Cryptantha Index
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Boraginaceae Images On Wayne's Word

The borage family (Boraginaceae) is widely distributed in temperate and tropical regions, with about 134 genera and 2650 species. It includes many wildflowers in the Colorado and Mojave deserts of North America, typically with connate corollas in scorpioid (coiled) inflorescences, and with stems and leaves covered in glistening hairs (trichomes). In tropical regions there are hardwood trees with beautiful, showy flowers. The family is well represented in San Diego County, both in native annuals and cultivated perennials. Two of the most common genera are the so-called "popcorn flowers" Plagiobothrys and Cryptantha. Dna evidence indicates that root parasites of the genus Pholisma (Lennoaceae) and members of the Hydrophyllaceae (Phacelia, Nemophila, Nama, etc.) are closely related to the Boraginaceae. In fact, in the revised Jepson Manual II: Vascular Plants of California (2012), all these species are placed in the Boraginaceae as of 2012; however, in the online Jepson eFlora (accessed April 2024) the latter genera are back in their previous families Lennoaceae & Hydrophyllaceae. Note: Many genera additions to the Boraginaceae based on DNA cladistics in the 2012 Jepson Flora are still included in World Flora Online (WFO).
   Tom Chester Anza-Borrego Cryptantha Key     Mike Simpson Cryptantha Home Page     Wayne's Word Cryptantha Image List 

2. Two Remarkable Native Sand Plants In Boraginaceae
In Jepson eFlora They Are Back in the Lennoaceae!

1. Native Wildflowers In The Boraginaceae

Species of Cryptantha are often referred to as white "forget-me-nots," although some people call them "popcorn flowers." I think the latter common name is best applied to members of the closely related genus Plagiobothrys. Both genera have white flowers in tight coiled clusters called scorpioid cymes. In my humble opinion, the flower clusters of Plagiobothrys look a little more like popped grains of corn, technically referred to as exploded endosperm. This is particularly true of spreading, low-growing varieties of P. collinus, appropriately named California popcorn flower. From a distance, the white flower clusters do resemble popcorn scattered on grassland and open ground in coastal sage scrub. In spite of what you might have heard, I must say that the popcorn analogy is not a good way to separate white forget-me-nots from popcorn flowers.

Left: The colorful blister beetle (Lytta magister) feeding on a desert Cryptantha (probably C. angustifolia).

There is a slight resemblance between the tight, coiled flower clusters of Cryptantha intermedia and Orville Redenbacher's Popcorn; however, there is a big difference in taste! Actually, you should not eat the flowers. The irritating hairs (trichomes) could cause an unpleasant reaction.

Plagiobothrys collinus var. californicus. Members of the genus Plagiobothrys are the true popcorn flowers. In my opinion, the flower clusters with distinct yellow centers (throats) resemble popcorn more than the flowers of Cryptantha species, particularly from a distance. Note: The above species may be the very similar P. collinus var. fulvescens.

Popcorn flowers (Plagiobothrys collinus) compared with a popped grain of corn (inset).

To accurately separate the genus Cryptantha from Plagiobothrys you must carefully examine the nutlets to see if they have a keel (ridge) above the elevated attachment scar or a distinct groove above the recessed attachment scar. One to four nutlets are enclosed within five calyx lobes covered by glistening hairs (trichomes). Depending on the species, the nutlets range in size from 0.5 to several millimeters. This requires a good 10x or 20x hand lens or a dissecting microscope.

Note: Some of the extreme macro images with black backgrounds were taken on a black range top with a
Nikon D-90 SLR and a Phoenix RF46N Ring Flash using a 60mm Micro Nikkor AF-S F/2.8G ED Macro Lens.

Comparison of the nutlets of California popcorn flower (formerly Plagiobothrys californicus var. fulvescens) with the common "white forget-me-not" (Cryptantha intermedia). Depending on the photo, I use grains of ordinary table salt for a size relationship in Wayne's Word ultra-macro images.

Nutlet of Plagiobothrys compared with Cryptantha. Plagiobothrys has a raised (elevated) attachment scar at the base of a ridge (keel). Cryptantha has a recessed attachment scar at the base of a narrow groove. The bottom line here is: If your nutlet has a distinct groove it is undoubtedly a Cryptantha.

Alkannin: A Naphthoquinone from species in the Boraginaceae, incl. Plagiobothrys & Alkanna.

I once told my students that cryptanthas are more densely covered in hairs than Plagiobothrys, but that statement doesn't necessarily hold true. Stems and leaves of Plagiobothrys nothovulvus and our local P. arizonicus contain a reddish dye that rubs off on your fingers; however, our common desert annual Cryptantha micrantha (purple-rooted forget-me-not) also gives off this reddish dye. Several members of the Boraginaceae are known for this deep red phenolic dye called alkannin (also spelled alkanet), including dyer's bugloss (Alkanna tinctoria). It was used to dye textiles, and as a coloring for vegetable oils, medicines and wine, and is commonly used today as a food coloring. In addition, it reportedly has wound healing and anti-inflammatory properties. The term "bugloss" applies to several species in the Boraginaceae with blue flowers and hairy foliage. It is derived from a Greek word meaning ox-tongue in reference to the rough, hairy leaves.

Specimens of Plagiobothrys nothovulvus and Cryptantha micrantha from the Palomar College Herbarium. Note the reddish alkannin stain around the roots of both specimens. Although the plants were collected nearly forty years ago, the dye is still visible on the herbarium paper.

Red dye on fingers after fondling the basal stem & leaves of Arizona Popcorn flower.

There are other annuals in the Boraginaceae, such as Pectocarya and Harpagonella with minute white flowers in scorpioid cymes; however, they have very distinctive fruits (nutlets) that are much different from Cryptantha and Plagiobothrys. White or blue-flowered "forget-me-nots" (Myosotis) resemble Cryptantha, but they are not native to San Diego County.

The "stickseeds" (Hackelia and Lappula) also occur outside San Diego county. They have nutlets with barbed prickles, thus making them very effective hitchhikers. In fact, I gave Hackelia floribunda a rating of 4 Sock Removal Difficulty Units (SRDUs) on my Wayne's Word Top 17 Hitchhiking Plants. I once camped on the western slope of the Panamint Range, Inyo County on a very cold winter night. I was finally forced to climb out of my camper shell at midnight and put on long underwear. Unfortunately, I stood in a patch of stickseed (probably Lappula redowskii) during this clothing transfer resulting in one of the most miserable nights I ever spent in my entire life. Ironically, I was on a quest for another serious hitchhiker, the pink-flowered Arizona devil's claw (Proboscidea parviflora), reportedly introduced high in the Panamint Range in the late 1800s by the legendary Shoshone named Hungry Bill.

As I stated above, to identify a species of Cryptantha you must observe the nutlets. They can still be green, but they must be mature enough to show minute outer bumps called tubercles. Assuming you have eliminated all other genera in the borage family, here are some of the questions presented to you in taxonomic keys in Munz and the Jepson Manual: Do you have 1, 2 or 4 nutlets? Are your nutlets roughened by minute tubercles or are they smooth and shiny? Are all of your nutlets the same (homomorphic) or do you have dissimilar nutlets (heteromorphic)? Are the margins of your nutlets winged of knifelike with thin edge, or are they rounded? Is the shape of your nutlets triangular-ovate or oblong-lanceolate? There are other questions about the style length related to nutlets, size of flowers (diameter of corolla limb) and details of the glistening hairs (trichomes) on the calyx. For example, if you spot a Cryptantha in Coyote Creek with a large calyx 10 mm in length, long trichomes with bulbous bases, and one shiny, smooth nutlet it is undoubtedly the rare and endangered C. ganderi, named after the famous San Diego naturalist Frank Gander. Another similar-appearing species in Coyote Creek C. barbigera typically has four tubercled nutlets. Mike Simpson's Cryptantha page at SDSU has marvelous images of all 22 species native to San Diego County. Tom Chester has prepared on nice key to Cryptanthas of Anza-Borrego arranged in an easy to follow table. I have included the following images to show the variation in flower size and nutlets of several species. For more species, see the table at the bottom of this page.

Gander's cryptantha (Cryptantha ganderi), one of the many annual species of Cryptantha in San Diego County. Like other species of Cryptantha, the foliage and inflorecence are covered with bristly trichomes. This species typically has only one nutlet (rather than four), and the nutlet is smooth and shiny (rather than tubercled or papillate). The single nutlet is shown in the inset image (white arrow). The bristly sepals surrounding the nutlet are up to 10 mm long, longer than most other species in San Diego County. The long, conspicuous trichomes have bulbous bases. C. ganderi has an R-E-D code of 3-3-2 on the CNPS Rare & Endangered List 1B. It is endemic to sandy riverbeds of Borrego Valley and Anza-Borrego State Park.

Cryptantha intermedia showing roughened (tuberculate) nutlets.

Cryptantha clevelandii showing smooth and shiny nutlets.

Small-flowered form of Cryptantha micrantha. This is certainly one of the smallest desert wildflowers, rivaled in minuteness by certain flowers of the everlasting tribe (Inuleae) of the Asteraceae (e.g. Stylocline) and unisexual flowers of Chamaesyce (Euphorbiaceae). The U.S. penny (one cent) is 19 mm in diameter. Photographed with Nikon D-90 and 60 mm Micro-Nikkor AF-S F 2.8G ED Macro Lens with Phoenix Ring Flash.

Tubercled nutlets of three species of Cryptantha: A. C. intermedia, B. C. muricata and C. C. micromeres. The cuboidal grain of table salt (NaCl) is about 0.3 mm on a side.

Cryptantha micromeres from the southern Merriam Mountains. The nutlets are only 0.5 mm long, slightly larger than an average grain of table salt (NaCl) that is 0.3 mm on a side. The spreading corolla limb is approximately 1.0 mm in diameter. By way of comparison, the world's largest flower (Rafflesia arnoldii) of the Malay Archipelago is 3,000 mm in diameter! This is the smallest Cryptantha flower and one of the smallest "complete" flowers in San Diego County. Our local desert species (C. micrantha) is almost as small. The flowers of our closely-related Pectocarya species are also minute. Complete flowers have all 4 major floral parts, including calyx, corolla, androecium (stamens) and gynoecium (pistil). There are smaller flowers, such as those in the euphorbia and arum (duckweed) families, but they are incomplete and imperfect (unisexual).

Cryptantha Images Listed By Species

  1. Cryptantha angustifolia: Most Common Sp. In Anza Borrego

  2. Cryptantha barbigera in Smoke Tree Wash
    Cryptantha barbigera in Coyote Creek

  3. Cryptantha clevelandii in Southern Merriam Mountains

  4. Cryptantha decipiens in Moonlight Canyon Wash

  5. Cryptantha ganderi in Coyote Creek
    Cryptantha ganderi in Coyote Creek

  6. Cryptantha intermedia in Southern Merriam Mountains
    Cryptantha intermedia in Southern Merriam Mountains

  7. Cryptantha maritima in Domelands of Imperial County

  8. Cryptantha micrantha in Coyote Creek

  9. Cryptantha micromeres in Southern Merriam Mountains

  10. Cryptantha muricata in Southern Merriam Mountains

  11. Cryptantha nevadensis in Coyote Creek

  12. Cryptantha oxygona Near Gorman (L.A. County)

  13. Cryptantha pterocarya (cycloptera?) in Smoke Tree Wash
    Cryptantha pterocarya cycloptera in Moonlight Canyon Wash

  14. Cryptantha utahensis in Moonlight Canyon Wash
    Cryptantha utahensis in Coyote Creek (Boulder Alley)

Plagiobothrys: A Cryptantha Look-Alike

  1. Plagiobothrys arizonicus (Popcorn Flower) at Scissors Crossing   

Yellow Fiddleneack (Amsinckia) In Local Coastal Sage Scrub

A prolific southern California annual wildflower that competes with weedy grasses and forbs in vacant fields and disturbed areas.

Like the genus Cryptantha, the ovary composed of four one-seeded nutlets. The simple (unbranched style) arises from the gynobase rather than from the apex of the ovary. It is appropriately called a gynobasic style. The calyx is covered by glistening, sharp-pointed, bristly hairs (trichomes).

The Comb Burs (Pectocarya) and Grappling Hook (Harpagonella)

Comb burs (Pectocarya linearis ssp. ferocula) have very minute white flowers like the smallest cryptanthas; however, their nutlets are very different.

The comb burs (Pectocarya) are common in the Anza-Borrego Desert. Like Cryptantha and Plagiobothrys, they have minute white flowers; however, the nutlets are very different. The four spreading (divergent) nutlets have the shape of a tiny X. Each nutlet is fringed with minute prickles (hairs) along the margins. If you spot a small annual in a sandy wash with tiny X-shaped structures along its stem, it is most certainly a Pectocarya. In order to identify the species, you must have a mature nutlet. The coastal gabbro endemic called grappling hook (Harpagonella palmeri) has a very unique fruit with barbed projections radiating out in all directions. It has a rating of 10 Sock Removal Difficulty Units (SRDUs), surpassed only by the ultimate hitchhiker from Madagascar (Uncarina grandidieri). The latter species belongs to the sesame family (Pedaliaceae) along with our devil's claws (Proboscidea).

Pectocarya linearis ssp. ferocula, a coastal comb bur that also grows in the Anza-Borrego desert. The barbed nutlets make effective little hitchhikers.

The fruits of Harpagonella palmeri resemble miniature grappling hooks. The numerous barbed projections make them very effective hitchhikers. In fact, they have a SRDU rating of 10 (sock removal difficulty units), the second highest rating on the official Wayne's Word Top 17 Hitchhiking Plants.

Phacelia: One of the Largest Genera of Flowering Plants in California
With Well Over 100 species In The State (120 Species In Jepson eFlora).

As I have stated previously on this page, whether this wildflower genus is placed in the families Boraginaceae or Hydrophyllaceae depends on which flora you are using. Jepson eFlora places it in the Hydrophyllaceae with many other genera of wildflowers & shrubs in California. World Flora Online places it in the Boraginaceae. Obviously both families are closely related according to DNA cladograms (phylogenetic trees). In fact, some references consider the Hydrophyllaceae a subfamily Hydrophylloidiae within the Boraginaceae. Therefore, many species in the Palomar College Arboretum & local hills of coastal sage scrub could fall into the latter subfamily, including Emmenanthe, Eriodictyon, Eucrypta, Nemophila, Phacelia, Pholistoma, and Wigandia.

2. Remarkable Parasitic Sand Plants Moved To Boraginaceae in 2012 Jepson;
In The Latest Online Jepson eFlora They Are Now Back In The Lennoaceae!

Additional members of the traditional Boraginaceae in the Anza-Borrego Desert region of San Diego County include yellow fiddlenecks (Amsinckia), the coldenias (Tiquilia formerly placed in the Ehretiaceae) and Heliotropium (formerly placed in the Heliotropaceae). In the new Jepson Manual II (2012), the amazing root parasites (Pholisma) and the numerous wildflowers in the phacelia family (Hydrophyllaceae) are included in the Boraginaceae, making this one of largest and most diverse plant families in southern California; however, this is all changed again in the online Jepson eFlora (accessed April 2024). I am still including the root parasites Pholisma on this page because they are so truly fascinating! Note: Many genera additions to the Boraginaceae based on DNA cladistics in the 2012 Jepson Flora are still included in World Flora Online (WFO).

One of the most interesting of all dune plants, and certainly one of the most bizarre wildflowers in North America is "sand food" (Pholisma sonorae). This amazing parasitic flowering plant grows in the Algodones Dunes of southeastern California and adjacent Arizona, and in the sand dunes of El Gran Desierto in Sonora, Mexico (north of Bahia Adair in the Gulf of California). Within this area, the plants grow on sand dunes produced by wind transport of sand from the beaches of ancient Lake Cahuilla and the Colorado River delta. Another unusual species of sand food (Pholisma culiacana) is endemic to rocky, subtropical thorn scrub 500 miles (800 km) south in Sinaloa, Mexico. The disjunct distribution of these two species may be explained by plate tectonics. Floras west of the San Andreas fault and Gulf rift, in southern California and Baja California, have been displaced northward at least 500 km since the Miocene epoch (about 30 million years ago). Assuming that P. culiacana is more like the tropical progenitor of both species, P. sonorae could have diverged into a more xeric (drought resistant) species west of the San Andreas fault as peninsular California moved northward. Present day populations of P. sonorae east of the fault could have migrated there by shifting sand dunes in the region.

Sand plant (Pholisma arenarium), a close relative of sand food (P. sonorae) is occasionally seen in sandy areas of the Colorado Desert region. Like sand food, it is parasitic on the roots of nearby shrubs. Unlike sand food, it typically prefers sandy riverbeds and washes rather than sand dunes.

3. Cultivated Boraginaceae Species From Other Countries

Other Species In The Borage Family (Boraginaceae)
Including Cultivated Species From Other Countries

In addition, there are tropical trees in this family with beautiful, dark-grained wood. In fact, flowering trees of the genus Cordia are commonly seen in the Hawaiian Islands, southern Florida and the Caribbean. Although the name is pronounced the same, the type genus Borago is not derived from the Spanish word for bighorn sheep (borrego). Borago officinalis is an ancient European herb possibly originating in Syria and naturalized in San Diego County. It can be seen at the Herb Garden at the San Diego Wild Animal Park and San Diego Botanical Garden in Encinitas. The fresh leaves and blue flowers are eaten in salads and a nutritious oil is obtained from the seeds. Soothing medicinal teas are also made from the flowers of Borago and the closely related Echium in Iran, Pakistan and India.

Ziricote (Cordia sebestena) on Ambergris Caye off the coast of Belize. The beautiful hardwood is used for carvings.

Ziricote (Cordia sebestena), an evergreen, hardwood tree native to the Florida Keys and Caribbean region. It belongs to the large Borage Family (Boraginaceae), a family that includes mostly herbaceous wildflowers like the forget-me-nots (Myosotis and Cryptantha). The orange-red, crinkled petals are fused into a funnel-shaped corolla that tapers into a slender tube (see detached corolla on leaf). Another species called "kou" (C. subcordata) is native the Polynesian region. The beautiful wood and orange blossoms are similar to the Caribbean species; however, the leaves are not as stiff and rough as in C. sebestena.

Pride of Madeira (Echium fastuosum = E. candicans), another striking shrub native to the Canary Islands. The borage family (Boraginaceae) includes many species of wildflowers in California, including white forget-me-nots (Cryptantha and Plagiobothrys) and yellow fiddlenecks (Amsinckia). Some of these occur in the Arboretum and adjacent coastal sage scrub after spring rains. The family also includes a beautiful hardwood tree that is native to the Caribbean Islands (Cordia).

"Tower of Jewels" (Echium wildpretii)) native to subalpine volcanic slopes of the Canary Islands, a truly magnificent member of the Boraginaceae. The silvery rosettes of leaves & flower stalk are reminescent of the silver sword (Argyroxiphium sandwicense ssp. macrocephalum), a member of enormous sunflower family (Asteraceae) near 10,000 ft. summit of Haleakala on Hawaiian Island of Maui. .

Like other genera in the Boraginaceae, the ovary of Echium is composed of 3-4 one-seeded nutlets. The genus is derived from the Latin "echis" or viper, from resemblance of nutlets to the head of a viper.

Dense, glistening trichomes adorn the inflorescence of Wigandia urens in the Palomar College Arboretum. The flowers are produced in coiled (scorpioid) cymes typical of the waterleaf & borage families; however, cladistical DNA evidence suggests this species is best placed in the subfamily Hydrophylloideae within the Boraginaceae.

Like our California phacelias and turriculas, the stiff hairs (trichomes) of wigandias can be irritating to sensitive skin. It is best not to handle wigandia plants without gloves. In fact, they are sometimes called "mala mujer" in Mexico; however, they are not nearly as painful or "bad" as the "mala mujeres" of the euphorbia family (Euphorbiaceae), including species of Cnidoscolus. Like prickly pear and cholla cacti, the general rule of "look but don't touch" also applies to the hairy stems of wigandia. As long as wigandias are pruned back and viewed from a distance, they pose no serious threat to people. They are truly remarkable shrubs with foliage reminiscent of the rain-soaked New World tropics.

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