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Flower structure follows a basic plan with modifications, which results in enormous and sometimes overwhelming diversity. Flower diversity is much more manageable if one first understands the basic flower plan. Lillies, having large, complete flowers with no fused parts, are used in this section as model organisms to illustrate flower morphology. Subsequent sections include altered flower structures, resulting in significant variation.

Complete flowers comprise multiple sepals, petals, stamens, and one or more pistils arranged in concentric whorls. The outermost structures are the sepals. Sepals are leaf-like. They enclose and protect a developing bud. Sepals are generally thicker than petals. They hold in moisture and block insects and fungal and bacterial spores from entering the bud.

Moving inward from the sepals is a ring of petals. Petals attract pollinators. They are often thin, delicate, and colorful. 

The third whorl, located inside the ring of petals, is composed of stamens. A stamen has an anther at the top supported by a stalk called the filament. The anther produces pollen, and the filament raises the anther to a location where a pollinator or the wind can easily pick up the pollen.

Finally, the innermost structures are pistils. Some flowers have only one pistil. Others, such as buttercups, can have many pistils.  At the top of the pistil is the stigma, which receives pollen. The base of the pistil is the ovary, which will form the fruit containing seeds at maturity. The style connects the stigma and ovary. The ovary photographed here is "superior" because it is located above the base of the other floral whorls.

Consider the arrangement of flower structures when viewing an unfamiliar species.  Pistils are in the center surrounded by stamens, surrounded by petals, surrounded by sepals. It is all about structural arrangement: location, location, location. 

Some of the showiest flowers in the Sierra Nevada are in the lily family, Liliaceae. Lilies have three sepals, three petals, six stamens, and a single pistil having three compartments (carpels). The lily from the previous section is a Calochortus lily. The Calochortus genus includes mariposa lilies, star tulips, and globe lilies. Forty-two Calochortus species (57 taxa) out of approximately 70 total species are native to California, making the state the hub of Calochortus diversity. 

Butterfly Mariposa Lily (Calochortus venustus) is a California endemic species that grows in the western Sierra foothills and California's coastal mountains. The primary petal color may be white, yellow, pink, or red, with diverse interior markings, including a deep red spot on each petal. The Butterfly Mariposa Lily may or may not have a secondary color spot further up on the petals.  “Mariposa” is Spanish for butterfly, which refers to the colorful petals of these plants.

Mariposa lilies have nectar-producing glands near the base of each petal. These nectaries can be useful in distinguishing between similar species. The Butterfly Mariposa Lily’s nectaries are squarish in shape.

The Superb Mariposa Lily (Calochorus superbus) and Butterfly Mariposa Lily are similar in appearance and have ranges that overlap. Both species have beautiful, highly variable petal colors and a deep red spot often capped with yellow on each petal.  The great diversity of petal color and markings within each species, yet significant similarity between the two species, makes them difficult to distinguish.

A notable difference between the Butterfly and Superb Mariposa Lilies is that the nectar gland on the lower portion of each petal is crescent-shaped on the Superb Mariposa Lily rather than square as with the Butterfly Mariposa Lily.  The secondary spot on the upper petal is always absent on the Superb Mariposa Lily.

Gold Nuggets or Yellow Mariposa (Calochortus luteus) could be confused with yellow varieties of the previous two mariposa lilies. This plant has deep yellow petals with red-brown markings. A thin circle of dense, short hairs near the base of the petals distinguishes this plant from other species. Gold Nuggets grow at lower elevations up to only 2,300 feet. Gold Nuggets and Superb Mariposa hybridize.

Leichtlin’s Mariposa Lily (Calochortus leichtlinii) is a common plant with a higher elevation range and later blooming season than the previous species. The lily petals are generally white with sometimes a tinge of blue or pink; anthers are white. Leichtlin’s Mariposa Lily has a maroon-black spot of variable size located above yellow nectaries.  

The previous Calochortus lilies can grow to almost two feet, while the Sierra Mariposa Lily (Calochortus minimus) is a small species, reaching only about 4 inches in height. These plants produce up to five white flowers with white, blue, or purple anthers and have one grass-like leaf that can grow up to eight inches long. The Sierra Mariposa Lily is found exclusively in moist, open woodlands of the Sierra Nevada Mountains.

Yellow Star Tulip (Calochortus monophyllus) is another low-growing, less than eight inches, Sierra endemic species. This plant grows in wooded locations below 4,000 feet in the northern western foothills. 

Yellow Star Tulip is easily identified by its deep yellow color and densely hairy petals. There is often a maroon spot at the base of each petal.

White Globe Lily or Fairy-Lantern (Calochortus albus) grows in the western Sierra foothills and coastal mountains. It may range from eight to 32 inches in height. Due to its nodding and closed flowers, this plant may not initially look like a Calochortus species.  

Like the other Calochortus species, the White Globe Lily has linear grass-like leaves. When the flower is flipped upright and opened, a structure similar to that of other Calochortus species is revealed.

Yellow Star Tulip and White Globe Lily live in foothill woodlands on the western slope of the Sierra Nevada Mountains. Each has 20 chromosomes (2n=20) and a similar flowering period.  Sometimes, these two plants grow just a few feet apart. Awkward-appearing plants may also be found in these locations, often with partially nodding or partially open pale yellow flowers. These unusual plants are likely hybrids of the Yellow Star Tulip and White Globe Lily. 

Some lilies are particularly spectacular because their large sepals change from green to the color of the petals as they mature. This gives the impression of six petals, which increases the pollinator’s attraction to the plant. When sepals and petals look similar, they are known as tepals. The origin of the tepal (sepal or petal) can still be recognized by the locations of their whorls on the flower. Here, Sierra lilies with six tepals are highlighted.

The first four lilies presented are of the Lilium genus. They are all tall plants with leaves that whorl around their stems.

Kelly’s Lilly (Lilium kelleyanum) grows along streams and in wet thickets from 7,000-11,000 feet in elevation. This plant can be found in the Northern and Southern Sierra but is absent in the Central Sierra counties. The Kelley’s Lily's tepals are typically solid yellow-orange with red-maroon spots.

The Alpine Lily (Lilium parvum) grows in wet meadows and thickets at elevations up to 9,500 feet in the Sierra and Cascade ranges.  A single plant may grow up to 5 1/2 feet tall and have numerous small funnel-shaped flowers. 

Flowers of the Alpine Lily are yellow-orange to red-orange with maroon spots. Tepal color is generally a paler yellow-orange near the center and deeper or darker orange at the tips.

California Leopard Lily (Lilium pardalinum) has colors similar to the Alpine Lily but larger nodding flowers with more retracted tepals and longer stamens.  This plant is found in mountain and coastal regions from Southern Ontario to Southern California at low to mid-elevations.  A similar-appearing plant, Humboldt’s Lily (Lilium humboldtii ), grows in the northern counties of the western Sierra foothills.

The largest blossom in California belongs to the Washington Lily (Lilium washingtonianum), named to honor Martha Washington. It grows in dry, open forests below 6,200 feet in California and Oregon. The tepals of this plant's fragrant flower are white, rarely pink, with yellow midribs and small purple spots.

As flowers mature, the ovaries develop into fruits containing seeds.  (In some plants, structures surrounding the ovary also contribute to the fruit.) Within the lily family, one of two fruit types is produced. The fruit of plants like the Washington Lily is a dry seed pod known as a capsule. The capsule opens at maturity, releasing seeds stacked within three chambers. The other fruit type found in the lily family is a fleshy berry. See Hooker's Fairy Bells as an example.

Brown Bells or Brown Fritillary (Fritillaria micrantha) grows exclusively in the Sierra's yellow pine and red fir forests. The plant’s slender stem grows up to three feet with thin alternate leaves, several spring-blooming flowers above, and one to three whorls of four to six leaves below. Brown Bell’s small maroon or green flowers may be tricky to spot. In non-flowering years, the plant will produce one large nurse leaf, which photosynthesizes to trap energy for storage in the bulb. 

Fawn lilies in the Erythronium genus are generally smaller forest plants with basal leaves. Tuolumne Fawn Lily (Erythronium tuolumnense) is a rare plant with distribution limited to a few populations in Tuolumne County and bordering regions of Calaveras County. It grows in woodlands, especially on shady, moist, steep canyon slopes.  

Tuolumne Fawn Lily generally grows up to a foot in height. Here, a smaller-than-typical specimen is dwarfed in appearance by a giant banana slug. These organisms share the same moist forest habitat. 

Do not disturb the Tuolumne Fawn Lily. Native lilies are available from reputable nurseries without harming native populations.

The Purple or Plain Leaf Fawn Lily (Erythronium purpurascens) is a graceful plant that grows in open forests of the northern counties in the Sierra Nevada. Plants have two wavy leaves and a stalk with one to six flowers. Tepals are white with a yellow base. The white regions turn pink or purple as the flower ages.

Bride’s Bonnet (Clintonia uniflora) is a forest plant most commonly found between 3,000 to 5,000 feet. It spreads by rhizomes to produce an attractive ground cover. The plant is shade-tolerant and grows well at the edge of canopy openings. Each Bride’s Bonnet plant has 2 to 3 leaves and one (occasionally two) white star-like flowers. The plant produces blue berries eaten by thrush, grouse, and deer.

Hooker's Fairy Bells (Prosartes hookeri) is a forest species with attractive foliage growing up to two feet in height. This plant produces small, bell-shaped, green to cream-colored flowers on the underside tips of branches. At this location, the flowers are often hidden and overlooked. 

In sharp contrast to the inconspicuous flowers produced by Hooker’s Fairy Bells in late spring/early summer, attractive bright red, egg-shaped berries appear to drip off the stem tips in late summer. These berries provide food for ground-foraging birds and slugs. 

Flowers can vary greatly, making the identification of structures complicated. Here, we test the concentric whorls model of floral structures on the complex flower of Hartweg’s Iris (Iris hartwegii). 

Hartweg’s Iris is a locally common, endemic, medium-sized plant that grows on woodland slopes in the western foothills of the Sierra Nevada Mountains. It blooms from late spring to early summer. 

The flower of Hartweg's Iris is typically cream, yellow, or lavender with yellow or purple veins. It is sometimes called the "Rainbow Iris" due to these color variations. 

At first inspection, an iris appears to have three whorls of petals. The outermost and innermost whorls are not petals but modified sepals and styles, respectively. Three sepals (falls) comprise the outermost, largest, and often most colorful whorl.  The 2nd whorl contains three upright actual petals (banners).  

The 3rd whorl contains stamens. On the iris, stamens may be hidden. Three stamens are located above the three sepals and below the three styles. The anthers are large and long. The styles are modified and look like narrow, notched petals. The barely discernible tissue below the notch is the stigma. 

Irises have "inferior" ovaries. An iris's ovary is at the bulge located below, where the sepals, petals, and stamens separate.

The unique structure of irises aids in pollination. When a bee lands on the sepal, it is guided to the nectar through a tunnel created by the proximity of the sepal and style, as well as colorful lines known as nectar guides. Inside the flower, a bee deposits pollen from a previously visited flower onto the stigma. When the bee backs out of the flower, it picks up fresh pollen from the anther.

Although the sepals and the pistil are highly modified, an iris flower follows the four concentric whorls model. The central structure is the pistil, surrounded by a ring of three stamens. A ring of three petals surrounds the stamens, and sepals surround the petals.

The Western Blue Flag Iris (Iris missouriensis) is widely distributed throughout the Western United States. It has a similar flower structure to Hartweg’s Iris but is taller, up to two feet, and prefers wetter environments. The flower is typically purple, occasionally white, with distinct nectar guides on the sepals. Western Blue Flag Iris spreads into large populations in seasonally moist, grazed meadows as livestock avoid the plant. 

Not all members of the iris family, Iridaceae, have complicated flowers. During spring in the Sierra foothills, you may spot Western Blue-eyed Grass (Sisyrinchium bellum ) growing in wet meadows or along streams.  The flower of this plant looks distinctly different from an iris, yet it shares the characteristics of floral parts in multiples of three and an inferior ovary. Western Blue-eyed Grass has long, narrow leaves and branched stems. Tepals are wedge-shaped, becoming wider toward the tips.

In summer, a plant similar to Western Blue-eyed Grass, Idaho Blue-eyed Grass (Sisyrinchium idahoense), blooms in moist grassy areas. This species has more elliptic-shaped tepals that narrow toward the tips.

All flowers in the previous sections have radial symmetry. This occurs when flower structures radiate from the center like the spokes on a wheel. When some floral parts on a given flower have different sizes or shapes with like structures arranged on opposite sides of a single plane, bilateral symmetry results; flowers having bilateral symmetry can only be divided by a single line to produce mirror images on either side of the division.

The California Centaury (Zeltnera venusta) produces a flower with radial symmetry. The small flowers may be pink, magenta, or occasionally white with a white throat. The anthers are bright yellow and twist into a corkscrew shape after releasing pollen. The bright colors and twisted ribbon-like anthers give this plant a festive appearance.

The California Centaury is relatively common in the Sierra up to 6,000 feet. The plant grows from one-20 inches in grasslands and forest openings.  

In contrast, the flower symmetry of Hernandez Bluecurls (Trichostema rubisepalum) is distinctively bilateral. The pistil and stamens are long and arching. Purple petals with long white hairs are of three sizes. The narrowest petals flank the stamens. Broader petals are positioned beside the narrow petals, and between these is a single, often speckled, lip petal. 

Hernandez Bluecurls is a member of the mint family, Lamiaceae, growing below 4,500 feet in the western Sierra Nevada foothills and Southern Diablo Mountains. The plant is typically found in or along dried serpentine creekbeds of its limited range in Tuolumne, Mariposa, and San Benito Counties. The stem and leaves of Hernandez Bluecurls are covered with hairs and glands. When touched, the plant will emit a mild vinegar aroma.

Due to its specific environmental requirements and limited distribution, Hernandez Bluecurls are unlikely to be encountered. Although its cousin, Vinegarweed (Trichostema lanceolatum), is common and grows from the western Sierra foothills westward to the Pacific Ocean. 

Vinegarweed grows in dry, open locations and flowers from late summer to mid-autumn. The plant is appropriately named. From a distance, vinegarweed appears as an insignificant, green, weedy plant among dried amber grasses. At closer proximity, a robust acidic odor can be detected. Only at the closest inspection can the plant’s other-worldly beauty be revealed. 

Vinegarweed has the same physical traits as Hernandez Bluecurls, although greatly exaggerated. Flower structures, most notably the pistil and stamens, are longer, glands are more numerous, and the vinegar-like odor is more robust. The odor comes from volatile oils, which deter herbivores and may inhibit seed germination of competitive plant species. Native Americans prized Vinegarweed for its medicinal properties.

Some flower variations result from the fusion of floral structures. Fusion is the attachment of multiple flower parts, forming a single structure. For example, the plants addressed in the Flower Symmetry section have the bases of their petals and sepals fused to form tubes. Here, monkeyflowers of the lopseed family, Phrymaceae, are highlighted as representative plants with prominently fused sepals and petals.

Monkeyflowers are germane to considerations of California wildflowers. There are about 200 species of monkeyflowers worldwide, with more than 50% native to California, many exclusively in California. Monkeyflowers are highly diverse and live in a variety of habitats. The majority of monkeyflowers have showy blossoms, even if they are small. For some species, most of the above-ground plant is comprised of flowers. When such plants grow in mass, they produce a carpet of color. 

A monkeyflower blossom has five fused petals that form a corolla tube, with the petals becoming free at the tips to produce a bilateral, trumpet-like flower shape. The petals emerge from a whorl of fused sepals, collectively known as the calyx. Monkeyflowers have four stamens in two pairs and a single pistil. The stigma is two-lobed and will shut when stimulated. 

Monkeyflowers of the Sierra are in the genera Diplacus and Erythranthe.  The photo of the Cardinal Monkeyflower  

(Erythranthe cardinalis) shows a plant with an obvious flower stem that extends past the photo. In contrast, Kellogg’s monkeyflower (Diplacus kelloggii) is a plant with an inconspicuous flower stem. Monkeyflowers with a flower stem shorter than the calyx are placed in the genus Diplacus, whereas plants with a flower stem longer than the calyx are classified as Erythranthe.

Kellogg’s Monkeyflower grows from one inch to one foot tall on unstable slopes below 4,000 feet in the Sierra Nevada and Cascade ranges. Its spring blossom is reddish-purple or magenta with darker pigment towards the flower throat. Two lines on the lower petal, anthers, and the spotted corolla tube are yellow. Kellogg’s Monkeyflower has leaves arranged oppositely on the stem like all monkeyflowers.

A face has bilateral symmetry. An imaginary line running from the forehead to the chin and transversing over the nose will produce mirror images of the right and left sides of the face. It has been suggested that the blossoms of some monkeyflowers, such as Kellogg’s Monkeyflower or this Yellow-lipped Pansy Monkeyflower (Diplacus pulchellus), look like monkey faces. The corolla tube opening represents the mouth, the side petals resemble cheeks, and the lower petal is the chin or lower lip. 

The “face-like” Yellow-lip Pansy Monkeyflower grows exclusively in moist, open locations where the soil has been disturbed. The small plant is uncommon. Its range is centered in Calaveras, Tuolumne, and Mariposa counties. The flower throat is magenta, the upper petals are lavender to magenta, and the lower petal is yellow with magenta spots. The color of the side petals may match either the upper or lower petals.

Purple Mouse Ears, or Brownies (Diplacus douglasii) are small annual monkeyflowers that could be easily overlooked if not for their bright magenta petals. This plant received its common name from its diminutive size and two large pink upper petals resembling mouse ears. Purple mouse ears grow on slopes, often in serpentine soil, below 4,000 feet in the Sierra foothills, Cascade Mountains, and coastal regions.

Taking some imaginary license, you may think of purple mouse ears not as a mouse or monkey face but as a fire-breathing dragon. Under this scenario, the corolla tube represents the mouth, and the yellow anthers and lines inside the magenta corolla tube's throat are the dragon’s flames.

In sharp contrast, flowers from a plant of this same species viewed from the side could be imagined as a choir with its members looking upward and singing an angelic hymn.

Tricolor Monkeyflower (Diplacus tricolor) is a spring-blooming, annual plant most often found in seasonally wet depressions of the Central Valley. However, it may also be found in the lower western Sierra foothills, usually below 2,000 feet. The petals are purple, with central magenta markings on each petal. The lower petal ridges are yellow.

Bolander's Monkeyflower (Diplacus bolanderi) grows in open and disturbed locations in the central Sierra foothills and northern coastal mountains up to 5,500 feet. Plant size varies considerably from one to 35 inches in height. The plant is densely hairy. Flowers have a swollen calyx and pink petals with a white throat.  

Torrey's Monkeyflower (Diplacus torreyi) is endemic to and common in the Sierra Nevada Mountains. It grows in open areas up to 6,500 feet. The outer corolla tube is dark pink with light pink unfused tips flaring outwards. Two yellow ridge stripes bordered in magenta extend from the flower throat to the lower lip petal. Other markings are generally absent.

Some people detect a foul odor when the leaves or stems of the Skunky Monkeyflower (Diplacus mephiticus) are rubbed or crushed. This small plant, less than 6 inches tall, grows in sagebrush openings and on granite slopes of the Sierra Mountains and Great Valley. The flowers may be yellow or pink, with red lines and spots on the lower petals.

Layne's Monkeyflower (Diplacus layneae) is a California endemic species that grows on barren granite or serpentine soils in mountain regions up to 6,500 feet. The plants are hairy and one-11 inches tall. The flower has a calyx with dark ribs, pink petals with dark interior markings, and a white throat.

Granite-crack Monkeyflower (Diplacus graniticola) is similar to Layne's Monkeyflower.  Granite-crack Monkeyflower has a limited distribution extending from Hetch-Hetchy south to central Tulare County at elevations of 1,000-7,000 feet. It and Layne’s Monkeyflower sometimes grow in close proximity and bloom at similar times. Granite-crack Monkeyflower may have originated from the hybridization of Layne’s and Skunky Monkeyflowers.

Granite-crack Monkeyflower grows almost exclusively in granite crevices. Unlike Layne's Monkeyflower, it has a yellow throat and a compact habit. On Granite-crack Monkeyflower, the space between the leaves (internode) is less than the length of the leaves. In contrast, Lanye’s Monkeyflower has an open growth habit, with leaves spaced further apart on the stem.

Only six percent of monkeyflowers are small shrubs, yet they may be most familiar as they are frequently grown in native gardens and easy to spot on roadside slopes. Orange Bush or Sticky Monkeyflower (Diplacus aurantiacus grandiflorus) grows in the Sierra foothills and California coastal regions on slopes and in disturbed locations. Its leaves are sticky and curled with harry undersides. Orange Bush Monkeyflower is a stunning plant that produces many yellow-orange or pale-orange flowers.  

Rock Bush Monkeyflower (Diplacus calycinus) is a Southern Sierra species growing in foothills east and southeast of Fresno. As the name suggests, It grows in rocky crevices or well-drained locations below 4,500 feet. Like the Orange Bush Monkeyfower, Rock Bush Monkeyflower produces a profusion of flowers. Both bush monkeyflower species are available for sale at native plant nurseries.

At the opposite end of the monkeyflower size spectrum is Least-flowered Monkeyflower (Diplacus leptaleus). This plant can grow to 5.5 inches but may be much smaller and is easily overlooked. Least-flowered Monkeyflower grows in open locations with gravely granite soils above 6,500 feet. The tiny flowers may be pink, yellow, or white.

Brewer’s Monkeyflower (Erythranthe breweri) is another extremely small plant. Unlike the least-flowered Monkeyflower, it has notched petals and grows in moist environments. Due to their small sizes, both species require close inspection. The flower stem of Brewer’s Monkeyflower is longer than the calyx, so it is classified in the Erythranthe genus. All of the following monkeyflowers are stem-flowered monkeyflowers of the Erythranthe genus.

Some Erythranthe monkeyflowers are bicolored with three yellow lower petals, the center petal having red spots, and two upper petals of a contrasting color. Yellow and White or Bicolor Monkeyflower (Erythranthe bicolor) is one such species. This plant prefers moist clay soils, has linear leaves, and grows to 10.5 inches.

The bicolor variety of Yellow and White Monkeyflower is easy to identify from the two upper white petals. Flowers may also have all yellow petals, making identification more difficult. The all-yellow plants may be interspersed within a yellow and white population, or entire populations may have the all-yellow petaled plants. 

Montia-like Monkeyflower (Erythranthe montioides) is a central and southern Sierra bicolor species. The lower petals are yellow, and the upper petals are maroon, but all five petals may be yellow. The all-yellow variety resembles the all-yellow-petaled Yellow and White Monkeyflower. However, the Montia-like Monkeyflower is smaller, growing to only six inches, and prefers granite soils.

Primrose Monkeyflower (Erythranthe primuloides) is a common Sierra plant that grows in wet meadows, seeps, and streambanks from 2,000 to 11,000 feet. This small perennial spreads to form mats comprised of leaf rosettes; often, the leaves have long, upright hairs. The flowers are yellow, usually with red spots on the lower three petals.

Slender-stemmed Monkeyflower (Erythranthe filicaulis) is a rare and fairly threatened species.  Its distribution is limited to moist, loamy soils at 4,000-5,700 feet in Tuolumne and Mariposa Counties. The species is sometimes called the Hetch Hetchy Monkeyflower because it can be found in the general vicinity. The narrow-stemmed plant grows from 1.5-12 inches.  The flower is pink-purple with a dark purple throat and two yellow lines on the lower petal.

The Many-flowered Monkeyflower (Erythranthe floribunda) is common in the Sierra Nevada and Coastal Mountains. It grows below 8,000 feet in wet, open locations. The annual plant is densely hairy, erect, and branched. Many-flowered monkey flowers bloom in spring and summer, with yellow flowers having red markings. Variably toothed leaves up to an inch are broadest at the base and somewhat heart-shaped.

Musk Monkeyflower (Erythranthe moschata) could be confused with Many-flowered Monkeyflower. It has similar flowers, but the red markings are more lines than dots. The plant is a hairy perennial that spreads by rhizomes and has a musk-like odor. It grows in partial shade of wet environments. Musk Monkeyflower's one-half to two-inch toothed leaves are tapered at the blade base and tip.

California Blushing Monkeyflower (Erythranthe erubescens) is a large monkeyflower that grows up to three feet tall along stream banks, seeps, and wet depressions at elevations of 4,500-11,500 feet. The petals are hairy and light pink, with yellow lines and red spots on the lower petal ridges. A less common and similar-appearing sister species, Lewis’s Monkeyflower (Erythranthe lewisii), has deeper pink flowers.

The Cardinal or Scarlet Monkeyflower (Erythranthe cardinalis) is closely related to the California Blushing and Lewis’s Monkeyflowers. The tunnel-like yellow throats of the latter monkeyflowers are perfect for bee pollinators. In contrast, the backward-bending, red-orange petals, exposed stigma, and anthers of the Cardinal Monkeyflower are appropriate for hummingbirds. Pollinator preference for specific flower color and shape prevents closely related species from interbreeding.

Several plants within the Erythranthe genus have yellow flowers with the basic flower shape depicted in the photograph above. The flower is two-lipped, with the top two petals forming the upper lip and three lower petals forming the bottom lip. The lower petal is the largest, has two humped ridges, and forms a landing pad for pollinators. The landing pad is hairy and has variable red spots. Here, four species with this flower structure and occurring in the Sierra are compared.

Large Mountain Monkeyflower (Erythranthe tilingii) is a summer-blooming plant that grows above 4,500 feet along streams and in wet meadows. Plants may be solitary up to 14 inches tall or spread by rhizomes to form mats. Opposite leaves may or may not be toothed. The yellow corolla may be as large as 1.5 inches, with or without red dots on the lower lip. Flower stems originate independently at nodes and do not share a flower stalk.

Seep or Common Yellow Monkeyflower (Erythranthe guttata) is similar to Large Mountain Monkeyflower. Both plants are perennials with similar leaf and flower shapes. Seep Monkeyflower can be found at lower elevations, has a longer bloom time, grows taller, over two feet, and has smaller flowers, usually less than one inch. The most definitive distinguishing feature is that the Seep Monkeyflower has multiple flowers attached to a central stalk (raceme).

The Small-leaved Monkeyflower (Erythranthe microphylla) resembles a miniature version of the Seep Monkeyflower and grows in similar wet locations, such as moist meadows and seeps. Unlike the Seep Monkeyflower, the Small-leaved Monkeyflower is an annual and lacks rhizomes.

The Small-leaved Monkeyflower grows to about one foot tall and has flowers and leaves that are much smaller than those of the typical Seep Monkeyflower. However, it could be easily mistaken for a small specimen of the larger species.

Cut-leaved Monkeyflower (Erythranthe laciniata) is a small-flowered, thin-stemmed plant that can reach a height of up to 15 inches. It is easily identified by its distinctive lobed leaves on the lower stem. The range of Cut-leaved Monkeyflower is limited to the Sierra Nevada, where it grows almost exclusively in granite seeps above 3,000 feet.  

Flower modifications are not always dramatic, but they can still be puzzling. An example is the Marsh Grass of Parnassus (Parnassia palustris) in the Parnassiaceae family. This late summer bloomer is broadly distributed in North America and Eurasia. Its elongated, egg-shaped leaves are located at the base of the plant; the stem has a single bract and terminates with a single white flower.

Flowers of the Marsh Grass of Parnassus appear unremarkable from a distance, but closer inspection reveals five beautiful petals with intricate trough-like veins. The petals are creamy white; the veins appear gray or translucent. The flower also has five sepals, five pollen-producing stamens, and a central pistil with a four-lobed stigma. All normal, but then the flower becomes curious….

Between each typical-appearing stamen is a yellow glandular structure. Broad at their bases, these structures flare out with several slender stalks, each supporting a terminal gland. These unusual-appearing structures are in the same whorl as the stamens. They are themselves stamens modified to attract pollinators rather than produce pollen—the combination of these “staminodes “ and the distinctly veined petals results in a stunning flower.

Staminodes are also found in penstemons of the plantain family, Plantaginaceae. Penstemons have just one staminode per flower, much different from those on the Marsh Grass of Parnassus. As penstemons are common and bloom throughout the summer, one has more opportunities to observe staminodes in penstemons. California has over fifty native penstemon species subdivided into numerous varieties. They are well represented in the Sierra Nevada Mountains.

Penstemons are short-lived perennials: herbs, subshrubs, or shrubs that do well in dry, low-nutrient soils. Leaves of penstemons lack leaf stalks and come in various shapes with smooth or toothed margins. Most often, leaves are arranged oppositely on the stem, occasionally whorled, and primarily located at the base of the plant. 

The flowers may be grouped in whorled clusters or arranged along the stem. The petals are of various colors: blue, purple, pink, white, yellow, or crimson. They attract pollinators such as bees, butterflies, moths, and hummingbirds.

A penstemon flower has bilateral symmetry with five fused sepals and petals.  The flower has four fertile stamens having paired anthers, two on each side of the central pistil—the stamen’s filaments often arch within the floral tube like a rib cage. A single staminode, a sterile fifth stamen that lacks an anther, is present and responsible for the name penstemon, which roughly translates to “almost a stamen”; e.g., a peninsula is “almost an island.”

Depending on the species, staminodes can be densely hairy, sparsely hairy, or hairless. Staminodes with prominent hairs gave this plant group its common name, beardtongue. Imagine the staminode as a hairy tongue in the mouth of the penstemon flower. Sometimes, the anthers and lower lip petals are also hairy.

Staminodes are thought to be vestigial structures in some species. For others, especially those with hairy staminodes, it has been suggested that hairs may block out unwanted insects, act as a lever mechanism to shake pollen free from anthers, or temporarily trap pollinators to promote pollen transfer.

In some species, tiny glandular hairs are present on the outer petal. Similar hairs may also be present on stems, sepals, and leaves. Observe the photo above closely to see tiny hairs on most above-ground plant surfaces.

A common, easily distinguishable, and favorite penstemon is Mountain Pride (Penstemon newberryi). Mountain Pride is a small shrub growing to about a foot tall in rocky locations at mid-to-high elevations. The plant provides beautiful pink or magenta splashes of color contrasting the gray granite outcrops of mountain passes. The lower petals, staminode, and anthers have woolly hairs. Mountain pride is a summer bloomer with roundish, slightly toothed leaves.

Davidson’s Beardtongue (Penstemon davidsonii davidsonii) is a low-growing (less than 5 inches), mat-forming penstemon of the High Sierra. Pinkish or bluish-lavender flowers are hairy inside and rise above hairy stems, having small, thick, crowded, elliptic to roundish, evergreen leaves. This mid to late summer bloomer grows at 6,500-12,500 feet in open rocky locations.

Sierra Penstemon (Penstemon heterodoxus) is common at subalpine and alpine elevations of the Sierra. Clusters of small tubular flowers whorl around the stem. The exterior petal surfaces have tiny glandular hairs. The flower’s inside floor and staminode have brownish-yellow hairs. Corolla lobes are short and flair slightly. The upper two lobes flair the least. Buds are frequently deep purple. Leaves often fold lengthwise. At the base, they form a dense rosette.

Meadow Penstemon (Penstemon rydbergii oreocharis) looks similar to Sierra Penstemon but can be distinguished by the lack of glandular hairs on the outer petal surface. Meadow Penstemon may be found in meadows and along streambanks, often forming large populations. 

Showy or Royal Penstemon (Penstemon speciosus), as its common names suggest, is an impressive plant found in dry, rocky sites of subalpine and sagebrush ecosystems in the Western United States. Plants may grow to two feet tall at lower elevations with open growth. Plants are smaller at subalpine elevations. Leaves are narrow, green to green-gray, and often folded lengthwise.

The petal color of Showy Penstemon is blue, purple, or pink, with a lightly pigmented throat and reddish-purple lines (nectar guides) on the lower petals. Flowers are frequently densely arranged on one side of the stem. This species is a beautiful, robust choice for rock gardens.

Bridges’ or Beaked Penstemon (Penstemon rostriflorus) differs from the other species presented in both flower color and shape. The bright, downward-pointing, orange-red flowers attract hummingbird pollinators and, appropriately, have long tubular flowers that lack the landing pad typical of bee-pollinated flowers. Bridges Penstemon is a summer bloomer growing to three feet in sagebrush, woodland, and subalpine locations. In California, the plant is limited to central and southern regions.

Gaping Keckiella or Gaping, Yawing, or Bush Penstemon (Keckiella breviflora glabrisepala) was previously classified in the Penstemon genus and is often grouped with penstemons. Keckiella species are more vine or shrub-like than penstemons and, unlike penstemons, have hairs at the base of their filaments. Keckiella species possess other typical penstemon characteristics, including the sterile staminodes.

Gaping Keckiella is common in the Sierra Foothills and may grow as high as 9,000 feet. It is a shrub with numerous wondering flower stems reaching six feet under ideal conditions. Gaping Keckiella’s petals are hairy above, white, tinged with rose, and have purple-pink lines. The upper two petals are arching. The lower three deeply lobed petals retract down and back. Stems, leaves, and sepals are hairless. A similar variety Keckiella breviflora brevifloria has glandular hairs on the sepals. 

Some plants have flowers lacking one or more of the four primary structure whorls described earlier. Such flowers are known as incomplete. Ginger plants of the pipevine family, Aristolochiaceae, provide good examples. These flowers lack petals.

Growing on well-drained slopes in red fir and yellow pine forests of the Sierra Nevada Mountains, you may encounter Hartweg’s Wild Ginger  (Asarum hartwegii). This low-growing plant makes a beautiful ground cover with its variegated, heart-shaped leaves. 

The flower of Hartweg’s Wild Ginger is often overlooked as it grows close to the ground and may be blocked from view by its leaves. The flower has no petals. The flower shape is created by harry, maroon, partially fused sepals having long, pointed lobes, which visually blend into the litter layer. 

Hartweg’s Wild Ginger self-pollinates; hence, the plant does not need to waste resources to produce petals. Flies also sometimes lay eggs in the flowers and may inadvertently pollinate them during their visit. The flies are likely attracted to the flower’s musty odor and the sepals' deep red-brown (carcass-like or fungal-like) coloration. This cross-pollination by flies appears to play only a minor role, with self-pollination being more common.  

Wild ginger plant species employ ants for seed dispersal. The seeds of wild gingers have a highly nutritious lipid appendage known as an elaiosome. Ants harvest and carry the seeds to their nests to feed the elaiosome to ant larvae. The seeds are unharmed, dispersed, and planted in an ideal location for germination.  Over 11,000 plant species produce elaiosome-bearing seeds.

Lemmon’s Wild Ginger (Asarum lemmonii) is a California endemic species that grows in the same forests as Hartweg’s Wild Ginger.  The two species can be easily distinguished because Lemmon’s Wild Ginger has leaves with no variegation, has sepals with shorter lobes, and grows in wet locations. 

Note: these plants are not closely related to culinary ginger.  Native Americans used wild ginger for medicinal purposes. The plants are considered toxic.

California Pipevine or Duchman’s Pipe (Aristolochia californica), like its ginger relatives, lacks petals and has an unusual flower shape formed from green/brownish-red fused sepals. This climbing vine can be found at lower elevations, up to 2,300 feet, in the Northern Sierra Nevada foothills. California Pipevine is of particular ecological significance because it is the only plant on which the Pipevine Swallowtail Butterfly lays eggs.

Whereas ginger species lack petals, Fendler’s Meadow Rue (Thalictrum fendleri) of the buttercup family, Ranunculaceae, has flowers lacking two structural whorls: petals and either pistils or stamens.  A flower having pistils or stamens, but not both, is imperfect. The flower of Fendler’s Meadow Rue is considered both incomplete and imperfect.

Tassel-like male flowers consist of four sepals with 15 to 28 dangling stamens that move with the slightest breeze. The petals and pistils are absent. The female flower is less showy. It has four sepals and several pistils, with green, cream, or pink stigmas. Female flowers lack petals and stamens. Fendler’s Meadow Rue is wind-pollinated and does not need showy petals to attract animal pollinators.

Fendler’s Meadow Rue grows in shade or partial shade with widespread distribution in both mountain and coastal regions of California. Lobed, compound leaves grow in a mound from 1-2 feet high. Young leaves are in beautiful, tightly compacted clusters but then spread out as leaf stems (petioles) lengthen. Flower stalks rise above to a height of 4-5 feet.

Plants like Fendler’s Meadow Rue have separate male and female flowers; some other species have two distinct complete flower morphologies, having all four flower whorls. Foothill Jepsonia (Jepsonia heterandra) of the saxifrage family, Saxifragaceae, provides one example. Foothill Jepsonia grows from El Dorado County southward to Mariposa County. In winter/spring, each plant presents two to three attractive, round, slightly lobed, toothed leaves, which generally die back during the dry summer.

In autumn, Foothill Jepsonia plants with withered leaves utilize stored resources to produce flower stalks. Reddish, hairy stems support numerous flowers having five white petals with pink veins and five partially fused, pale, green-veined sepals. Each flower has ten stamens and a pistil, forked with two distinct stigmas and styles. About one-half of the plants will possess flowers with large stigmas elevated above anthers.

The other half of the Foothill Jepsonia population produces flowers with anthers rising above tiny stigmas. Foothill Jepsonia’s flowers are small. Therefore, careful observation is required to note the differences. This condition of a species having two distinct flower morphs with different style lengths relative to stamen lengths is known as heterostyly.

Heterostyly promotes cross-pollination, creating greater genetic variation in offspring.  In theory, a pollinator visiting a flower with a short stamen filament will transfer more pollen if the subsequent flower visited has a short style. Similarly, a pollinator picking up pollen from a flower with a long stamen filament will more efficiently transfer pollen to flowers with long styles. Thus, pollination is most successful between plants with reciprocal anther and stigma heights. 

Making sense of a flower’s morphology may be complicated by structures not part of the flower’s four concentric whorls. This is true of Castilleja species of the broomrape family, Orobancaceae, including paintbrushes and owl's clovers. These plants have colorful leaves called bracts, which serve to attract pollinators. Think of poinsettias. The prized red color of this favorite Christmas plant comes not from petals but from the red leaves surrounding small yellow flowers.

Paintbrush plants have colorful bracts located below and surrounding the flowers. The paintbrush flowers are tubular, inconspicuous, and overshadowed by the showy bracts. Pictured here are two examples of paintbrush flowers and bracts.

California has approximately forty-five Castilleja species divided into dozens of additional varieties. The secret to Castillejas’ success is partly their hemiparasitic means of obtaining nutrition. Castilleja plants are capable of photosynthesis, but their roots also penetrate the roots of nearby plants to steal nutrients. Although this parasitism harms host plants, it may also shape and benefit community diversity by keeping dominant host plant populations in check. 

Wyoming Paintbrush (Castilleja linariifolia) is widespread throughout the western United States. It grows up to three feet in dry, rocky locations such as sagebrush scrub. In most locations, the bracts are reddish-orange. However, on the eastern slope of the Sierra Nevada Mountains, the plant has a unique reddish-pink color.

Great Red Indian Paintbrush (Castilleja miniata miniata) grows up to 2 1/2 feet tall in moist or wet locations above 5,000 feet. The bracts of Giant Red Indian Paintbrush are red to orange. The lower leaves are lance-shaped, broader at the base, and tapering to a pointed tip. Only the red bract leaves have lobes. Habitat and the lower leaves lacking lobes can help distinguish this plant from similar species.

Wavy Leaf Paintbrush (Castilleja applegatei), widely distributed in California’s mountain regions, is highly variable and divided into four varieties. It is a summer bloomer found in dry locations. An identifying feature of the Wavy Leaf Paintbrush is its leaves with prominent curving margins.

Lemon’s Paintbrush (Castilleja subinclusa lemmonii) is readily identifiable by its distinctively pink bracts. Its stems are usually unbranched and grow up to eight inches in height. Lemon’s Paintbrush is a mid to late summer bloomer growing above 5,000 feet, most commonly in moist sub-alpine and alpine meadows.

Parrothead Indian Paintbrush (Castilleja pilosa) is a summer bloomer that grows to 14 inches tall in sagebrush scrub and alpine locations. Its bracts may be pale green or pinkish purple with white margins. Parrothead Indian Paintbrush is similar to Dwarf Alpine Paintbrush. A distinguishing difference is that the tips of Parrothead Indian Paintbrush bracts are rounded, and Dwarf Alpine Paintbrush bracts taper to a point.

Owl’s clovers also belong to the genus Castilleja. Plants referred to as paintbrushes frequently have long flowers resembling paintbrushes' tips, whereas Owl's clovers have stout flowers, some of which may bring to mind a perched owl. Flowers have five petals. Three lower fused petals form a pouch, and two fused upper petals form a beak. The dense, bicolored lobed bacts of an owl’s clover can resemble the clustered flowers of some clover species.

Purple Owl's Clover (Castilleja exserta) is a common spring bloomer with wide distribution. It grows up to 18 inches in open, grassy areas. Bracts are often purple but may also be pink, yellow, or white. Several owl’s clovers are highly variable in appearance and resemble one another. A hooked flower beak with shaggy hair identifies Purple Owl’s Clover.

Dense Flower Owl's Clover (Castilleja densiflora) is similar to Purple Owl’s Clover. This less common species has a straight flower beak with sparse hairs. Note the flower differences in the insert photos of the previous two images.

Thin Lobed Owl's Clover, Pale Owl's Clover, or Valley Tassels (Castilleja lineariloba) is an endemic plant of the Western Sierra foothills. Bracts are typically green with white tips but may also be pinkish. Stems have soft glandular hairs. 

Steps to follow when encountering a plant with an unfamiliar or unusual flower: 1. utilize the model of concentric whorls (Calochortus lilies); 2. be mindful that sepals may look like petals (tepal lilies); 

Flowers of yellow Pretty Face (Triteleia sp.) and pink Sierra Onion (Allium campanulatum) have tepals. 

3. recognize that structures may be significantly modified (iris’, Marsh Grass of Parnassus, and penstemons); 

The unique flowers of Brewer’s Miterwort (Pectiantia breweri) result simply from petals reduced to thin branched structures. 

4. anticipate bilateral symmetry and fused structures (bluecurls, monkeyflowers, and penstemons); 

Monkshood’s (Aconitum columbianum) unusual bilateral flowers have sepals, which differ in size and shape and are pigmented like petals. The upper sepal forms the hood.

5. understand that structural whorls may be absent (gingers); 6. know that a single species may have different types of flowers (Fendler’s Meadow Rue and Foothill Jepsonia) and 7. be aware that bracts may look like petals (paintbrushes and owl’s clover).

Warrior's Plume, or  Indian Warrior (Pedicularis densiflora) employs colorful bracts and red flowers to attract pollinators.