BrodiaeaPhylogeny

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Comments On Cladistic Groupings Of 14 Brodiaea Samples
From Monterey & Kern Counties To The Mexican Border
© W.P. Armstrong 04 July 2017
Based On 14 Brodiaea Collections I Made For The Unpublished
Phylogenetic Analysis Of Brodiaea by A.R. Diebold, et al. 2012

Note: I do not have permission to post this unpublished article or DNA cladograms. This brief summary is only an attempt to explain the groupings of my 14 samples in the cladogram. Some samples appear where I would expect them because of their morphological relationships based on my extensive images of flowering specimens. Other samples I cannot explain. For example, samples of Brodiaea terrestris ssp. kernensis from Kern County and the San Diego mountains that appear closely related morphologically are widely spaced from each other in the cladogram.

Groupings Of My Collections

1. Brodiaea terrestris ssp. terrestris (Coastal & Inland)

WPA1: BTT Coastal Monterey Co.
WPA2: BTT Inland SLO Co. (La Panza Area)

2. Brodiaea terrestris ssp. kernensis (Coastal)

WPA5: BTK Coastal SD Co. (Otay Mesa)
WPA13: BTK Coastal SB Co. (Gaviota Pass)

3. Brodiaea terrestris ssp. kernensis (Montane)

WPA3: BTK Montane SD Co. (Cuyamaca Lake)
WPA4: BTK Montane SD Co. (Laguna Mtns)
WPA6: BTK Central Kern County

4. B. orcuttii, B. filifolia, B. santarosae & 2 Hybrids

WPA12: B. orcuttii SD Co. (San Marcos)
WPA15: B. orcuttii SD Co. (San Marcos)
WPA11: B. filifolia SD Co. (San Marcos)
WPA14: B. filifolia SD Co. (San Marcos)
WPA8: B. santarosae Riverside Co. (SRP)
WPA9: BO x BF SD Co. (San Marcos)
WPA16: BO or BF x BTK (San Marcos

1. Brodiaea terrestris ssp. terrestris (Coastal & Inland)

Although they are listed as the same species, they were not grouped together in closely related clades. Based upon careful observations of cloned specimens and images of flowering plants, the two samples did exhibit morphological variation.


2. Brodiaea terrestris ssp. kernensis (Coastal)

These coastal forms of BTK from Santa Barbara & San Diego Counties were grouped very close to each other. This is not surprising because they appeared very similar morphologically based on my images of flowering plants. They were also grouped very close to B. jolonensis in the cladogram, suggesting a possible kinship. Perhaps this is why the Jepson Manual 2nd Edition lists this brodiaea with white staminodes as a possible undescribed taxon under B. jolonensis. I have also observed individuals with white and purple staminodes in the same population. Morphology data from Tom Chester and myself indicate this taxon is more closely related to B. terrestris ssp. kernensis. The latter species also has higher polyploid chromosome numbers (6n = 36, 7n = 42, & 8n= 48) compared with B. jolonensis (2n=12).

My tentative sporophyte 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.

  Brodiaea jolonensis Does Not Occur In Southern Californa  
Brodiaea jolonensis Does Not Exist In Southern California
San Marcos Sterile Hybrid: BTK x B. filifolia or B. orcutii?


3. Brodiaea terrestris ssp. kernensis (Montane)

The groupings of these 3 collections are perplexing. BTK from Cuyamaca Lake and the Laguna Mountains, San Diego County, were not closely grouped. Even more surprising was the grouping of BTK from Kern County which was near the bottom of cladogram. I must admit that I have no explanation for this unless there was an error in DNA sequencing or my samples.


4. Brodiaea orcuttii, filifolia, santarosae Plus Hybrids

Brodiaea orcuttii and B. filifolia, plus hybrids between them, are closely related and are grouped close to each other and B. santarosae. This arrangement is what I would expect.

Key To Brodiaea Species in Mainland & Insular Southern California
South of San Luis Obispo & Kern Counties to the Mexican Border
© W.P. Armstrong 26 May 2009

 Pictorial Key To Brodiaeas In San Diego County 

1  Staminodes 0 in all flowers.....B. orcutii

1' Staminodes present in 90-100% of flowers.....(2)

2  Staminodes filiform or uniformly tapered from base to tip.....(3)

2' Staminodes oblong to rectangular in outline, tapering only near tip if at all.....(5)

3  Filaments 0-1.5 mm; staminodes 1.0-4.5 mm long, reflexed against perianth.....B. filifolia

3' Filaments 2-8 mm; staminodes 0.0-7.0 mm long, recurved to erect.....(4)

4  Perianth length 19-24 mm; style 8.0-9.5 mm; ovary 4.0-5.0 mm; anther 5.0-5.5 mm; San Marcos,
San Diego County.....B. filifolia x B. orcuttii

4' Perianth length 24-36 mm; style 10.5-17 mm; ovary 3.5-8.2 mm; anther 5.4-8.9 mm; Santa Rosa Plateau,
Elsinore Peak (Riverside County) to Miller Mountain (San Diego County).....B. santarosae

5  Staminodes gen erect, edges flat to inrolled, sometimes hooded, gen purple.....(6)

5' Staminodes recurved, edges flat, not hooded, pointed at tip, gen white; Mission Trails Regional Park,
San Diego County.....B. elegans ssp. elegans

6  Staminodes inrolled or hooded at apex, lvs 2-3; mainland & insular.....B. terrestris ssp. kernensis.....(7)


7 Staminodes typically hooded at apex; coastal and insular; Santa Barbara Co. to n. Baja Calif.....Coastal BTK*

7' Staminodes flattened or inrolled along margins; Kern County and montane southern California.....Montane BTK*


6' Staminodes flat, cuspidate at apex, lvs 1; San Clemente Island.....B. kinkiensis

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 from Tom Chester thus far indicates that BTK is one variable species. 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


Images of Brodiaea Species in Mainland & Insular California
Monterey, San Luis Obispo & Kern Counties to the Mexican Border
© W.P. Armstrong, 18 October 2007
**  In general, coastal popultions of BTK have hooded staminodes; however, there is tremendous
variation in staminode shape in other populations, such as the Santa Rosa Plateau and Cuyamaca
Lake.  Potted plants with non-hooded, inrolled staminodes from the Laguna Mountains developed
hooded staminodes like coastal BTK when grown in Escondido. Click Here To View Image.
The flower color for Brodiaea species on this page is blue-purple to violet. In all the following images
I have attempted to match their original color;  however, they may appear different on your monitor.
Brodiaea jolonensis (Monterey County)

  Brodiaea jolonensis Not in Southern Calif.  
Brodiaea jolonensis in Monterey County


Brodiaea terrestris ssp. terrestris (Coastal Monterey County)


Brodiaea terrestris ssp. terrestris (Inland San Luis Obispo County)

  B. terrestris ssp. terrestris in San Luis Obispo County  


Brodiaea terrestris ssp. kernensis (Kern County)

  Brodiaea terrestris ssp. kernensis in Kern County  


Brodiaea terrestris ssp. kernensis (Coastal BTK In Southern California)

  Coastal BTK in San Marcos (Coastal San Diego County)  
Coastal BTK above Gaviota Pass (Santa Barbara County)


Brodiaea terrestris ssp. kernensis (Montane BTK In Southern California)

Brodiaeas at Kearny Mesa & Cuyamaca Lake
  Staminode Variation in BTK at Cuyamaca Lake  


Brodiaea terrestris ssp. kernensis (Santa Rosa Plateau, Riverside County)

Brodiaeas on Santa Rosa Plateau & Elsinore Pk
  Staminode Variation in BTK at Santa Rosa Plateau  


Brodiaea filifolia (San Diego & Riverside Counties)

Brodiaea filifolia in San Marcos, San Diego Co.
  Brodiaeas on Santa Rosa Plateau & Elsinore Pk  


Brodiaea orcuttii (Coastal & Montane San Diego County)

  Brodiaea orcuttii in San Marcos, San Diego Co.  
Brodiaeas at Kearny Mesa & Cuyamaca Lake


Brodiaea santarosae (Santa Rosa Plateau, Riverside County)

The flower of Brodiaea santarosae has longer perianth segments than other Brodiaea species in southern California. The total spread of this flower is nearly 50 mm. This species has long filaments like B. orcuttii, but has much larger flowers. It has no staminodes or slender staminodes that vary in length. It is similar to the fertile hybrid between B. filifolia and B. orcuttii, except the flowers are generally about 40 percent larger than the hybrid. See following image comparison between this species and the hybrid.

  Brodiaeas on Santa Rosa Plateau & Elsinore Pk  
Brodiaea santarosae on the Santa Rosa Plateau
Variation in Flowers of Brodiaea santarosae


Brodiaea santarosae compared with B. filifolia x B. orcuttii

  San Marcos Brodiaea Hybrids  
1. Fertile Hybrid: B. filifolia x B. orcuttii
2. Fertile Hybrid: B. filifolia x B. orcuttii
3. Sterile: BTK x B. filifolia or B. orcutii

  Hybrid On Santa Rosa Plateau  
1. A Sterile Hybrid: B. santarosae x BTK


Brodiaea elegans ssp. elegans (San Diego County)

Is Brodiaea elegans in San Diego County?
  B. elegans at Mission Trails Regional Park  


Brodiaea kinkiensis (San Clemente Island)

  Brodiaea kinkiensis on San Clemente Island  


Brodiaea species native to southern California: (A) B. orcuttii, (B) B. filifolia, (C) B. santarosae, (D) B. terrestris ssp. kernensis, (E) B. kinkiensis, (F) B. elegans ssp. elegans.

Index Of Brodiaea Pages On Wayne's Word
PDF of B. santarosae Article Published in Madrono Vol. 54: 187-198 (2007)

  1. Floras Citing Brodiaeas In Southern California
  2. Key To Brodiaea Species In Southern California
  3. Brodiaea Images: Central & Southern California
  4. Key To Brodiaea Species in San Diego County
  5. Is Brodiaea jolonensis in San Diego County?
  6. Is Brodiaea elegans in San Diego County?

    Brodiaeas In San Marcos          

  7. San Marcos Vernal Pools & Brodiaea Populations
  8. More Images From San Marcos Vernal Pool Area
  9. Coastal BTK in San Marcos During Spring 2006
  10. Coastal BTK x B. filifolia or B. orcuttii In San Marcos
  11. San Marcos Brodiaeas After March Rains of 2006
  12. San Marcos Vernal Pools During Spring Of 2008
  13. San Marcos Vernal Pools During Spring Of 2010
  14. San Marcos Vernal Pools During Spring Of 2011
  15. San Marcos Vernal Pools During Spring Of 2012
  16. Statement of Brodiaea Research in San Marcos

    Hybrid Brodiaeas In San Marcos          

  17. Fertile Brodiaea filifolia x B. orcuttii In San Marcos
  18. Fertile San Marcos Hybrid Grown In Flower Pot 1
  19. Fertile San Marcos Hybrid Grown In Flower Pot 2
  20. Sterile Hybrid Brodiaea (Coastal BTK x BF or BO)

    Brodiaeas In Central & S. California          

  21. Staminode Variation In BTK At Camp Pendleton
  22. Kearny Mesa, Mission Trails & San Marcos
  23. Brodiaeas at Kearny Mesa & Cuyamaca Lake
  24. Staminode Variation in BTK at Cuyamaca Lake
  25. Brodiaea terrestris kernensis in Kern County
  26. B. terrestris terrestris in San Luis Obispo County
  27. Brodiaea jolonensis in Monterey County
  28. Coastal BTK in Santa Barbara County (Gaviota Pass)   
  29. Coastal BTK in Santa Barbara County (Figueroa Mt)
  30. Brodiaea kinkiensis on San Clemente Island
  31. Pollen Of B. elegans Variant at Mission Trails
  32. Anthers Of BJ? vs. Coastal & Montane BTK
  33. The Vascular Pattern of Perianth Segments
  34. Vascular Strands of BJ From Monterey County
  35. Vascular Pattern of #662 From Otay Mesa
  36. Brodiaea Leaves On Sprouting Potted Plants
  37. Brodiaeas Grown In Pots Spring 2008-09 (1)
  38. Brodiaeas Grown In Pots Spring 2008-09 (2)

    Brodiaea santarosae            

  39. Floral Variation in Brodiaea santarosae
  40. Images Of Basalt On The Santa Rosa Plateau
  41. Brodiaea santarosae Lecture (Blue Background)
  42. Brodiaea santarosae Lecture (White Background)
  43. Images Of Brodiaeas on the Santa Rosa Plateau
  44. Brodiaeas on Santa Rosa Plateau & Elsinore Pk
  45. Brodiaeas on the Santa Rosa Plateau (2006)
  46. Brodiaea santarosae On Santa Rosa Plateau (1)
  47. Brodiaea santarosae On Santa Rosa Plateau (2)
  48. Brodiaea santarosae On SRP: Pollinator Bee
  49. Staminode Variation in BTK at Santa Rosa Plateau    
  50. Brodiaea santarosae cf. B. filifolia & B. orcuttii
  51. Rock Formations On The Santa Rosa Plateau
  52. Rock Lichens On The Santa Rosa Plateau
  53. Miller Mt. Location For Brodiaea santarosae
  54. Santa Rosa Plateau Wildflowers 6 May 08
  55. Santa Rosa Plateau Wildflowers 22 May 08
  56. Brodiaea Hybrid On The Santa Rosa Plateau
  57. Brodiaeas On Miller Mt & Santa Rosa Plateau