Plants that are restricted to one or more specific localities within a region are termed endemic. For example, the Torrey pine is native as well as endemic to California. Although commonly cultivated, it only grows wild on the sandstone bluffs between Del Mar and La Jolla in San Diego County, and on Santa Rosa Island. Because of the many isolated mountain ranges and valleys in California, approximately 22 percent of the flora is endemic to the state. Some endemic plants in California include the California cypresses (Hesperocyparis), Santa Cruz Island ironwood (Lyonothamnus floribundus ssp. asplenifolius), Death valley sage (Salvia funerea), Death Valley locoweed (Astragalus funereus), Panamint daisy (Enceliopsis covillei), and Death Valley gilmania (Gilmania luteola). Death Valley is especially rich in endemics because of its isolation from other regions. It is surrounded by high mountain ranges and hundreds of miles of arid desert. In fact, one of the rarest endemic wildflowers in California is called "rock lady" (Maurandya petrophila syn. Holmgrenanthe petrophila), a member of the plantain family (Plantaginaceae). This unusual plant grows in crevices on vertical limestone walls in steep canyons of the Grapevine Mountains bordering Death Valley.

Naturalized: True naturalized plant species are introduced from another region (outside California). Because they are able to grow wild and reseed themselves, they are considered part of the California flora. Some naturalized species are considered very destructive to the natural environment because they are extremely invasive and may replace the native species. This is especially serious when the loss of native plants threatens the survival of native animals that depend on the plants for food. Naturalized plants that are considered undesirable are often referred to as weeds. It has been estimated that more than 16 percent of the California flora is composed of naturalized species. Naturalized "weeds" often have ingenious methods of dispersal, including hitchhiking on animals and airborne seeds. A single tumbleweed plant (Salsola tragus) may produce 20,000 to 50,000 seeds within numerous small fruits, each surrounded by a circular, papery border. Mature plants readily break off at the ground level and are pushed along by strong gusts of wind. As they roll along hillsides and valleys, the seeds are scattered across the landscape. The introduction of Saharan mustard (Brassica tournefortii) into the Anza-Borrego desert region is an ecological disaster. The South African shrub called boneseed (Chysanthemoides monilifera ssp. monilifera) was introduced into the Palomar College Arboretum in the 1970s and has become locally naturalized. It is listed for California in the Flora of North America but not in the Jepson Manual.

Johnson Canyon & Panamint Range along the western border of Death Valley. I parked at canyon entrance above alluvial fan for one night. The following day I hiked to the snow-covered ridge to find this plant, an ascent of about 2,500 feet.

Serpentinite is a metamorphic rock formed on the sea floor at tectonic plate boundaries by the process of serpentinization. During this process, most chemical reactions essential to basic biochemical pathways of life take place. Hence, serpentine thermal vents are candidate (hypothesis) where life on Earth originated. Biological chemicals essential for metabolism of life have been discovered in these thermal vents, including Acetyl-CoA. There are currently (2024) two other solar system bodies where there is evidence for serpentinization: Mars and the Saturn moon Enceladus.

Twenty (18 depending on latest reference) of the more than 100 species of Pinus on earth. All of these pines are native to the state of California. 1. Monterey Pine (P. radiata), 2. Bishop Pine (P. muricata), 3. Santa Cruz Island Pine (P. remorata), 4. Whitebark Pine (P. albicaulis), 5. Limber Pine (P. flexilis), 6. Beach Pine (P. contorta), 7. Lodgepole Pine (P. murrayana), 8. Western White Pine (P. monticola), 9. Knobcone Pine (P. attenuata), 10. Bristlecone Pine (P. longaeva), 11. Foxtail Pine (P. balfouriana), 12. Four-Leaf Pinyon (P. quadrifolia), 13. Two-Leaf Pinyon (P. edulis), 14. One-Leaf Pinyon (P. monophylla), 15. Ponderosa Pine (P. ponderosa), 16. Coulter Pine (P. coulteri), 17. Digger Pine (P. sabiniana), 18. Torrey Pine (P. torreyana), 19. Jeffrey Pine (P. jeffreyi), 20. Sugar Pine (P. lambertiana). Note: This image is scanned from a Kodachrome 35mm color transparency taken in 1974. Taxonomic changes have been made on some of these species. Note: In the Jepson Flora of California 2nd Edition (2012), Pinus remorata is now considered a synonym of P. muricata. Another species (left image) called the Washoe pine (P. washoensis), with cones similar to a miniature Jeffrey pine, is now recognized as a variety of the ponderosa pine (P. ponderosa). In addition, the beach and lodgepole pines are now recognized as subspecies of P. contorta, rather than separate species. This gives a grand total of 18 species, 7 subspecies and 3 varieties.

The Angiosperm Phylogeny Group (APG) has proposed some significant changes in the classification of many traditional angiosperm families, including the placement of all duckweeds in the Araceae rather than the Lemnaceae. Nomenclatural changes are cited under the APG II system (2003) and superceeded by APG III system (2009). These changes are based on computer-generated evolutionary trees or cladograms. Thousands of data characters have been used, including morphology, anatomy, flavonoids, allozymes, and DNA sequences from chloroplast genes and introns. The Jepson Manual Second Edition essentially follows the changes summarized in the following reference by W.T. Judd, et al. 2008. The incorporation of these changes into textbooks, floras, checklists and herbarium collections is a formidable task, not to mention relearning hundreds of new names.

Judd, W.S., Campbell, C.S., Kellogg, E.A., Stevens, P.F., and M.J. Donaghue. 2008. Plant Systematics: A Phylogenetic Approach (Third Edition). Sinauer Associates, Inc., Sunderland, Massachusetts. 611 p.   Angiosperm Phylogeny Group (2003). "An Update of the Angiosperm Phylogeny Group Classification for the Orders and Families of Flowering Plants: APG II." Botanical Journal of the Linnean Society 141 (4): 399-436.   Angiosperm Phylogeny Group (2009). "An Update of the Angiosperm Phylogeny Group Classification for the Orders and Families of Flowering Plants: APG III." Botanical Journal of the Linnean Society 161 (2): 105-121.

Many traditional phylogenetic groupings of species within families and genera are not monophyletic and are inconsistent with modern cladistical analyses based on DNA. In other words, the groupings are paraphyletic or polyphyletic, and do not show all species within a group descending from a common ancestor. Monophyly is the natural evolutionary pattern in which all species are descended from a common ancestor. In order to have consistent computer-generated, monophyletic cladograms, it is sometimes necessary to change paraphyletic and polyphyletic groupings by moving species into different genera, and by moving genera into different families. Many of the taxonomic revisions in the Jepson Manual 2nd Edition (2012) were made in order to have consistent monophyletic groupings. This is why Spirodela punctata was placed in the genus Landoltia and why the Lemnaceae was placed in the familiy Araceae.

The cladogram (left) is from D.H. Les and D.J. Crawford (1999). It has high boot strap values and is based on molecular (rbcL) data from chloroplast DNA. It clearly shows that a grouping composed of 3 species of Spirodela is paraphyletic. This is why S. punctata was placed in the monotypic genus Landoltia.

In 1999, D.H. Les and D.J. Crawford proposed the new genus Landoltia containing one species L. punctata, formerly Spirodela punctata. This species is morphologically intermediate between Lemna and Spirodela. According to Les & Crawford, it represents an isolated clade distinct from both Lemna and Spirodela. Without this change, the genus Spirodela would be paraphyletc.

According to Professor Dr. Elias Landolt (personal communication, 2001), the creation of the new genus Landoltia is not necessary based on a purely morphological point of view; however, based on DNA and enzymatic studies, the change is warranted in order to form phylogenetically consistent taxa. The inclusion of a fifth genus Landoltia appears to be necessary based upon a comprehensive analysis of the Lemnaceae by D.H. Les, D.J. Crawford, E. Landolt, J.D. Gabel, and R.T. Kimball (2002). More that 4,700 characters were studied, including data from morphology and anatomy, flavonoids, allozymes, and DNA sequences from chloroplast genes (rbcL, matK) and introns (trnK, rpl16).

The Angiosperm Phylogeny Group (APG) has proposed some significant changes in the classification of many traditional angiosperm families, including the placement of all duckweeds in the Araceae rather than the Lemnaceae. Nomenclatural changes are cited under the APG II system (2003) and superceeded by APG III system (2009). These changes are based on computer-generated evolutionary trees or cladograms. Thousands of data characters have been used, including morphology, anatomy, flavonoids, allozymes, and DNA sequences from chloroplast genes and introns. The Jepson Manual Second Edition (2012) essentially follows the changes summarized in the following reference by W.T. Judd, et al. 2008. Since the genus Landoltia was proposed by D.H. Les and D.J. Cawford in 1999, several classic papers on the phylogeny of the duckweed subfamily (Lemnoideae) and other aroids (Araceae) have used the name Landoltia. In my opinion, the name Landoltia is warranted because it is consistent with the objectives of the Jepson Manual 2nd Edition (2012) based on phylogenetic studies using plastid DNA.

An argument for replacing the names Landoltia punctata and Spirodela punctata with the previous name Spirodela oligorrhiza has been made by Daniel B. Ward (2011). In order to make sure we are referring to the same species, Ward has suggested calling this "Lesser Greater Duckweed" to avoid confusing it with the larger species of Spirodela (S. polyrrhiza & S. intermedia) called "Greater Duckweeds." In this article I will simply call it LG Duckweed instead of Lesser Greater Duckweed.

• Ward, D.B. 2011. "Spirodela oligorrhiza (Lemnaceae) is the Correct Name for the Lesser Greater Duckweed." J. Bot. Res. Inst. Texas 5 (1): 197-203.

Spirodela punctata (Meyer) Thompson was named by C.H. Thompson in 1898 based on a collection from the 1938-1842 Wilkes Expedition, labeled Orange Harbor, Tierra del Fuego. Whether this collection actually came from the tip of South America is debatable. The parenthetical author G.F.W. Meyer described this species earlier as Lemna punctata from a type specimen collected in Guyana, South America in 1818. Unfortunately, Meyer's original type specimen was lost. According to Ward (2011), LG Duckweed does not occur in the areas where these collections were made: The Tierra del Fuego collection was mislabeled and the Guyana collection was not LG Duckweed! Futhermore, he states that the only native Spirodela in South America is S. intermedia. Since Meyer's type speciment was lost, Ward neotypified the species as Lemna punctata G.F.W. Meyer. Thompson's binomial is still Spirodela punctata (Meyer) Thompson; however, this no longer refers to LG Duckweed. It is now the correct binomial for the South American Spirodela intermedia. The correct name for LG Duckweed now becomes Spirodela oligorrhiza (Kurz) Hegelmaier, a name published by Hegelmeier in 1868. Hegelmeier apparently never saw the South American specimens discussed above, so his name is probably based on the true LG Duckweed.

Ward's 2011 neotypification will make Landoltia a synonym of Spirodela and no longer available for the intended LG Duckweed. The restoration of separate generic status for LG Duckweed now known as Spirodela oligorrhiza (Kurz) Hegelm. will require the creation of a new genus name. The binomial Spirodela punctata (Meyer) Thompson will now refer to the South American species known as Spirodela intermedia W. Koch. By neotypification the name Landoltia also becomes correctly applied to Spirodela intermedia.

According to E. Landolt (Personal Communication, 2012), the name change proposed by Ward in untenable:

• "I think this problem cannot be solved definitely. The main problem is the fact that it is not possible to decide which species Meyer was describing under the name of Lemna punctata. Certainly, it has to be a species of the genus Spirodela sensu lato because we don't know any other species within the Lemnoideae with pigment cells ("punctata"). The description of Meyer is very rudimentary. I could not find any herbarium specimen collected by Meyer. His description could concern Spirodela oligorrhiza, Spirodela intermedia or Spirodela polyrrhiza. I collected all of these species in northern South America. The description fits best for Spirodela oligorrhiza because it mentions 2-to 3 roots per frond. Most individuals of S. oligorrhiza in nature have 2 to 5 roots. S. polyrrhiza and S. intermedia mostly have more than 8 roots (up to 18). Only very rarely and only in very young fronds they show less then 5 roots. L. punctata was collected by Meyer in Guyana. On the other side, S. intermedia is known from the neighbouring state Surinam and surely is indigenous in the region. S. polyrrhiza and S. punctata might be introduced to South America. Today, S. punctata is frequent in the regions of Rio and Sao Paulo, in Venezuela, Colombia and Ecuador. I have collected S. polyrrhiza in Colombia and Ecuador. Even if S. punctata is introduced into South America it is not known at which year the introduction took place for the first time. It looks like S. punctata would be easily distributed by ship from harbour to harbour and from there by bird to places within a continent."

  "I can understand that Thompson choose a new type for Lemna punctata. The correctness of his decision is not disputed. I checked the neotype the collection of Wilkes from copies in four different Herbara. It is clearly the species which is now called "punctata". It is not important if the material was collected in Orange Harbor or somewhere else. Because it is not possible and will probably never be possible to decide the identity of Lemna punctata with certainity it is not advisable to change the correctly published neotype of Thompson. If we change the type of L. punctata again we will have a terrible chaos in nomenclature. Therefore I am not following the proposal of Ward."

My objection to Ward's proposed neotypification is based on two primary points. (1) He is neotypifying Meyer's lost type specimen with Spirodela intermedia. It is impossible to know with 100% certainty which species Meyer was describing under the name Lemna punctata back in 1818. It could have been the "LG Duckweed" that we know as Landoltia punctata (Spirodela punctata = Spirodela oligorrhiza), or it could have been another species of Spirodela such as S. intermedia. Why complicate this taxonomy based on speculation. (2) Cladistical analysis has clearly shown that Spirodela punctata belongs in a separate genus (Landoltia), otherwise the grouping of Spirodela with 3 species is paraphyletic. The trend in modern floras such as the Jepson Manual Second Edition (2012) is for consistent monophyletic groupings.

Cladogram From Chis Pires & Kenneth Sytsma (2002)

Themidaceae: A. Santa Rosa basalt brodiaea (Brodiaea santarosae), a recently described species from the Santa Rosa Plateau of Riverside County. B. Ithuriel's Spear (Triteleia laxa) from Kern County. C. Golden stars (Bloomeria crocea) in San Diego County.

A New Brodiaea From The Santa Rosa Plateau: Santa Rosa Basalt Brodiaea MADROÑO Vol. 54: 187-198 (2007)   Santa Rosa Basalt Brodiaea FREMONTIA Vol. 37 (2): 20-27 (2009)

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Significant Changes To The Snapdragon Family (Scrophulariaceae)

Computer generated monophyletic clades based on chloroplast DNA have resulted in major changes to the Scrophulariaceae. Plantago, Penstemon, Veronica, Linaria, Antirrhinum, Keckiella, & Digitalis are now placed in the Plantaginaceae. Mimulus with its thigmotropic stigma is placed in the Phrymaceae. Traditional genera retained in the Scropulariaceae include Verbascum and Scrophularia. Other genera placed in the Scrophulariaceae include Buddleja and Myoporum. Indian paintbrush (Castilleja), Indian warrior (Pedicularis), and owl's clover (Orthocarpus) are placed in the parasitic family Orobanchaceae with the broomrapes (Orobanche). Other closely-related families representing separate clades are the Paulowniaceae, Lentibulariaceae, Acanthaceae and Bignoniaceae. Members of the Martyniaceae (Proboscidea, Martynia & Ibicella) are included in the Pedaliaceae with Uncarina, Harpagophytum & Sesamum.

Scrophulariaceae: (A) California bee plant (Scrophularia californica ssp. floribunda), (B) butterfly bush (Buddleja davidii) and (C) prostrate myoporum (Myoporum parvifolium). Significant changes have been made to the once enormous snapdragon family (Scrophulariaceae). Traditional genera retained in the Scropulariaceae include mullein (Verbascum) and figwort (Scrophularia). Buddleja (formerly of the Loganiaceae) and Myoporum (formerly of the Myoporaceae) are now placed in the Scrophulariaceae.

Orobanchaceae: Although (A) Indian paintbrush (Castilleja affinis ssp. affinis) and (B) Indian warrior (Pedicularis densiflora) were once placed in the snapdragon family (Scrophulariaceae), they are now placed in the parasitic broom-rape family (Orobanchaceae) along with (C) chaparral broom-rape (Orobanche bulbosa). The latter holoparasitic plant is attached to the roots of chamise (Adenostoma fasciculatum)--see roots of host shrub in photo. The Orobanchaceae are considered monophyletic, including the complete parasites lacking chlorophyll (holoparasites) and the hemiparasites with green leaves (Castilleja and Pedicularis). According to Judd et al. (2008), this affinity is supported by DNA cladistic analysis, hemi to holoparasitic habit, hair morphology and possibly their racemose inflorescences.

Wayne's Word Article On Parasitic Flowering Plants     Wayne's Word Article On Mycotrophic Flowering Plants

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Major DNA Changes In Plantaginaceae
Many Species Transferred From Other Plant Families
[See Globularia x indubia Planted At Palomar College]

As I stated above, computer generated monophyletic clades based on chloroplast DNA have resulted in major changes to families, such as the Scrophulariaceae. For example, Plantago, Penstemon, Veronica, Linaria, Antirrhinum, Keckiella, & Digitalis are now placed in the Plantaginaceae.

Plantaginaceae: (A) Showy penstemon (Penstemon spectabilis var. spectabilis) and white-lined sphinx moth (Hyles lineata), (B) Chinese houses (Collinsia heterophylla), and (C) and foxglove (Digitalis purpurea). These common genera were formerly placed in the snapdragon family (Scrophulariaceae) but have now been transferred to the Plataginaceae.

Hyles lineata & Penstemon spectabilis var. spectabilis

Plantaginaceae: (A) Red-seeded plantain (Plantago rhodosperma) and (B) Nutall's snapdragon (Antirrhinum nuttalianum ssp. nutallianum. Many familiar genera of the Scrophulariaceae, such as snapdragons (Antirrhinum), Chinese houses (Collinsia), penstemons (Penstemon and Keckiella), toadflax (Linaria), and speedwell (Veronica) are now placed in the plantain family (Plantaginaceae). Plantain seeds (Plantago) are the source of the soluble fiber used in diet supplements such as Metamucil® and Hydrocil®. 2 very unlikely families that were also moved into the Plataginaceae are Hippuridaceae (including "mare's tail" Hippuris) and Callitrichaceae (including "water starwort" Callitriche). I saw water starwort in the vernal pool near Palomar College many years ago; and I saw a population of mare's tail in a marsh along the Walker River north of Bridgeport, CA. Unfortunately I cannot find images of either in my extensive Kodachrome collection.

A. Common plantain (Plantago major). B. Minute psyllium seeds seeds of the genus Plantago (including Plantago psyllium and P. ovata) are used in laxatives and soluble fiber supplements. The husk around each seed contains a water-soluble mucilaginous gum. Ground seeds (right) imbibe water and swell, forming a thick, jelly-like, mucilaginous mass of soluble fiber.

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Phrymaceae: Red bush monkeyflower (Mimulus aurantiacus, formerly M. puniceus). This common shrub of coastal San Diego County was once placed in the snapdragon family (Scrophulariaceae). It is now placed in its own family, the Phrymaceae. The spreading stigma lobes are thigmotropic and close together with the slightest touch of your finger or an incoming pollinator, such as the bill of a hummingbird. This action decreases the chance of self pollination and favors cross pollination, especially if the incoming pollinator is covered with pollen from another monkeyflower blossom. When the bill or head of the hummingbird enters the blossom and touches the stigma it immediately closes. Pollen carried by the bird is trapped within the closed stigma lobes. As the bird probes for nectar deep in the corolla it also picks up fresh pollen from the anthers. But when it leaves, there is little chance of this newly acquired pollen touching the stigma because it is already closed, thus averting any self pollination.

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Chocolate (Theobroma cacao)

Major Changes In The Mallow Family (Malvaceae)

The mallow family contains some of the most beautiful wildflowers in San Diego County and some economically important species, including cotton & okra. Now several additional economically important families have been merged with the Malvaceae, including the chocolate family (Sterculiaceae: Cacao & cola nut), the basswood family (Tiliaceae: Basswood & jute), and the bombax family (Bombacaceae: Kapok & balsa).

Malvaceae: A. California Ayenia (Ayenia compacta), formerly in chocolate family (Sterculiaceae). B. Cheeseweed (Malva parviflora). C. Pink Flame Tree (Brachychiton discolor), formerly in the Sterculiaceae.

Economically Important Species in the Malvaceae: Left: Kapok (Ceiba pentandra), formerly in the bombax family (Bombacaceae). Center: Okra (Hibiscus esculentus). Right: Cotton (Gossypium) tetraploid hybrid).

California Native Wildflowers in the Malvaceae: Left: Bush Mallow (Malacothamnus fasciculatus) in Palomar College Arboretum. Center: Apricot mallow (Sphaeralcea ambigua var. ambigua). Right: Desert 5-Spot (Eremalche rotundifolia).

DNA phylogenetic trees (cladograms) clearly show that the chocolate family (Sterculiaceae), basswood family (Tiliaceae), and bombax family (Bombacaceae) are not monophyletic. They are better treated as subfamilies within the mallow family (Malvaceae). This is similar to placing the duckweed subfamily Lemnoideae within the Arum family (Araceae).

Monophyletic: A taxonomic group that represents a single branch (clade) in a cladogram, and having a common ancestor. For example, all birds and reptiles are thought to have descended from a single common ancestor and are monophyletic. Humans (Homo) and chimpanzees (Pan) are also monophyletic.

Terms Used For Taxonomic Groupings:

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Changes To The Martyniaceae & Pedaliaceae (APG II but not Jepson Manual)

Pedaliaceae (Martyniaceae in Jepson Manual). Two North American species of Proboscidea. These species were formerly placed in the Martyniaceae along with Martynia and Ibicella. They are now placed in the Pedaliaceae in some references, along with sesame (Sesamum indicum), Uncarina and Harpagophytum: Left. Proboscidea louisianica ssp. louisianica is a native annual in the eastern United States and is naturalized in coastal Coalifornia, including San Diego County. The yellow lines in the corolla throat are nectar guide lines that direct pollinator bees to the nectar source. Right: Proboscidea althaeifolia is a native perennial in the Colorado Desert of the southwestern United States and Baja California. It blooms during the scorching heat of July and August, long after most other desert wildflowers have bloomed and gone to seed. It occurs sparingly in Anza-Borrego Desert State Park of San Diego County.

Large, hitchhiker seed capsules of the Pedaliaceae. From left: Ibicella lutea (formerly placed in the Martyniaceae), Harpagophytum procumbens, and Uncarina grandidieri.

Devil's Claws: Hitchhikers On Big Animals   The Ultimate & Most Painful Hitchhikers Sesame: An Important Seed Plant

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A Number Of Familiar Names Have Changed In Jepson 2nd Edition
The Genus Lotus Has Been Changed To Acmispon

Deerweed (Lotus scoparius var. scoparius), a native shrubby perennial found throughout the Palomar College Arboretum and adjacent coastal sage scrub. Much to the chagrin of botanists like myself, this genus has been renamed and the new scientific trinomial is Acmispon glaber var. glaber. In the desert var. brevialatus, the keel is longer than the wings.

Cactaceae: The Cholla Cactus Genus Opuntia Has Been Changed To Cylindropuntia
Cyperaceae: Naked Stem Bulrushes Scirpus Have Been Changed To Schoenoplectus
Asteraceae: The Genus For Cheesebush Hymenolcea Has Been Changed To Ambrosia

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Where To See Wildflowers In Southern California Favorite Anza-Borrego SD-Riverside Witch Creek Fire California Special Santa Rosa Plateau Brodiaea Joshua Tree Marine Parasite

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Brassica fruticulosa In San Diego Couty

All text material & images on these pages copyright © W.P. Armstrong