, the size relationships in photo include the "head" of an ordinary straight pin, the "eye" of a sewing needle, and a millimeter ruler.

Brodiaea elegans ssp. elegans at Mission Trails Regional Park.

More Images From Mission Trails Regional Park


Hopefully the following discussion and images may explain the differences between B. elegans and other species of Brodiaea in San Diego County. In my opinion, the Jepson Manual key to Brodiaea species is somewhat confusing until you have seen all of the species. It is based on A Biosystematic Study of the Genus Brodiaea (Amaryllidaceae) by T. F. Niehaus, University of California Publications in Botany Volume 60, University of California Press, 1971.

In order to key out Brodiaea elegans, you must assume that staminodes (staminodia) are present and the perianth lobes are ascending. For "widely spreading" perianth lobes see Brodiaea filifolia at the following link.

Thread-leaf Brodiaea (Brodiaea filifolia)

The perianth tube is not narrowed above the ovary and the staminodes are clearly narrower than the perianth segments, so B. pallida and B. stellaris can be ruled out. The staminodes are wider at base and not wavy, therefore B. appendiculata and B. californica can be ruled out.

This brings us to the couplets at line 14 in the Jepson Manual key page 1180 (3rd printing, 1996): Staminodes standing away from stamens; perianth tube funnel-shaped; OR 14' Staminodes leaning in toward stamens; perianth bell-shaped. B. elegans definitely has staminodes that are widely separated from the stamens and a perianth tube that is funnel-shaped rather than bell-shaped. We also have a brodiaea in San Diego County that fits the couplet "stamens leaning in toward stamens."

The following image is B. jolonensis from Monterey County. I am virtually positive that this species is identified correctly. In the Jepson Manual this species keys out under 14': "The staminodes leaning in toward stamens." I wouldn't use the word "leaning," but there is a slight inward curve toward the stamens. This staminode position not only fits B. jolonensis, it also fits B. terrestris and B. coronaria.

Brodiaea jolonensis in Monterey County, California. Photograph taken near marshy creek along trail to Junipero Serra Peak in early June, 2004. The ovary of this flower was purple, unlike the green ovaries of Coastal BTK and Montane BTK in San Diego County. The apex of the staminodes is slightly curved inward (hooded) as in coastal San Diego populations of B. terrestris kernensis.

The following CalPhotos images show B. elegans with widely spreading staminodes and B. terrestris with staminodes leaning inward toward stamens:

CalPhotos Image of Brodiaea elegans
CalPhotos Image of Brodiaea terrestris

All of the Coastal BTK that I have examined in coastal San Diego County have staminodes with an inward curvature at the apex (hooded) that is similar to B. jolonensis. Their relative position to the stamens is about the same as B. jolonensis. They are closer to the stamens than B. elegans. Under the description for B. terrestris kernensis, the Jepson Manual includes the statement: "Staminodes leaning inward slightly but held away from stamens." See the following images of Coastal BTK in coastal San Diego County. The apices of all staminodes are curved inwardly, a characteristic referred to as "hooded."

Coastal BTK in San Marcos.

Coastal BTK on Otay Mesa.

Coastal BTK and Pogogyne abramsii on Kearny Mesa.

Another Image Of Coastal BTK on Kearny Mesa

Coastal BTK in the Ramona vernal pool area.


Brodiaea populations in the Laguna and Cuyamaca Mountans have staminodes that are erect and inrolled along margins. They appear different from populations of Coastal BTK because the staminodes are generally not hooded. In B. coronaria the staminodes are strongly inrolled along margins but they lean inward toward the stamens. The San Diego mountain populations have staminodes that are erect and held away from the stamens. In B. elegans the staminodes are more widely separated from the stamens and they are approximately the same length as stamens. Staminodes in the mountain populations are clearly longer than the stamens. In addition, flowers of the Laguna and Cuyamaca populations do not resemble the CalPhoto images of B. coronaria or B. elegans. It is possible that these mountain populations more closely resemble the true B. terrestris ssp. kernensis of Kern and Tulare Counties; however, they do not have the dentate anther connective described by Niehaus for B. terrestris. Instead they have a prominent V-shaped or U-shaped notch in the anther connective as in Coastal BTK. Until we understand their taxonomic relationship with the true ssp. kernensis of Kern and Tulare Counties, we are calling the southern California populations Coastal BTK and Montane BTK.

Montane BTK in the Laguna Mountains.

The staminodes are erect and inrolled along margins; however, they are not curved inward at the apex (hooded) as in coastal San Diego populations of Coastal BTK.

Montane BTK at Cuyamaca Lake (17 June 2005).

The staminodes are erect and hooded as in coastal San Diego populations of Coastal BTK.

The following CalPhotos image of B. elegans shows flat staminodes that are not inrolled and are standing away from the stamens. CalPhotos apparantly does not have the inrolled B. elegans ssp. hooveri. In the above image the inrolled staminodes are in the same relative position as B. terrestris. Their shape and position more closely resembles the CalPhoto images of B terrestris than B. coronaria.

CalPhotos Image of Brodiaea elegans
CalPhotos Image of Brodiaea coronaria
CalPhotos Image of Brodiaea coronaria
CalPhotos Image of Brodiaea terrestris
CalPhotos Image of Brodiaea terrestris

Montane BTK in the Laguna Mountains.

At this time, I am at a loss to explain the differences in the Brodiaea populations from the Laguna and Cuyamaca Mountains compared with coastal San Diego populations of BTK. Although they both have the V-shaped or U-shaped anther connective, Montane BTK lacks the minute lobe at the base of the notch that is found in some populations of Coastal BTK. Mountain populations generally lack the hooded apex of staminodes typical of Coastal BTK. In addition, the floral tube appears slightly more funnel-shaped than bell-shaped. Perhaps mountain populations are more representative of the trueB. terrestris ssp. kernensis described for central California. Until we understand the taxonomy of true ssp. kernensis of Kern and Tulare Counties, we are calling the southern California populations Coastal BTK and Montane BTK.

With the exception of B. filifolia, B. orcuttii and B. elegans, the only other possible coastal brodiaeas for San Diego County are Coastal BTK and B. jolonensis. I believe B. jolonensis can be ruled out because no coastal brodiaeas have its characteristic purple ovary, and the vascular pattern of inner perianth segments is not characteristic of B. jolonensis. Based on the vascular pattern of Niehaus #662 collected on Otay Mesa, we believe this species is also Coastal BTK rather than B. jolonensis.

Flowers of Brodiaea jolonensis from Monterey County showing the distinctive purple ovaries. According to Niehaus (1971), this is the only species of Brodiaea in California with purple ovaries. In fact, no populations of Montane BTK or Coastal BTK in San Diego County have ovaries this color.

The use of purple ovary color to identify Brodiaea jolonensis ideally applies to fresh flowers. Ovaries of other species in dried herbarium specimens often turn purplish-brown. The left photo of Brodiaea ovaries was taken from herbarium specimens all collected in San Marcos. A. Coastal BTK; B. San Marcos hybrid? brodiaea; C. Brodiaea filifolia.

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 indicates 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" is clearly evident in B. terrestris ssp. terrestris of central and northern California, but does not appear that useful in southern California populations of BTK.

Dentate Connective of Coastal BTK
Chromosome Numbers Of The San Marcos Brodiaea


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