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Vascular Pattern of BTK Perianth Segments

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)

The midvein of Brodiaea perianth segments contains three primary vascular bundles. Each bundle is composed of numerous vessels with spirally thickened secondary cell walls that appear like coiled springs under a compound microscope. The bundles of some species separate into strands of vessels. According to Niehaus (1971), the number of strands is characteristic of certain species. In BTK, vascular bundles of inner perianth segments separate into multiple strands, up to three strands per bundle. Bundles of B. jolonensis typically do not separate into strands, although I have observed bundles with two strands. Because of overlapping vascular strands and the limited depth of field with compound light microscopes, it is often difficult to capture the well-defined strands of all three vascular bundles in a single image. Strands are not always separated from each other along a vein, and appear to merge in places. In damaged or poorly dried (wrinkled) perianth segments, strands may be broken. Individual vessels may have separated from the strands or be out of focus. In general, B. jolonensis has fewer well-defined strands compared with Coastal BTK in southern California.

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."

Table Of Contents:
  1. BTK in San Marcos
  2. BTK on the Santa Rosa Plateau
  3. BTK on Catalina Island
  4. BTK on San Miguel Island
  5. BJ in Monterey County


1. BTK in San Marcos

Magnified horizontal view (100x) of an inner perianth segment of Coastal BTK in San Marcos showing three primary vascular bundles each composed of multiple strands of tracheary elements (vessels). The strands consist of vessels with spirally thickened walls. Niehaus (1971) represented this strand pattern as 3-3-3. Although the triplet banding for each bundle is difficult to interpret from this image, the fact that each bundle contains several strands is proof that this is not B. jolonensis which has a total of only 3 or 4 strands. The perianth segment was cleared by soaking it in NaOH.

Horizontal magnified view (100x) of an outer perianth segment of Coastal BTK from San Marcos showing three primary vascular bundles. Niehaus (1971) represented this strand pattern as 1-2-1. The outer bundles consists of single strands while the center bundle consists of two strands. The strands consist of several vessels with spirally thickened walls. B. jolonensis also has this 1-2-1 strand pattern in its outer perianth segments. The staminodes are attached to the outer perianth segments.

Vascular Strands Of Brodiaea Perianth


2. BTK on the Santa Rosa Plateau

Magnified view (100x) of an inner perianth segment of Coastal BTK showing three primary vascular bundles. Each bundle is subdivided into three strands of tracheary elements (vessels). The strands consists of several vessels with spirally thickened secondary walls. Niehaus (1971) represented this strand pattern as 3-3-3. B. jolonensis has noticeably fewer strands, a total of three or four. The perianth segment was cleared by soaking it in NaOH and stained with safranin. The strands were darkened slightly using Adobe Photoshop. Spiral thickenings in the vessel walls appear as transverse bands under high magnification. In fact, the triplet strands superficially resemble an aerial view of three sets of railroad tracks.

Left: Magnified view (100x) of an inner perianth segment of Coastal BTK from the Santa Rosa Plateau showing three primary vascular bundles. Each vascular bundle contains multiple strands of tracheary elements. The strands consist of vessels with spirally thickened walls. Although the triplet strands are not well-defined on all three bundles, they are fairly distinct in the right image. Niehaus (1971) represented this strand pattern as 3-3-3. Right: Magnified view (400x) of the far right vascular bundle (red square) showing three strands of tracheary elements (vessels). The perianth segments were cleared by soaking them in NaOH. No stains or color enhancements were used on this image.


3. BTK on Santa Catalina Island

Magnified view (100x) of an inner perianth segment of Coastal BTK from Santa Catalina Island showing three primary vascular bundles (shown in three transparent color bands). Each bundle is subdivided into multiple strands of tracheary elements (vessels). The strands consist of several vessels with spirally thickened walls. Niehaus (1971) represented this strand pattern as 3-3-3. The fact that each bundle contains several strands indicates that this is not B. jolonensis which has a total of only three or four strands in the entire vein. The perianth segment was cleared by soaking it in NaOH. Spiral thickenings in the vessel walls appear as transverse bands under high magnification. This photo is from a pressed specimen labeled B. jolonensis.

Magnified view (100x) of an inner perianth segment of Coastal BTK from Santa Catalina Island showing seven distinct vascular strands (highlighted in transparent colors). Each strand consist of one-several vessels with spirally thickened walls. The lower blue band probably represents more than one strand. B. jolonensis can be ruled out here because it has a total of only three or four strands in the entire vein. This photo is from a pressed specimen labeled B. jolonensis.

Vascular Pattern of Brodiaea jolonensis

Magnified view (400x) of an inner perianth segment of Coastal BTK from Santa Catalina Island showing multiple strands of tracheary elements (vessels). The strands consist of vessels with spirally thickened walls. More vascular strands were directly above this view. The perianth segment was stained with safranin. This photo is from a pressed specimen from Santa Catalina Island labeled B. jolonensis.


4. BTK on San Miguel Island

Brodiaea flower from San Miguel Island. The floral dimensions, vasculature of inner perianth segments, and green ovary indicate that it is Coastal BTK and not B. jolonensis. Flower courtesy of Michael Charters.

Magnified view (400x) of an inner perianth segment of Coastal BTK from San Miguel Island showing 3 primary vascular bundles. Each bundle is subdivided into three strands of tracheary elements (vessels). The strands consists of vessels with spirally thickened secondary walls. Niehaus (1971) represented this strand pattern as 3-3-3. B. jolonensis has noticeably fewer strands, a total of three or four, compared with a total of nine for BTK. The perianth segment was cleared with 10% NaOH and stained with safranin.

Magnified view (400x) of an inner perianth segment of Coastal BTK from San Miguel Island showing multiple strands of tracheary elements (vessels). The strands consists of vessels with spirally thickened secondary walls. B. jolonensis from Monterey County has noticeably fewer strands, a total of three or four, compared with a total of nine for BTK. The perianth segment was cleared with 10% NaOH and stained with safranin.


5. Brodiaea jolonensis from Monterey County

Pressed flower of B. jolonensis showing prominent midvein on perianth segments. Inner segments are opposite the anthers, while outer segments are opposite the staminodes.

Inner perianth segment of Brodiaea jolonensis from Monterey County. Segment cleared with 10% NaOH and stained with safranin to highlight the vascular strands.

Microscopic view of the inner perianth segment of Brodiaea jolonensis from Monterey County. There are three distinct vascular strands, each composed of several intertwined vessels with spirally thicked secondary cell walls. This appears to fit the 1-1-1 strand pattern described by Niehaus. (Magnification 300 x).

Magnified view (400x) of an inner perianth segment of Brodiaea jolonensis from Monterey County showing three strands of tracheary elements (vessels). Each strand consist of vessels with spirally thickened walls. This pattern appears to fit the 1-1-1 pattern described by Niehaus (1971). In Coastal BTK there are three triplet sets of strands, a total of nine strands. Perianth segment cleared with 10% NaOH and stained with safranin. The bundle of raphide crystals was selectively sharpened in this image.

Magnified view (100x) of an inner perianth segment of Brodiaea jolonensis from Monterey County showing four strands of tracheary elements (vessels). Each strand consist of several vessels with spirally thickened walls. This pattern appears different from the 1-1-1 pattern described by Niehaus (1971). In Coastal BTK there are three triplet sets of strands, a total of nine strands. Although this specimen does not fit Niehaus' 1-1-1 pattern for B. jolonensis, it is clearly different from the 3-3-3 pattern of Coastal BTK in southern California.

Magnified view (400x) of an inner perianth segment of Brodiaea jolonensis from Monterey County showing four vascular strands. Each strand consists of several vessels with spirally thickened walls. This pattern is different from the 3-strand pattern described by Niehaus. The four strands appeared to merge into three strands in some places along the vein, and this may explain Niehaus' 1-1-1 pattern. In Coastal BTK there are three triplet sets of strands, a total of nine strands. Although this image does not exactly fit Niehaus' 1-1-1 pattern for B. jolonensis, it is clearly different from the 3-3-3 pattern of Coastal BTK in southern California. The four strands were colorized with Adobe Photoshop.

Magnified view (100x) of an inner perianth segment of Brodiaea jolonensis from Monterey County showing 4-6 vascular strands. Each strand consists of several vessels with spirally thickened walls. This pattern is different from the 3-strand pattern described by Niehaus. The strands appear to merge and separate along the perianth segment. Although this image does not exactly fit Niehaus' 1-1-1 pattern for B. jolonensis, there are clearly fewer strands than typical Coastal BTK in southern California.

Comparison of BTK and B. jolonensis Vascular Patterns


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