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 Brodiaea jolonensis   Floras Citing Brodiaeas   Key To S. Calif. Brodiaeas   S. Calif. Brodiaea Images    Index Of Brodiaea Pages 

Brodiaea Species in San Marcos
© W.P. Armstrong, 27 May 2007
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)

  1. Images of Coastal BTK in San Marcos  
  2. Conclusions: San Marcos Coastal BTK    
  3. Sterile Hybrid With Flat Staminodes
  4. Possible Origin Of The Sterile Hybrid
  5. Fertile B. filifolia x B. orcuttii Hybrid
  6. Brodiaea orcuttii in San Marcos
  7. Brodiaea filifolia in San Marcos

1. Coastal BTK (B. terrrestris ssp. kernensis) in San Marcos

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

The status of these lovely wildflowers is rather dismal. They are surrounded by buildings in a rapidly growing area zoned for industry and shopping centers. Some of the vernal pools have been damaged by deep tire ruts and the dumping of debris. Raised topography (mounds) near vernal pools once supported burrowing owls. During years with sufficient rainfall, some pools are teeming with San Diego Fairy Shrimp (Branchinecta sandiegonenis).

  See Plants & Animals Of The San Marcos Vernal Pools  

Coastal BTK (Brodiaea terrestris ssp. kernensis) photographed 30 April 2006 in San Marcos. This species appears identical to populations on Otay Mountain (San Diego County) and the Santa Rosa Plateau (Riverside County). It does not have the dentate anther connective of B. terrestris ssp. terrestris. Although it has a distinct notch in the anther connective (between the anther sacs), similar to the V-shaped notch of B. jolonensis in Monterey County, the overall flower shape and dimensions are quite different. Its vasculature is different and its chromosome number is also higher than the reported numbers for B. jolonensis. This species is clearly not B. jolonensis.

A comparison of the anther axes of Brodiaea jolonensis, B. terrestris ssp. terrestris in Monterey County, and coastal & montane "BTK" in southern California. Some Coastal BTK in San Diego County have a minute lobe at the base of the V-shaped or U-shaped notch. B. terrestris ssp. kernensis in Kern County also has a dentate anther axis. In Chris Pires' treatment of the genus Brodiaea in Flora of North America On-line, the dentate anther connective tissue is not used in the separation of Brodiaea jolonensis from B. terrestris.

Coastal BTK (Brodiaea terrestris ssp. kernensis).

  Compare With B. jolonensis in Monterey County  

Coastal BTK (Brodiaea terrestris ssp. kernensis). Note the minute "dentate" appendage at the base of the notch in anther connective. This could be described as a W-shaped notch.

Coastal BTK (Brodiaea terrestris ssp. kernensis). The grass in photo is false brome (Brachypodium distachyon), a naturalized annual from southern Europe.

Coastal BTK in San Marcos: More than 97% of the pollen grains are fertile. Fewer than 3% are smaller, transparent blanks. Mature grains are approximately 50 micrometers (0.05 mm) long. An unfortunate flower thrip was dining on the anthers when the pollen sample was taken. This animal became part of the stained slide (lower left quadrant). (Magnification approximately 200 x.)

  See Table Of Relative Cell Sizes  

Coastal BTK in San Marcos: More than 98% of the pollen grains are fertile. Fewer than 2% are smaller, transparent blanks (red arrows). These percentages are not based on this image. They are based on several anthers from fertile BTK flowers. Mature grains are approximately 50 micrometers (0.05 mm) long. (Magnification approximately 100 x.)

Coastal BTK (Brodiaea terrestris ssp. kernensis) with fertile anthers.


2. Conclusions Regarding Coastal BTK In San Marcos

The species of Brodiaea with hooded staminodes appears to be Coastal BTK (Brodiaea terrestris ssp. kernensis) and not B. jolonensis. There is certainly some variability in the flowers of this species in San Marcos. The notch in the anther connective tissue may be V-shaped, U-shaped or W-shaped, with a minute appendage at the base of the notch. No flowers have the distinctive dentate connective of Brodiaea terrestris ssp. terrestris in Monterey County.

The ovary color, general floral dimensions and vascular pattern of Coastal BTK are different from B. jolonensis in Monterey County. This is discussed in more detail by clicking the red B. jolonensis tab at the top of this page. The chromosome number is at least 36 or more, too high for typical B. jolonensis in Monterey County. Coastal BTK identical to the San Marcos population occur elsewhere in coastal San Diego County, including Otay Mountain. Chromosome counts of the Otay Mountain population are currently under investigation by Dale McNeal of the University of the Pacific . Although no exact numbers have been determined at this time, preliminary studies indicate that the number may be higher than determinations made by Theodore Niehaus for Otay Mesa.

Two additional species of Brodiaea occur in the San Marcos field, including B. filifolia and B. orcuttii. The latter species generally occurs in the vernal pools, but may also grow intermixed with B. filifolia and Coastal BTK throughout the property.

Coastal BTK in the San Marcos population have distinctly hooded staminodes and anthers with generally 98% or higher pollen viability. These flowers would be considered fertile under Benson's criteria (1962). Smaller flowers with strap-shaped staminodes occur sparingly in the population. These flowers are sterile and their anthers contain no pollen grains. They appear to be sterile hybrids, possibly between Coastal BTK and B. filifolia or B. orcuttii. B. filifolia even appeared in a flower pot with corms of the transplanted hybrid. According to Niehaus (1971), hybridization occurs within sympatric populations of Brodiaea species. The progeny of these hybrid crosses generally have sterile flowers with no pollen. Hybrids can multiply asexually by cormlets. In addition, the field also contains a fertile hybrid between B. filifolia and B. orcuttii.

Coastal BTK (Brodiaea terrestris ssp. kernensis) in San Marcos photographed 26 April 2007. This was a relatively dry spring so very few flowers were in bloom.


Brodiaea Hybrids In San Marcos

3. Coastal BTK x Brodiaea filifolia or Brodiaea orcutii
   A Sterile Hybrid With Strap-Shaped Staminodes

Left: Typical Coastal BTK (Brodiaea terrestris ssp. kernensis) showing hooded staminodia and fertile, pollen-bearing anthers. Right: Smaller hybrid with strap-shaped staminodia and sterile anthers without pollen. This may be a hybrid between Coastal BTK and B. filifolia or B. orcuttii which also occur intermixed in the field. In fact, both B. filifolia and B. orcuttii appeared in a flower pot with corms of the transplanted hybrid.

Both of these species appeared in a flower pot with corms of the transplanted hybrid.

This appears to be a hybrid between Coastal BTK (Brodiaea terrestris ssp. kernensis) and B. filifolia or B. orcuttii which occur nearby. Note the strap-shaped staminodes that are not hooded as in typical coastal BTK. The flower was completely sterile with no mature pollen grains, giving credibility to the hybrid hypothesis. The clear liquid inside the reduced anther sacs contained cellular debris and scattered nuclei, but no well-formed pollen grains.

This appears to be a hybrid between Coastal BTK (Brodiaea terrestris ssp. kernensis) and B. filifolia or B. orcuttii which occur nearby. The strap-shaped staminodes are not hooded as in typical coastal BTK. The flower was completely sterile with no mature pollen grains, giving credibility to the hybrid hypothesis. In his "A Biosystematic Study of the Genus Brodiaea (Amaryllidaceae)", Univ. of Calif. Publications in Botany Vol. 60 (1971), Niehaus reported a cross between B. terrestris and B. coronaria which had a 100 percent seed set. The hybrid progeny of this cross had flowers that were morphologically intermediate between those of the two parents. Pollen fertility of each hybrid offspring was obtained, and the majority were completely sterile. According to Lyman Benson (Plant Taxonomy, The Ronald Press, New York (1962): "If, for example, more than half the pollen grains are abortive, probably something is amiss with the meiotic process preceeding pollen-grain formation." There are several causes for hybrid sterility, including the incompatibility between parental chromosomes during meiosis resulting in failure to pair up properly during synapsis of prophase I.

This appears to be a hybrid between Coastal BTK (Brodiaea terrestris ssp. kernensis) and B. filifolia or B. orcuttii which occur nearby. In fact, both B. filifolia and B. orcuttii appeared in a flower pot with corms of the transplanted hybrid. Note the strap-shaped staminodes that are not hooded as in typical coastal BTK. The flower was completely sterile with no mature pollen grains, giving credibility to the hybrid hypothesis.

This appears to be a hybrid between Coastal BTK (Brodiaea terrestris ssp. kernensis) and B. filifolia or B. orcuttii which occur nearby. Note the thin, envelope-shaped anthers that appear very different from typical BTK in San Marcos. The anther sacs were devoid of pollen grains. This was truly a sterile flower.

Left: Fertile, pollen-bearing stamen of Coastal BTK (Brodiaea terrestris ssp. kernensis). Right: Possible hybrid between Coastal BTK and B. filifolia or B. orcuttii which occur nearby. The small, thin anther sacs of the hybrid were devoid of pollen grains.

Vascular bundle composed of vessels in filament of Coastal BTK "hybrid."

Ovule-bearing ovaries of 3 Brodiaea species: A. Sterile hybrid of B. terrestris ssp. kernensis x B. filifolia or B. orcuttii showing minute aborted ovules. This is conclusive evidence that this is indeed a sterile hybrid incapable of producing seeds. B. Fertile hybrid of Brodiaea filifolia x B. orcuttii showing mature ovules. C. Brodiaea santarosae with mature ovules.

A sterile hybrid on the Santa Rosa Plateau of Riverside County. The flattened staminode has an elongate, subulate central tooth. One of the parents is undoubtedy Brodiaea terrestris ssp. kernensis that grows nearby. The other parent is either B. santarosae or B. filifolia which grow in the vicinity. If B. filifolia is a parent, then perhaps B. orcuttii is one of the parents of the San Marcos sterile hybrid because it lacks the elongate, subulate tooth.

   See Hybrid Brodiaea on the Santa Rosa Plateau   
Hybrid Brodiaea on the Santa Rosa Plateau


4. Origin & Of San Marcos Sterile Hybrid

Hypothesis #1

One possible explanation for hybrid sterility may be synaptic failure at prophase I of meiosis. Niehaus (1971) studied several populations of "Brodiaea jolonensis" in southern California and published sporophyte chromosome numbers of 36. We are certain that these populations are Coastal BTK and not B. jolonensis. My tentative count for Coastal BTK in San Marcos is at least 36. If these populations are hexaploid (6n = 36) with a base number of 6 (n = 6), a cross between a tetraploid (4n) BF (or BO) would result in a 5n (pentaploid) hybrid. Preliminary chomosome counts of Coastal BTK from Otay Mesa by Dale McNeal at University of the Pacific (personal communication, 2006) indicate that the number may be greater than 40. An exact count is difficult because small chromosomes are often obscured by larger ones.

Microsporogenesis in the San Marcos BTK showing the first division of a pollen mother cell (microsporocyte). Cytoplasmic division (cytokinesis) has not occurred yet. The two chromosome clusters (2 sets of chromosome doublets) contain at least 36 chromosomes, possibly more depending on how you count overlapping chromosomes. There are very small chromosomes that may be obscured by the larger ones. Brodiaea species are known to have base numbers of 6 and 8. [500 x]

Diploidization Of Old Polyploids

Summarized from: "Advances in the Study of Polyploidy Since Plant Speciation."
by D.E. Soltis, P.S. Soltis, and J.A. Tate. New Phytologist 161: 173-191 (2003).

Old genomes that are polyploid with respect to the base number and amount of genetic material, may function as diploids with respect to the level of gene expression and chromosomal characteristics. These "old polyploids" may have become "diploidized" by the loss, mutation or suppression of duplicate genes. Other causes for diploidization may include genomic rearrangements and transposons. This can drastically change the chromosome properties of a species. For example, an odd polyploid with a base number of six might have a sporophyte number of 5n = 30. This pentaploid would tend to be sterile because of an odd chromosome set at synapsis of prophase I. However, if this plant behaves as a diploid with 2n = 30, it would be fertile with two sets of 15 chromosomes during meiosis. It would simply have a diploid (sporophyte) number of 2n = 30 and a haploid (gametophyte) number of n = 15.

Hypothesis #2

With base numbers of 6 and 8 chromosomes, most species of Brodiaea are technically polyploids with multiple sets of chromosomes. For example, B. terrestris ssp. kernensis in Kern County is an octoploid with eight sets of chromosomes, and B. orcuttii is tetraploid with four sets of chromosomes. [B. jolonensis in Monterey County is diploid with two sets of chromosomes.] According to Niehaus (1971), the lack of quadrivalents in meiosis plus the differences in size of individual mitotic chromosomes at different ploidy levels suggests that these are "old polyploids." That is, the tetraploids, hexaploids, and octoploids have been in existence long enough to change by translocations, deletions, and so on. These differences apparently are enough to ensure that during meiosis only bivalents will occur in the higher ploidy levels. In bivalents, only two sets of maternal and paternal chromosomes associate during synapsis of meiosis, in contrast to quadrivalent where four sets of homologous chromosomes associate during meiosis.

If Brodiaea species behave as diploids, then BF and BO each have n = 12 and 2n = 24, and BTK has n = 18 or a higher number. In fact, BTK could be 2n = 36, 38, 40, 42, 44, 46, 48, etc. and still be theoretically fertile with two sets of homologous chromosomes. A hybrid between BF or BO and BTK would be sterile because of the non-homologous pairs of chromosomes that differ in number. Hybridization may occur, but the resulting hybrid offspring grown from seed may by sterile without viable pollen. For example, a cross between BF and BTK could result in a hybrid with 12 BF chromosomes and 18 BTK chromosomes which would not match up during synapsis of prophase I. This hypothesis is a plausible explanation for the sterile hybrids we have observed in San Marcos.

  More Images Of The Sterile San Marcos Hybrid Brodiaea  


5. Fertile Brodiaea filifolia x B. orcuttii Hybrid in San Marcos

A comparison of the flowers of B. filifolia (left) and B. orcuttii (right). B. filifolia has short filaments only 1.0 mm in length and small, subulate staminodes (white arrow). B. orcuttii has longer filaments up to 8.0 mm in length and no staminodes. Because of their reduced staminodes, Niehaus considers these to be more advanced species. They both have the same chromosome number (n = 12) and cross pollinate producing a fertile hybrid with staminodes and long filaments (see next section). One or both species also cross pollinate with nearby Coastal BTK (B. terrestris ssp. kernensis) in San Marcos, producing sterile hybrids (see above section) with strap-shaped staminodes and reduced anthers that produce no pollen.

A Brodiaea filifolia x B. orcuttii hybrid in San Marcos. It has the flower size and long filaments of B. orcuttii, but has the flower shape and filiform staminodes of B. filifolia. Flower dimensions: Perianth = 24 mm, style = 8.5 mm, filament = 5.5 mm, anther = 5.5 mm, ovary = 4.9 mm, staminode = 2.5 to 4.5 mm. The flower dimensions are smaller than the population resembling B. orcuttii and B. filifolia on the Santa Rosa Plateau. The anthers definitely contained pollen. The fertility can be explained because the cross involves two tetraploid hybrids with the same sporophyte chromosome number of 24. The hybrid is probably also tetraploid (4n = 24). Ovaries of a potted hybrid plant contained abundant mature ovules, conclusive evidence that this is indeed a fertile hybrid. See image B above.

1.   Brodiaea santarosae on The Santa Rosa Plateau
2.   Brodiaea santarosae on The Santa Rosa Plateau
3.   Lecture Notes On Santa Rosa Basalt Brodiaea

San Marcos Brodiaea filifolia x B. orcuttii hybrid showing slender, erect staminodia. The filiform staminodes resemble B. filifolia, but the long filaments resemble B. orcuttii.

San Marcos Brodiaea filifolia x B. orcuttii hybrid showing slender, erect staminodia. The filiform staminodes resemble B. filifolia, but the long filaments resemble B. orcuttii.

Brodiaea filifolia with erect staminodia. The filaments are short (1.0 mm), unlike the B. filifolia x B. orcuttii hybrid (see above) which has filaments of 5.5 mm. Flower dimensions: Perianth = 20 mm, style = 6.0 mm, filament = 1.0 mm, anther = 4.0 mm, ovary = 5.0 mm, staminode = 4.5 mm. This flower has all the characteristics and dimensions of B. filifolia except the erect staminodes. In most B. filifolia flowers examined from this field, the subulate staminodes are reflexed against the perianth (see following images).

  More Images Of The Fertle San Marcos Hybrid Brodiaea  


6. Brodiaea orcuttii in San Marcos

Brodiaea orcuttii: This species has no staminodes and is considered more highly evolved according to Niehaus (1971). The flowers are larger than B. filifoloia.

Brodiaea orcuttii: This species has no staminodes and is considered more highly evolved according to Niehaus (1971). The flowers are larger than B. filifoloia.

In June 2006, numerous flower beetles [Coleoptera--Melyridae--Dasytinae] were feeding on the flowers of Brodiaea orcuttii. These beetles are obviously major pollinators for this species. This species was identified by Bob Allen as soft-winged flower beetle (Emmenotarsalis quadrimaculata).

Vascular pattern of inner perianth segment of Brodiaea orcuttii. According to Niehaus (1971) there are two outer strands with a double strand in the center (1-2-1). In this image there are possibly two strands in the center; however, farther along the midvein, the lower vascular bundle separates into multiple strands. Segment cleared with 10% NaOH but not stained. (200 x).

Brodiaea orcuttii in San Marcos (May 19, 2007).

Brodiaea orcuttii in a fenced vernal pool area adjacent to Fry's Electronics in San Marcos. The site is managed by the San Diego Zoological Society. Photo taken 1 May 2007.


7. Brodiaea filifolia in San Marcos

Brodiaea filifolia with subulate staminodes reflexed against perianth.

Brodiaea filifolia with subulate staminodes reflexed against perianth.

Brodiaea filifolia, a rare species with small, filiform or subulate staminodes that typically spread outwardly (away from the stamens).


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