Notes on genealogical relationships, identification, and medicinal properties and/or uses

Angiospermae (angiosperms or flowering plants) = Magnoliidae Novak ex Takht.
Most non-flowering seed plants (gymnosperms) disperse their microspores (male spores) as pollen and megaspores (female spores) as part of the seeds through the aid of wind, while most flowering seed plants (with some exceptions) do all this through the aid of animals. In flowering seed plants, it can be said that microspores are dispersed as pollen during the stage that produces flowers, while megaspores are dispersed as part of the seeds during the stage that produces fruit. It is often assumed that the ovule provides protection for the developing megagametophyte (female gametophyte) and the seed (resulting from fertilization of the egg of megagametophyte) provides protection for the developing embryo that is thrust into or embedded within megagametophyte tissue. Perhaps, one advantage of the flowering seed plants is their ability to protect seeds within fruits until the time is right for dispersal. Even in fleshy, edible fruits, the seeds can be protected until they become hard (lignified), indigestable, and the fruits are ready to be consumed by animals. In turn, animals can assist by eating the fleshy-fruits so that the indigestable seeds can be spread over greater distances. In some flowering plants, the fruits can be shed as units, frequently containing several sections called carpels with multiple seeds. In others, the fruits (or contained carpels) can break open, allowing the seeds to become individually liberated from the fruit. It is commonly noted that the gametophytes of flowering plants, packaged within pollen grains and ovules, are usually even more minute (with fewer cells) than those of other seed plants. In contrast, sporophytes can develop into giant flowering trees, such as the African baobab (Adansonia digitata), the South American ombu (Phytolacca dioica), the Indian banyan (Ficus bengalensis), and the Australian Eucalyptus regnans, that are enormous, but do not grow as tall or massive as some of the sporophytes of certain non-flowering seed plants in the family Cupressaceae, such as the the coast redwood (Sequoia sempervirens), the giant sequoia (Sequoiadendron giganteum) and the Montezuma bald cypress (Taxodium mucronatum). The sporophytes can be secondarily reduced in size, as in the lemnids (duckweeds), but in this group, the tiny sporophytes are still larger than the minute female gametophytes of flowering plants.

Apiaceae - Lindl. - Family Name
Common name

Berberidaceae - Juss. - Family Name
Common name
Barberry family - - (NM)
Perennial herbs, shrubs, or small trees, deciduous (seasonally shedding leaves) or evergreen (bearing leaves thoughout the year); stems with 3-parted or unparted (simple) spines (derived from leaves at nodes) or without spines; branches often simple (appearing like wands); leaves usually spirally arranged on stem, alternate (rarely opposite), simple or pinnately compound (with leaflets arranged in a feather-like fashion), lobed or unlobed, leaf or leaflet margins entire (without teeth) or toothed with teeth sometimes bearing apical spines, lobes of leaves sometimes bearing spines, stipules [pair of often tiny leaf-like appendages at base the leaf stalks (petioles)] common; the cluster of flowers most commonly an indeterminate inflorescence called a raceme comprising a single axis with indefinate (indeterminate) growth from bottom upward that bears stalked (pedicellate) flowers or more rarely various other inflorescence types, such as umbels with flower stalks (pedicels) attached close together or at one point, or flowers solitary in axil (angel between stem and leaves); the individual flowers small, radially symmetrical (divided into equal halves along any diameter), perfect (bisexual with both male and female parts); perianth often bicyclic (both sepal-like and petal-like parts in two whorls or series, although sometimes slightly helical), sometimes also with 6 (to 7) whorls of distinct parts [often 3-merous (with 3 parts per whorl), but sometimes each whorl of 2 or 4 or more rarely 5 parts], the parts of the outer 2 whorls commonly referred to as sepals and those of the inner 2 (4 or 5) whorls commonly referred to as petals [the range in the number of perianth parts (sepals and petals) in family variable (e.g., Podophyllum L. with sepals 6-9 and petals 6-9)], sometimes with nectar secreting cells at the bases of 6 inner petals; male parts (stamens) bicyclic (in two whorls) and opposite inner most whorls of petals; these stamens mostly 6 with each of the two whorls 3-merous (comprising 3 stamens) or more rarely 4 with each of the two whorls 2-merous [the number of stamens sometimes more by proliferation (e.g., Podophyllum with 12-18)]; anthers (terminal pollen bearing parts of stamens) each dithecal (two lobed) and tetrasporangiate (each lobe with two pollen bearing cavities eventially merged as one cavity), these anthers opening to release pollen by basal valves (flaps) or sometimes longitudinal slits; female part (pistil) superior (located above perianth parts), with apparently a single carpel (sometimes interpreted as derived from the union of 2-3 carpels); pre-seeds (ovules) 1 to many attached marginally or near base of one (rarely two) locules (cavities) within the basal part of pistil called the ovary that ripens (once ovules are fertilized) as a small (seed containing) fleshy (somewhat juicy) simple fruit called a berry (rarely a dry fruit).
The major characters that distinguish this family from others include a multicyclic (= multi-seriate) perianth composed of many parts in 3-(2- or 4-)merous whorls differentiated into distinct (outer) sepal-like segments (collectively called the calyx) and (inner) petal-like segments (collectively called the corolla). The innermost petal-like parts are nectariferous, bearing a group of cells that secrete nectar (a fluid rich in sugar and other nutrients). The stamens are bicyclic (or biseriate) and the pistil (= gynoecium) is single, superior, and apparently (?) composed of one carpel (but see below).
The whorled, trimerous flowers (with each whorl often of 3 segments) are themselves (with or without opposite stamens) probably ancestral for the broader (huge) lineage referred to as the Mesangiospermae (= mesangiosperms). This may explain why the 3-merous, whorled condition is so common among early branching members of the major sublineages (magnoliids, monocots, and eudicots) of the huge mesangiosperm group; and the barberry family is an early branching member of the eudicots.
For example, the multicyclic (or multi-seriate) perianth of the barberry genus Berberis can include two outer, 3-merous whorls (6 yellow, more spreading, sepal-like parts, falling off almost immediately after the flowers open) and two inner, 3-merous whorls (6 yellow, more erect, more persistent, petal-like parts with basal nectaries). The so-called 'C' nectaries at base of the petal-like parts may be at least one shared, derived character of the subfamily Berberidoideae Kosteletzky (including the genera Ranzania and Berberis); however, the number of perianth parts of this lineage can range from only 3 to as many as 12. A bicyclic calyx of distinct sepal-like parts is somewhat differiated from a bicyclic corolla of distinct petal-like parts; and the 6 stamens opposite the surrounding, often more erect petal-like parts are also bicylic (in two 3-merous whorls). Stemens sensitive to touch may be another shared, derived character of the subfamily Berberidoideae.
The carpel [the structure in which the enclosed seeds (developing from the fertilized ovules) are produced] is considered the smallest unit of the female part of the flower called the pistil. Simple pistils are composed of only one carpel. Compound pistils are composed of more than one united carpel. Although members of the barberry family appear to have flowers with pistils composed of only one carpel, these pistils may actually be composed of 2-3 united carpels. Since the fruit of the family usually with a fleshy fruit wall and one to many seeds develop (once ovules are fertilized) from a single ovary of a single flower, it is considered a fleshy, simple fruit called a berry (the basic, although not the exclusive fruit type of the family).
A diversity of chemicals called alkaloids (some with medicinal potential, including the most common yellow pigmented, benzyl-isoquinoline alkaloid called berberin) are found throughout the family Berberidaceae. For example, the branches, roots and stems of all species of Berberis s.l. have a yellow center due to the presence of berberine. Although not all species of Berberis s.l. are singled out as medicinal and listed in the paragraphs or records of the Spanish-English dictionary of names, the alkaloid berberine, present in rhizomes (underground stems) and bark of probably all Berberis species, has marked antibacterial effects (often employed in eyewashes) and can also be used as a bitter tonic. Even though berberine is poorly absorbed by the body, preparations containing this alkaloid can be effective for oral treatment of various gastrointestinal infections, especially bacterial dysentery. Some of these types of alkaloids or other possible chemical constituents (e.g., certain bioactive ligans, such as podophyllotoxin, and a few other types of alkaloids) likely account for the most common medicinal uses of species in the genera Berberis, Caulophyllum, and Podophyllum.
Eight species of Berberis s.l. (the genus defined in the broad sence, including Mahonia s.str. and one other recently segregated genus) are the only members of the family in New Mexico.

Berberis - L. - Genus Name
Since 1818, botanists have been debating whether Mahonia should be in the same genus as Berberis or not. The genus Berberis L. (1754) and its segregated genus Mahonia Nuttall (1818) can be considered the two largest members of family Berberidaceae.
Those species characterized as evergreen (not seasonally leaf shedding) shrubs or rarely small trees most often with stems not spiny and leaves pinnately compound (composed of three or more leaflets) are sometimes referred to by the genus name Mahonia Nuttall; however, B. fendleri (formerly sometimes called Mahonia fendleri (Gray) Wooton & Standley) is a deciduous (seasonally leaf shedding) shrub with stems spiny and leaves simple (not composed of leaflets) and species of the unusual section Horridae Fedde, although possessing spineless stems and compound leaves, appear intermediate between the rest of Mahonia and Berberis s.str. on the basis of smooth, varnished dark red to reddish brown stems and several other characters (Ahrendt, 1961); therefore, botanists (since 1818) have disagreed on whether or not to recognize Mahonia as a separate genus from Berberis. Nevertheless, from analysis on the basis of DNA sequencing, Yu & Chung (2017) have provided a new circumscription of Mahonia, limiting this genus to core Mahonia, which could also be referred to as Mahonia s.str. However, the previously defined genus Mahonia, which could be referred to as Mahonia s.l., is a paraphyletic group. Therefore, an additional segregation (separation) from previously defined Mahonia of two newly defined genera named Alloberberis Yu & Chung (2017) and Moranothamnus Yu & Chung (2017) was necessary to render the new circumscription of the genus Mahonia s.str. monophyletic and distinct from a monophyletic genus Berberis s.str. All this manipulation, involving changes of scientific names, is easier to understand by inspecting the genealogical tree (see below) that has resulted from the DNA analysis of Yu & Chung (2017).
Yu & Chung (2017) classification of Berberis s.l.:
o Berberis [= Berberis s.str. = simple-leaved Berberis lineage (or clade)]
o Mahonia (= core Mahonia,
excluding Mahonia sect. Horridae + Berberis claireae + Berberis s.str.)
o Moranothamnus (= Berberis claireae) sister to Berberis s.str. (see phylogenetic tree)
o Alloberberis (= Mahonia sect. Horridae) sister to Moranothamnus + Berberis s.str.
[The newly proposed genus Alloberberis comprises the following 11 species:
(1) Alloberberis eutriphylla (Fedde) C.C.Yu & K.F.Chung
(2) Alloberberis fremontii (Torr.) C.C.Yu & K.F.Chung (in New Mexico)
(3) Alloberberis haematocarpa (Wooton) C.C.Yu & K.F.Chung (in New Mexico)
(4) Alloberberis higginsiae (Munz) C.C.Yu & K.F.Chung
(5) Alloberberis longipes (Standl.) C.C.Yu & K.F.Chung
(6) Alloberberis muelleri (I.M.Johnst.) C.C.Yu & K.F.Chung
(7) Alloberberis nevinii (A.Gray) C.C.Yu & K.F.Chung
(8) Alloberberis pimana (Laferr. & Marroq.) C.C.Yu & K.F.Chung
(9) Alloberberis pinifolia (Lundell) C.C.Yu & K.F.Chung,
including Alloberberis pinifolia var. coahuilensis (C.H.Mull.) C.C.Yu & K.F.Chung
(10) Alloberberis swaseyi (Buckley) C.C.Yu & K.F.Chung
(11) Alloberberis trifoliolata (Moric.) C.C.Yu & K.F.Chung (in New Mexico),
including Alloberberis trifoliolata var. glauca (I.M.Johnst.) C.C.Yu & K.F.Chung.]
o Berberis s.l. (= Mahonia s.s. + (Alloberberis + (Moranothamnus + Berberis s.s.))).
It is hard to tell whether the new segregate genus names Alloberberis and Moranothamnus of Yu & Chung (2017) in Berberis s.l. have become officially accepted, although it is apparent that some recent (2018) publications and web sites are using both segregate names; and Yu & Chung (2017) has certainly shed some doubt in the minds of some botanists on whether it was wise to except the merging of Mahonia into Berberis s.l. However, what might be debatable among some other botanists is that all these changes in names are unnecessary, since the 'lumping' of all these segregate names as the single genus called Berberis s.l. also results in a monophyletic group. Why make changes in previous names that clearly correspond to well established monophyletic groups? Because name changes should be minimized as much as possible, such changes involving polyphyletic or paraphyletic groups should be of much higher priority; and even if the scientific names of this 'splitting' classification of Yu & Chung (2017) become widely adopted, the sister relationships of these genera (Mahonia s.s. + (Alloberberis + (Moranothamnus + Berberis s.str.))) does not prevent subsequent 'relumping' of all of them into (again) the single genus Berberis s.l. (see phylogenetic tree). Although Moranothamnus (= Berberis claireae) has only recently been found to be the closest sister to Berberis s.str., it has long been known that the species of Berberis (or Mahonia) section Horridae (= Alloberberis) have a closer relationship to Berberis s.str. than the species of core Mahonia (Ahrendt, 1961). This has already been supported by analysis based on the DNA (internal transcribed spacer) sequencing of Kim et al. (2004). Together with the Japanese herb Ranzania japonica T. Ito, the genera Berberis and Mahonia (Kim and Jansen, 1998), as well as the recently proposed genera Alloberberis and Moranothamnus (Yu & Chung, 2017), comprise a monophyletic group (lineage or clade) with the same base chromosome number (x = 7) within the family Berberidaceae.
Species of paraphyletic Mahonia s.l. (core Mahonia + Alloberberis + Moranothamnus) are generally and easily distinguished (from Berberis s.str.) by the presence of spineless stems and pinnately compound leaves (each with leaflets arranged in a feather-like fashion and opposite to each other with a single terminal leaflet). Monophyletic Mahonia s.str. (= core Mahonia, excluding the segregated genera Alloberberis and Moranothamnus) has about 70 species found in Western North America (including Mexico), Central America, and Eastern Asia. Alloberberis [= Berberis (or Mahonia) section Horridae) has 11 species (see above) found in Western North America (e.g., the species until recently called Berberis fremontii Torr., B. haematocarpa Wooton, and B. trifoliolata Moric. native in New Mexico).
Species of Berberis s.str. are generally and easily distinguished by spiny stems and simple leaves (each without leaflets). Berberis (= Berberis s.str.) has about 500 species found in temperate and subtropical regions of the world (but not in Australia).
The Western Mexican genus Moranothamnus (with only one species sister to all the species of Berberis s.str.) has (like Mahonia s.s. and Alloberberis) spineless stems and compound leaves (each with leaflets).
Although Alloberberis [= Berberis (or Mahonia) section Horridae] has species that are mostly distributed in the Southwest United States and Northwest Mexico, the shrub species Alloberberis eutriphylla (Fedde) Yu & Chung can be found in the oak forests of Sierra de Zapalinamé in Northeast Mexico.
Identification of Berberis spp. in New Mexico (data from Allred and Ivy, 2012):
Any suspected species of Berberis s.l. should first be checked to see whether it possesses the family characteristic summerized above. If a plant to be identified is a shrub (even though woody aerial stems can sometimes be very short), bearing simple leaves or pinnately compound leaves (sometimes holly-like) with margins of leaflets toothed and often prickly or lobes with terminal prickles (only rarely without teeth or lobes), than provided this plant can also be accommodated by the family description, it is likely a member of Berberis s.l. The group of species called Berberis s.str. has usually spiny stems and always simple leaves that are deciduous. This group in New Mexico includes B. fendleri, B. thunbergii, and B. vulgaris. Both B. fendleri and B. vulgaris have 3-parted spines [although B. vulgaris can possess simple (unparted) spines]; and B. fendleri has leaf margins untoothed (entire) or with 3-12 teeth on each side, purple, two year old branches, and racemes bearing 4-15 flowers, while B. vulgaris has leaf margins with 16-30 teeth on each side, gray, two year old branches, and racemes with 10-20 flowers. The cultivated B. thunbergii can be distinguished from the rest of Berberis s.s. in New Mexico by spines simple (not 3 parted), leaves with margins entire (not spiny or toothed), and a cluster (inflorescence) with only 1-3 flowers.
The rest of the genus (sometimes called Mahonia) can be distingushed from Berberis s.s. by stems not spiny, evergreen (bearing leaves throughout the year), and individual leaves pinnately compound (with leaflets arranged in a feather-fashion and bearing one terminal leaflet).
B. trifoliolata [= Alloberberis trifoliolata (sometimes also called Mahonia trifoliolata (Moricand) Fedde)] can be distinguished from others in this paraphyletic group by leaves all with 3 leaflets (3-foliolate), usually covered with a translucent, waxy substance (glaucous), and the terminal leaflet without a stalk (sessile).
The rest of this paraphyletic group mostly possess leaves with 5-11 leaflets, often green (less wax covered), and the terminal leaflet stalked.
B. haematocarpa (= Alloberberis haematocarpa) and B. fremontii (= Alloberberis fremontii) possess a loose cluster of only 1 to 11 flowers.
B. haematocarpa [also called Mahonia haematocarpa (Wooton) Fedde] is distinguished by the terminal leaflet with a drawn out apical portion that projects much more than the lateral teeth and smaller berries solid and juicy, while B. fremontii [also called Mahonia fremontii (Torrey) Fedde] is contrasted by the terminal leaflet with an apical portion (not drawn out) that projects about the same as the lateral teeth and larger berries dry and swollen or expanded.
In contrast to both B. haematocarpa and B. fremontii,
B. repens [= Mahonia repens (Lindley) G. Don] and B. wilcoxii have dense clusters (racemes) of 25-50 flowers.
B. repens has leaflets thin and flexible with teeth of margins 6-24 on each side, while B. wilcoxii [= Mahonia wilcoxii (Kearney) Rehder] has leaflets thick and rigid with teeth of margins 3-5 on each side. Although all species of Berberis s.l. have clusters of small yellow flowers, both B. repens and B. wilcoxii have similar dense flower clusters and similar, dark blue berries. However, B. repens is a low growing shrub that trails or creeps close to the ground, while B. wilcoxii can grow up to 2 meters tall. Nevertheless, the prickly, holly-like, pinnately compound leaves and other similarities place both these species among certain other species growing outside New Mexico in a monophyletic of core Mahonia (Yu & Chung, 2017), which could be considered Mahonia s.str.
In New Mexico (data from Allred and Ivy, 2012), B. fendleri can be found wild in the northern mountains and plains on rocky slopes and within canyons; B. fremontii is wild in northwest parts of the state in pinon-juniper woodlands and plains grasslands; B. haematocarpa is wild in oak woodland, grassland, and desert shrubland; B. repens is wild and widespread in foothills and mountains; B. thunbergii is only found cultivated in the state; B. trifoliolata is wild in the eastern and southern parts of state; B. vulgaris is cultivated and sometimes escapes in the wild; and B. wilcoxii is uncommon in the extreme southwest bootheel area (more common in the Sonoran desert of Arizona and Sonora).
The probably most common B. repens (= Creeping Mahonia) in ponderosa pine and mixed conifer forests of New Mexico is a low growing shrub that trails and spreads close to the ground with a short, woody, aerial stem that is smooth (not spiny), with prickly, holly-like, pinnately compound leaves that turn red to purple with the coming of winter, and with densely clustered, bright yellow flowers and waxy covered (glaucous), blue-black berries.
According to The Plant List (Available from: http://www.theplantlist.org), B. aquifolium Pursh is the accepted
species name; and B. aquifolium subsp. repens (Lindl.) Brayshaw
[or B. aquifolium var. repens (Lindl.) Scoggan] is an infraspecific
taxon of B. aquifolium.
Therefore, some botanists treat B. repens as a creeping or trailing
subspecies or variety of the species B. aquifolium (= tall Oregon grape).
The following phylogenetic tree for Berberis s.l. is consistent with the analysis of Yu & Chung (2017):

        /---- Berberis s.s.
    /---|      
    |   \---- Moranothamnus
/---|
|   \-------- Alloberberis
|
\------------ core Mahonia (= Mahonia s.s.)
  • Ahrendt, L. W. A. (1961) Berberis and Mahonia: a taxonomic revision. Botanical Journal of the Linnean Society, 57(369), 1-410.
  • Kim, Y. D., Kim, S. H., & Landrum, L. R. (2004) Taxonomic and phytogeographic implications from ITS phylogeny in Berberis (Berberidaceae). Journal of Plant Research, 117(3), 175-182.
  • Kim, Y. D., & Jansen, R. K. (1998) Chloroplast DNA restriction site variation and phylogeny of the Berberidaceae. American journal of Botany, 85(12), 1766-1778.
  • Yu, C. C., & Chung, K. F. (2017) Why Mahonia? Molecular recircumscription of Berberis s.l., with the description of two new genera, Alloberberis and Moranothamnus. Taxon, 66(6), 1371-1392.

    Chenopodium - L. - Genus Name
    See also the genus Dysphania.
    Family Amaranthaceae
    According to Kelly W. Allred (Downloadable Version 2010): - - (NM)
    o Pale goosefoot C. albescens Small is not in New Mexico,
    but C. albescens of New Mexico authors is C. atrovirens Rydberg.
    o C. album Linnaeus var. missouriense (Aellen) Bassett & Crompton
    is the only native form of an otherwise exotic C. album Linnaeus;
    and C. album L. var. lanceolatum (Muhlenberg ex Willdenow)
    Cosson & Germain is an introduced variety.
    o The introduced Mexican tea C. ambrosioides L. is now called
    Dysphania ambrosioides (Linnaeus) Mosyakin & Clemants.
    o The pitted goosefoot includes C. berlandieri Moquin-Tandon
    var. berlandieri, C. berlandieri Moquin-Tandon var. sinuatum
    (Murr) Wahlenberg, and C. berlandieri Moquin-Tandon var.
    zschackei (Murr) Murr.
    o The introduced Jerusalem oak goosefoot (= Jerusalem-oak)
    C. botrys L. is now called Dysphania botrys (L.) Mosyakin
    & Clemants.
    o The blite goosefoot (= strawberry blite) are introduced varieties
    including C. capitatum (Linnaeus) Ambrosi var. capitatum and
    C. capitatum (Linnaeus) Ambrosi var. parvicapitatum S.L. Welsh.
    o C. desiccatum A. Nelson includes C. desiccatum A. Nelson
    var. desiccatum (= desert goosefoot) and C. desiccatum A. Nelson
    var. leptophylloides (Murr) Wahl (= plains goosefoot).
    o The leafy goosefoot C. foliosum (Moench) Ascherson and oak-leaf
    goosefoot C. glaucum Linnaeus var. glaucum are introduced, but
    C. salinum Standley now called C. glaucum L. var. salinum (Standley)
    B. Boivin (= Rocky Mountain goosefoot) is native.
    o The fetid goosefoot C. graveolens Willdenow is now called
    Dysphania graveolens (Willdenow) Mosyakin & Clemants.
    o The foetid goosefoot C. hircinum Schrader and nettle-leaf goosefoot
    C. murale Linnaeus are introduced.
    C. hircinum (native to South America with a distinctive odor and large
    leaves, both lobed and toothed) was once reported in southern New
    Mexico. However, it has apparently not been reported to occur in the
    state since the early 1900s. C. murale (native to Europe and Asia) is
    not that well known in the state.
    o The Over's goosefoot C. overi Aellen is now called C. capitatum (L.)
    Ambrosi var. parvicapitatum S.L. Welsh (= blite goosefoot or
    strawberry blite).
    o The name C. pratericola Rydberg is no longer prefered, now including
    the varieties C. desiccatum A. Nelson var. desiccatum and C. desiccatum
    A. Nelson var. leptophylloides (Murr) Wahl.
    o C. quinoa Willdenow is occasionally grown in gardens, but it
    apparently does not fruit in New Mexico and is not known to escape.
    o The red goosefoot C. rubrum L. is the native variety C. rubrum L. var.
    humile (Hooker) S. Watson or the introduced C. rubrum L. var. rubrum.
    o Rocky Mountain goosefoot C. salinum Standley is now known as
    C. glaucum L. var. salinum (Standley) B. Boivin.
    o Standley's goosefoot C. standleyanum Aellen is actually a large plant
    of the species C. hians Standley.

    Chenopodium album - L. - Species Name
    Family     Amaranthaceae
    In New Mexico, either the native Chenopodium album Linnaeus var. missouriense (Aellen)
    Bassett & Crompton or the introduced C. album Linnaeus var. album and
    C. album L. var. lanceolatum (Muhlenberg ex Willdenow) Cosson & Germain.

    Chenopodium berlandieri - Moquin-Tandon - Species Name
    Family Amaranthaceae
    In Mexico, the flowering tops of this species (as well as certain others
    in the genus) were at least once used as laxatives. - - (AJP1885-Mexico-8)
    Seeds for cultivated plants of the species are sold under the names
    Aztec spinach, Huauzontle, or Red Aztec spinach.
    Some of these cultivated types can sometimes be found in gardens
    (or excaped from gardens) in New Mexico. The seeds, flowering tops,
    or leaves can be used as food similar to closely related C. album L.,
    C. nuttalliae Saff. = C. berlandieri ssp. nuttalliae, or C. quinoa Willd.

    Chenopodium graveolens - Willd. - Species Name
    = Dysphania graveolens (Willdenow) Mosyakin & Clemants
    Family Amaranthaceae
    Certain Native Americans in New Mexico refer to this species by a name that can be translated as 'strong odor leaf' (Stevenson, 1915). It has been reported by this author that they steeped the plant in water and the vapor was inhaled for headache. The isolation of two new sesquiterpenoids, (+)-8-alpha-hydroxyelemol and (+)-8-alpha-acetoxycryptomeridiol, was done by chromatographic fractionation of a chloroform extract of the aerial parts of the plant (Mata, 1993). In Mexico, a tea is ingested to treat gastrointestinal ailments, headaches and fevers, and it is used as a vermifugue. The anthelmintic properties demonstrated by the flavanone called pinocembrin (Mata, 1993) could be related to the use of this plant in Mexican folk medicine for treatment of worms.

  • Rachel Mata (1993) Chemical Studies and Biological Aspects of Some Mexican Plants Used in Traditional Medicine in recent advances in phytochemistry volume 27, Phytochemical Potential of Tropical Plants, Proceedings of the Thirty-second Annual Meeting of the Phytochemical Society of North America, Miami Beach, Florida, August, 1992.

    Clerodendrum (Clerodendron) - L. - Genus Name
    Family Lamiaceae
    [In subfamily Ajugoideae (= Teucrioideae)]
    Clerodendrum chinense is a member of the lineage called the Asian clade
    (mentioned below) of the genus Clerodendrum sensu stricto (s.s.).
    Volkameria ligustrina (= Clerodendrum ligustrinum) of Tropical America
    appears to be in a lineage called the Pantropical Coastal clade; and this entire
    lineage appears sister to another lineage called the New World clade that includes
    the genera Aegiphila Jacquin, Amasonia L., Ovieda L. (only one species:
    Clerodendrum spinosum (L.) Spreng. = Ovieda spinosa L.), and Tetraclea A. Gray.
    These two sister lineages form a larger lineage that is sister to the genus
    Clerodendrum sensu stricto (s.s.), including two other sister lineages called
    the Asian and African clades. Clerodendrum chinense belongs to the Asian clade;
    and Clerodendron ligustrinum is now often placed in the Pantropical Coastal clade
    genus Volkameria L. as Volkameria ligustrina Jacq.
    For more details see:
    o WeArn, J. A., & Mabberley, D. J. (2011).
    Clerodendrum confusion-redefinition of, and new perspectives for, a large labiate genus.
    Gard. Bull. Singapore, 63, 119-124.
    o Yuan, Y. W., Mabberley, D. J., Steane, D. A., & Olmstead, R. G. (2010).
    Further disintegration and redefinition of Clerodendrum (Lamiaceae):
    implications for the understanding of the evolution of an intriguing breeding strategy.
    Taxon, 59(1), 125-133.

    'Copangel' - - (Apachian/Madrean region)
    This is made as a combination, two-fifths of which is the ground roots
    of the perennial herb Chuchupate (Ligusticum porteri = Ligusticum madrensis)
    that is found in the region centred on the Sierra Madre Occidental of northern Mexico.
    The Mexican national health programme has done clinical studies, determining that this
    combination is effective in the treatment of peptic ulcers.

    Coreopsis - L. - Genus Name
    Family Asteraceae
    The genus "Coreopsis" as traditionally defined appears to have species scattered here and there throughout almost all the major lineages within the tribe Coreopsideae Turner & Powell. This scattering of species in separate, not directly related lineages throughout most of the tribe indicates that the traditional genus "Coreopsis" is polyphyletic. The tribal relatives of the various (related or unrelated) species of "Coreopsis" include species of the genera "Bidens", Cosmos, "Coreocarpus", Thelesperma, and many other genera. These genera are all related to each other, because they are all derived from within the same monophyletic tribe. However, there has been much difficulty in separating the traditional genus "Coreopsis" from the traditional genus "Bidens". Although the genera "Bidens" and "Coreopsis" (as traditionally defined) may each prove to be polyphyletic, these genera are almost certainly paraphyletic, because other distinctive genera such as "Coreocarpus" and Thelesperma appear to be derived from within them. The genus Cosmos may even be derived from within "Coreocarpus" as it is presently defined.
    Another more distant relative is the genus Dahlia L., including native Mexican, medicinal species, such as Dahlia coccinea Cav. Although Thelesperma, Cosmos, and Dahlia are likely well defined genera and Dahlia appears sister to the well defined genus Dicranocarpus, the genera "Bidens", "Coreopsis", and "Coreocarpus" (with some species closely related to Cosmos) do not appear at all monophyletic on the basis of the DNA analysis reported in several publications.
    The well-defined genus Thelesperma appears to be nested within a lineage including mostly eastern North American species of "Coreopsis" (e.g., C. basalis, C. grandiflora, C. lanceolata, C. palmata, C. tinctoria, and others) and some north temperate species of "Bidens" (e.g., B. cernua, B. frondosa, B. beckii, and B. tripartita), but not including members of the lineage of Coreopsis gigantea (Kellog) H.M. Hall and C. californica (Nutt.) Sharsmith (an uncommon species in southwestern New Mexico) only recently placed in the segregated genus Leptosyne DC. as Leptosyne gigantea Kellog and Leptosyne californica Nutt. According to DNA analysis, the two last mentioned species together with other relatives are not part of the lineages within the polyphyletic genus "Coreopsis" that are most closely related to the genus Thelesperma. However, even with the placement of C. californica and relatives in the seperate, segregated genus Leptosyne mostly restricted to the state of California, the rest of the genus "Coreopsis" (without these segregates) is still a highly polyphyletic group in its tribe. See discussion of the species of "Coreopsis" and also those of the polyphyletic genus "Bidens" that are most closely related to Thelesperma. Since "Coreopsis s.l." has a scattering of species in separate, not directly related lineages throughout most of its tribe, some other genera not found in New Mexico have been segregated from it, but the decision on which monophyletic group should be called Coreopsis s.s. is still pending.
    A head typical of the family Asteraceae but found in some other families as well is any compact cluster of stalkless (sessile) or short stalked (subsessile) flowers. The flower heads [also called capitula (plural), capitulum (singular)] of the family Asteraceae can be generally said to comprise (almost always) a surrounding involucre of phyllaries (involucral bracts) and 1-many (usually small) individual sissile or subsissile flowers called florets inserted with phyllaries and often paleae onto the receptacle (base or axis of the head). If only some of the flowers of the head in the lineage (or family) under consideration have petals united into a short tube with a distal, more elongated, more or less flattened, strap-shaped lip [a lamina (singular), laminae (plural)] with (0)3(-4), usually short, apical (distal) lobes, these more peripheral, petal-like flowers are called ray flowers (or ray florets) and a head bearing these ray florets together with often a larger number of more centrally located tubular flowers called disk flowers (or disk florets) is referred to as a radiate head. A head without ray florets and only tubular disk florets [each usually perfect = bisexual (often with five male parts called stemens and one female part called a pistil)] is referred to as a discoid head.
    The genera "Bidens", "Coreocarpus", "Coreopsis", Cosmos, Dahlia, Dicranocarpus, Thelesperma mentioned above, and others (whether well defined or not) are included in the well defined lineage often referred to as the tribe Coreopsideae Turner & Powell. This lineage is characterized by compact flower clusters that are referred to as either radiate or discoid heads, with each flower cluster internally possessing modified leaves (bracts) called paleae (or chaff) attached with several to many tiny flowers (florets) on a somewhat widened (mostly flat to convex or conical to rarely globular) receptacle and most members of the tribe externally possessing a calyculus (outer whorl of more leaf-like bracts) immediately adjacent to the base of the more inward one or two whorls of usually wider, more erect bracts collectively called the involucre that initially encloses and protects the early developing cluster of florets, which, of course, eventually becomes exposed as the florets become more mature. See further discussion on the nature of bracts and the involucre of the Aster family.
    The lineage of tribal relatives of species of "Coreopsis" often referred to as the tribe Coreopsideae has a worldwide distribution, with a New World center of diversity especially in Mexico. The involucre of species of "Coreopsis" is composed of one or two series (whorls) of bracts (phyllaries) that initially enclose and protect the cluster of often many, small, immature flowers. Although the structure referred to here as the calyculus is sometimes viewed as the outer whorl of phyllaries of the involucre (or the outer whorl of involucral bracts) by some authors, the calyculus characteristic of most of the tribal relatives of "Coreopsis" but not entirely limited in occurrence to the tribe is treated here as distinct from the involucre, which is a structure that more directly subtends (extends immediately below) the cluster of florets, entirely encloses them in initial development, and occurs in nearly all members of the family Asteraceae. If what is here called the calyculus is consided the outer bracts (or outer phyllaries) of the involucre, then such an involucre like that found in "Coreopsis" and most of its tribal relatives is usually said to be distinctly dimorphic [composed of inner and outer involucral bracts (phyllaries) that are unlike in appearance, number, size, shape, structure, or orientation]. In this case, the outer part of the involucre of "Coreopsis" is said to be composed of (3-)8 or more, distinct (completely separate), green phyllaries that are more or less herbaceous (soft and leaf-like in color and texture), while the inner part is said to be more or less globose (spherical) to cylindrical in shape, comprising more or less 8 phyllaries in 1 or 2 or slightly more series that are usually distinct from one another but rarely slightly connate (slightly fused basally not much more than about 1/10th of their length), and more or less membranous (thin, somewhat soft, and tending to be translucent) with more or less scarious (paper-like) margins.
    The species C. lanceolata is the only member of the genus "Coreopsis" in New Mexico with leaves almost entire (without lobes or with very few) and the rest of the species of the genus in New Mexico (C. californica, C. basalis var. wrightii, C. grandiflora, and C. tinctoria) have leaves pinnatifid (lobed at various points along margins like the sections of a feather). Although the basal leaves of both C. lanceolata and C. grandiflora are linear-lanceolate [intermediate between linear (long and slender with more or less parallel sides) to lanceolate (several times longer than wide, broadest toward the base and narrowed to the apex)], the leaves of C. lanceolata are mostly basal (growing from the base of a stem), rarely lobed and the radiate heads (daisey-like flower clusters) are on long (up to 40 cm) naked (leafless) peduncles (stalks of the flower clusters), while the leaves of C. grandiflora are mostly cauline (found along most of the stem), often lobed [pinnately divided (pinnatifid) with the lateral segments (lobes) linear] and the similar looking radiate heads are on short (less than 20 cm) peduncles.
    All species of "Coreopsis" have ray florets that are either pistillate (with a fertile "female" part called a pistil) or neuter (with a "female" part absent or non-functional) and, consequently, they all have radiate heads.
    [A pistillate ray floret, almost always lacking "male" parts (stamens each made up of a slender filament and a terminal, pollen bearing anther), is fertile with a female part (pistil made up of a basal ovary that ripens as fruit, a slender column attached above the ovary called the style with two apical style branches, and two pollen receptive stigmas along the inner surfaces of the style branches). In the aster family, the ovary becomes a small, seed-like fruit (called an achene or cypsela), when the egg cell of a single ovule (pre-seed contained within the ovary) is fertilized by sperm from the pollen. A neuter ray floret, also usually lacking stamens, is sterile with a non-functional ovary (sometimes much reduced) and with or without a style (styliferous or not styliferous).]
    For example, in "Coreopsis" of New Mexico, C. californica has pistillate ray florets with styles, while C. basalis var. wrightii, C. grandiflora, C. lanceolata, and C. tinctoria have neuter ray florets without styles like their closer relatives in the genus Thelesperma. C. basalis var. wrightii, C. lanceolata, and C. grandiflora have disk florets with five outer lobes, while C. tinctoria has disk florets with 4 outer lobes. The ray florets are usually yellow throughout (rarely with red-brown blotch near their base) and disk floret apices yellow in C. lanceolata and C. grandiflora (both perennials), while ray florets are usually yellow with red-brown or purple near their base (rarely yellow throughout) and disk floret apices reddish brown or purplish in C. basalis var. wrightii and C. tinctoria (both annuals). See further discussion of C. tinctoria and some of its closest relatives in the genus Thelesperma.
    (Species in New Mexico more closely related to Thelesperma than most other members of the polyphyletic genus "Coreopsis" include C. tinctoria, C. lanceolata, C. grandiflora, and C. basalis var. wrightii. Although also found wild in New Mexico, Coreopsis lanceolata is often cultivated in flower gardens. The rarer species C. californica in southwestern New Mexico is more distantly related to Thelesperma. Bohlmann et al. (1983) and Reichling and Thron (1989) have done some analysis of the secondary chemistry of species of "Coreopsis". Flavonoids and related chalcones and aurones, as well as unusual acetylenic compounds (polyacetylenes) and rare types of phenylpropanes have been identified. In the genus "Coreopsis", some of the chemicals called chalcones and aurones are responsible for the yellow color in the flowers. A similar chemistry is expected for species of "Bidens" and Thelesperma.)

    Coreopsis tinctoria - Nutt. - Species Name
    Family Asteraceae
    C. tinctoria can be distinguished from the rest of the wild species of the genus in New Mexico by the combination of an annual growth form with outer phyllaries (bracts of the calyculus) much shorter than inner ones (involucral bracts directly subtending the cluster of florets), ray flowers often with darkened red-brown or purplish base and the rays three lobed with middle lobe sometime further lobed into teeth but often larger (longer and wider) than the other two lobes, styles mostly blunt, disk corollas 4-lobed, and leaves once or twice pinnately divided (narrowly divided into somewhat slender and elongated lobes in a feather-like arrangement) with the terminal lobe often elongated more than the other lobes. See further discussion of C. tinctoria and some of its closest relatives in the genus Thelesperma.
    o The species Coreopsis tinctoria is reported to be used by the Kiowa (Vestal and Shultes, 1939), Lakota (Rogers, 1980) and Zuni (Stevenson, 1915) as a beverage (tea). When the flowering tops are boiled, the water eventually turns red or reddish. This may account for the use of this beverage by the Lakota to strengthen the blood. This species is also reported (Hamel and Chiltoskey, 1975) to have been known to the Cherokee as an antidiarrheal medicine. It could be of interest that species of the closely related genus Thelesperma is used routinely as a household tea and notably considered antidiarrheal by the Tewa of New Mexico. Coreopsis tinctoria is reported (Stevenson, 1915) to have been introduced by the Navajo to Zuni for use as a hot tea or coffee substitute. Like species of Thelesperma, it is often collected, folded, and tied into packets before drying it for later use. The dried packets are dropped in boiling water and then simmered. Both Coreopsis and Thelesperma are, of course, not only used as beverages but also as medicinals for various purposes by Native Americans. Like species of Thelesperma, the annual Coreopsis tinctoria is known to produce a yellow dye, which in acid conditions can take on a brownish, mahogany red color. This dye is not as useful for plant fibers, but it is commonly used to color yarn (wool). According to Reagan (1929), Coreopsis tinctoria is valued by the White Mountain Apache of Arizona for its dark, rich red dye.
    o Certain species of "Bidens" are also sometimes used to make tea or a coffee substitute similar to that of Coreopsis tinctoria Nutt. (= Golden tickseed), as well as almost any species of the related genus Thelesperma. Several species of "Bidens", some species of "Coreopsis", and probably most if not all species of Thelesperma are used as tea or coffee substitutes, providing a good example of how closely related plants are used for a similar purpose. Therefore, it should not be surprising that some Old World (European) species in this group (in genus "Bidens") are also known as household tea beverages.

    Dysphania - R. Brown - Genus Name
    Family Amaranthaceae
    In New Mexico, the introduced species Dysphania ambrosioides (= Mexican tea) and Dysphania botrys (= Jerusalem-oak), and the native species Dysphania graveolens (= Fetid goosefoot) have often been listed under the older names of Chenopodium ambrosioides, Chenopodium botrys, and Chenopodium graveolens, respectively.
    The aromatic (essential oil bearing) species of Chenopodium subgenus Ambrosia
    A.J. Scott with all sections of the subgenus has now been re-classified under the
    genus name Dysphania. According to the molecular (DNA) results (Kadereit et al.,
    2003, 2010; Fuentes-Bazan et al., 2012a,b), Dysphania aristata (L.) Mosyakin &
    Clemants should belong to the genus Teloxys [as T. aristata (L.) Moq.].
    o Kadereit, G., Borsch, T., Weising, K., & Freitag, H. (2003) Phylogeny of
    Amaranthaceae and Chenopodiaceae and the evolution of C4 photosynthesis.
    International journal of plant sciences, 164(6), 959-986.
    o Kadereit, G., Mavrodiev, E. V., Zacharias, E. H., & Sukhorukov, A. P. (2010)
    Molecular phylogeny of Atripliceae (Chenopodioideae, Chenopodiaceae):
    implications for systematics, biogeography, flower and fruit evolution, and the
    origin of C4 photosynthesis. American Journal of Botany, 97(10), 1664-1687.
    o Fuentes-Bazan, S., Mansion, G., & Borsch, T. (2012a) Towards a species level
    tree of the globally diverse genus Chenopodium (Chenopodiaceae). Molecular
    Phylogenetics and Evolution, 62(1), 359-374.
    o Fuentes-Bazan, S., Uotila, P., & Borsch, T. (2012b) A novel phylogeny-based
    generic classification for Chenopodium sensu lato, and a tribal rearrangement
    of Chenopodioideae (Chenopodiaceae). Willdenowia, 42(1), 5-24.

    Embryopsida Engler ex Pirani & J. Prado, 2012 (= embryophytes)
    It is difficult to provide a short definition of land plants called embryophytes and distinguish them (in a few words) from all other organisms without first discussing in some detail the alternation of generations involving the sporophyte and gametophyte and the nature of land plant spores. These organisms comprise a lineage of streptophytes s.l. that do not shed their fertilized egg (called the zygote) like most "algae", but retain the zygote protected within the female sex organ (e.g., archegonium) of the parent (zygote bearing) plant called the gametophyte or within the protective, highly condensed structure of the sporophyte called the seed. In the lineage of land plants called seed plants, the shared, derived character called the seed can be considered a modified female sporangium (nucellus) that is sheathed by one or two protective envelopes called integuments with (in presently living seed plants) only one tiny, sperm or pollen tube receptive opening called the micropyle. This modified female sporangium does not split open like those of "free-sporing plants" but retains a single, functional female spore that contains the embryo bearing female gametophyte. It just so happens that the highly condensed structure called the seed contains (within the wall of its single functional spore) a highly reduced, already fertilized, parent female gametophyte retaining the zygote that has already developed into an embryo! Most "algae" cannot possibly produce an embryo, because the zygote usually never undergoes the type of cell division called mitosis to produce a multicellular stage. Usually after the zygote of most "algae" is shed, it only internally undergoes the type of cell division called meiosis to directly produce spores. In turn, the spores can be released from the zygote. This is why embryophytes are commonly defined as all the photosynthetic organisms that retain on the parent gametophyte the zygote (unshed) to form an embryo contained, protected, and nourished within an archegonium or seed. In land plants, the embryo is generally enclosed within a protective covering of the sex organ of the mother gametophyte and, therefore, at least initially, the new sporophyte is dependent on the gametophyte. If the embryo is broadly defined as the first stage in the development the sporophyte plant that arises from mitosis of the single, fertilized egg cell called the zygote, it is true that some "algae" (e.g., certain green or brown algae) can be said to produce an embryo. Although the embryo of certain brown algae species in the genus Laminaria remains attached to a gametophyte for a short time, this attachment is only due to very temporary adhesion and not due to enclosure within the relatively small female sex organ or within the tissues of the parent gametophyte. However, the unique, land plant embryo can be distinguished from that of other embryo producing "algae" by the fact that it is enclosed and well protected within the archegonium or seed. Although not conserved (retained) in all members, the protection and nurturing of the embryo within the unique 'female' sex organ called archegonium of the parent gametophyte is an ancestral condition unique to this lineage of land plants. In certain, more derived seed plants, including Gnetum and Welwitschia plus all angiosperms, there has occurred a strong reduction in the number of cells of the 'female' gametophyte, resulting in the loss of the archegonium; and such a reduced 'female' gametophyte is called an embryo sac. Of course, the egg cell is still found among the remaining cells of the embryo sac. Therefore, the embryo (developing from the fertilized egg) is still embedded and to some extent nurtured and protected at least during the early stages of its multicellular development within the tissues of the parent gametophyte. However, unlike the ancestral condition, both the embryo and the often minute parent gametophyte without an archegonium are enclosed, nourished, protected within the seed. Therefore, the embedding of the embryo for nourishment and protection within the tissues of the parent gametophyte can be considered a shared, derived character (or synapomorphic character) that is both ancestral and unique to this lineage of land plants and that has been conserved (retained) in nearly all known members (from the simplest bryophytes and early polysporangiophytes to the most complex flowering seed plants). The production of an embryo of this type is one of the major characters that distinguishes land plants from all "algae"; and this is why these mostly land dwelling plants are more appropriately called embryophytes. Certainly, the majority of the organisms that are considered to be plants by most people fall into this group of land plants, today including what are commonly called liverworts, mosses, and hornworts, as well as the major hierarchical levels of organisms called vascular, seed, and flowering plants.
    The new consensus (as of 2018) from genetic (molecular) evidence is that monophyletic bryophytes are sister to monophyletic vascular plants. Therefore, embryophytes = bryophytes + vascular plants. See preferred genealogical tree for land plants. The newly defined lineage for bryophytes s.l. (including hornworts, liverworts, and mosses) has already been called Bryophyta Schimper s.l., while the bryophytes s.s. (= mosses) have been referred to by the division name Bryophyta Schimper s.s. or in the classification of Chase & Reveal (2009) the subclass Bryidae Engl., 1892. For some time to come, the paraphyletic sequence of liverworts sister to mosses + (hornworts + vascular plants) will probably still be embraced by most authors of current biology textbooks despite increased recently published contrary evidence based on genetic (molecular) analyses supporting monophyletic bryophytes sister to monophyletic vascular plants.
    The grade of presently living photoautotrophic eukaryotes called "algae" [representing independant (not directly related) lineages] can be distinguished from embryophytes by (1) single-celled sex organs (gametangia) or if multicellular, gametes (sex cells, such as sperm and egg) never surrounded by a layer of sterile cells, (2) single-celled sporangia (spore producing organs), and (3) zygotes (fertilized eggs) never developing into multicellular embryos while still within the female sex organs of gametophytes (organisms that produce gametes).

    Embryopsida (land plant) sexual and asexual reproduction, with central sexual life cycle
    (often referred to as a type of sporic life cycle)

    asexual reproduction (e.g., via sprout-like structures called gemmae of gametophyte)
    |
    mitosis
    |

    |
    mitosis
    |
    asexual reproduction (e.g., via gemmae of sporophyte in some vascular plants)

    Endosymbiosis
    All the eukaryote plants or "plant-like organisms" have originated from heterotrophic, biflagellate (bikont) ancestors (with two minute whip-like, tail-like structures for motility) that have become photoautotrophs by initially engulfing other photoautotropic (food producing) prokaryotes or eukaryotes; and these internally living, photoautotrophic (food-producing) organisms have become a permanent part of their host cells. This process called endosymbiosis, where one engulfed species of organisms can live inside the cells of other unrelated, host organisms and become incorporated as a permanent part (an organelle bound by two or three membranes) of their host cells, is touched on more in the section on the plant kingdom. This is a well supported example of an abrupt change, often considered a type of mutation that has involved a considerable amount of horizontal or lateral gene transfer from photoautotroph to heterotrophic host or somewhat from host to photoautotroph. The last or more recent common ancestor of all living eukaryotes descended from a line of organisms that had already incorporated via endosymbiosis an alpha-proteobacteria somewhat related to presently living purple bacteria, enabling almost all of the currently living eukaryotes (besides those that lost this ability) to carry out an oxygen dependent energy producing process called aerobic respiration. The resulting organelle called the mitochondrion is considered the power house of the eukaryotic cell. This has enabled these organisms to more completely extract energy from food in order to carry out various life processes. Like the prior example, this resulted in a bacterial endosymbiont enslaved by a eukaryotic host. Endosymbiosis is made possible (of course, without subsequent digestion of the engulfed organism) by the process called phagocytosis ('cellular eating'), a unique feeding characteristic found only in some eukaryotes, including some presently living "protozoa" and animals. Phagocytosis commonly involves engulfing (by invagination or infolding of the cell membrane) of undigested food particles that are packaged in membrane surrounded sacs large enough to be called vacuoles. In phagocytic organisms or phagocytes that engage in phagocytosis, internal digestion or more specifically intracellular (within cell) digestion takes place, because the food particle is digested (within the cell) often when the vacuole containing the particle fuses with a membrane encircled lysosome, another often smaller, sac-like structure containing the digestive enzymes. It might also be possible (although not yet proven) that endosymbiosis in the case of the mitochondria took place in the absence of phagocytosis by the fusion of prokaryotes (an archaea host with an alpha-proteobacteria). Mitochondria specialize in the oxidation (via the Krebs cycle) of 3-carbon organic acids that remain from the prior, partial oxidation of sugar in the metabolic pathway called glycolysis in the cytoplasm of eukaryotic cells. This provides an energy supply in the form of molecule called ATP that is immediately available for various energy requiring life processes.

    Green Plants (= viridophytes)
    A working classification of green plants (= "green algae" sensu lato plus land plants)
    primarily based on Lewis and Mc Court (2004), with slight modifications.
    A few traditionally defined paraphyletic or polyphyletic group names in quotes are
    provisionally retained until an improved resolution of genealogical relationships is achieved.
    Some names were obtained from http://www.algaebase.org/ and other sources.
    Kingdom: Chlorobionta Jeffrey 1982, emend. Bremer 1985, emend. Lewis and McCourt 2004
    (= viridophytes = Viridiplantae Cavalier-Smith, 1981)
    o Karol, KG., McCourt, RM., Cimino, MT., CF. Delwiche (2001) The Closest Living Relatives of Land Plants, Science Vol. 294, 2351-2353
    o Lewis LA, McCourt RM. (2004) Green algae and the origin of land plants, American Journal of Botany 91: 1535-1556.

    As a division of the kingdom of green plants (= Viridiplantae Cavalier-Smith, 1981), the lineage called streptophytes sensu lato (= Streptophyta Jeffrey 1967, sensu Leliaert et al. 2012) is composed of seven well supported sublineages represented by class names for the groups of organisms commonly called mesostigmatophytes, chlorokybophytes, klebsormidiophytes, charophytes sensu stricto, coleochaetophytes, conjugates, and embryophytes. The major sublineage of phragmoplastophytes (added here and comprised of the streptophytes that potentially undergo cell division via a phragmoplast) include the last four mentioned groups. The embryophytes are distinguished from all "green algae" (including the other streptophytes and phragmoplastophytes besides the embryophytes) by the development of an embryo from the fertilized egg cell (zygote) that is retained and well protected within the female sex organ (archegonium) or seed. See preferred streptophyte tree followed by supporting references. 
    Preferred land plant = embryophyte tree:

        /-------------Marchantiidae (liverworts) |
    /---|                                        |  
/---|   \-------------Bryidae (mosses)           | bryophytes
|   |                                            | 
|   \-----------------Anthocerotidae (hornworts) |
|            
|   /-----------------Lycopodiidae               | lycophytes     |
\---|                                                             | 
    |   /-------------monilophytes               |                | 
    \---|                                       euphyllophytes    | vascular plants 
        |    /--------gymnosperms                |  |             | = tracheophytes
        \----|                                   |  | seed plants |
             \--------Magnoliidae (angiosperms)  |  |             |
    Levisticum - Hill - Genus Name
    Family Apiaceae
    The genus Levisticum Hill is actually more closely related [in the same lineage (Sinodielsia Clade of subfamily Apioideae)] to such Chinese, medicinal species as Angelica sinensis (Oliv.) Diels (= Dong Quai, Danggui, Female ginseng) and Ligusticum sinense 'Chuanxiong' S.H. Qiu, Y.Q. Zeng, K.Y. Pan, Y.C. Tang & J.M. Xu (= Chuang Xiong, Gao Ben).

    Levisticum officinale - Koch - Species Name
    Family Apiaceae
    In Curtin (1965), the white flowered Angelica pinnata S. Watson (= Small-leaf angelica) is possibly confused with the yellow flowered L. officinale, which in Colorado is also called Osha by Spanish speaking people (Bye & Linares, 1986). Although sometimes called Osha (possibly also in New Mexico), L. officinale is not that closely related to Ligusticum porteri. L. officinale is actually more closely related to the Chinese medicinal species Angelica sinensis (Dang gui) and other species in the Sinodielsia Clade (see above). In China, the introduced and widely cultivated Levisticum officinale (native to Europe and Western Asia) is actually used medicinally as a substitute for native Dang gui. For identification purposes, see brief comparisons of Levisticum officinale with Ligusticum porteri.

    Ligusticum - L. - Genus Name
    Family Apiaceae
    Like several other genera of family Apiaceae subfamily Apioideae, the genus "Ligusticum" as traditionally defined is highly polyphyletic [assigned to a minimum of five major (some informally named) lineages, including the Acronema Clade, the Conioselinum chinense Clade, Pyramidoptereae Boiss., Selineae Spreng., and the Sinodielsia Clade (see Downie, et al., 2010)].
    The related medicinal species native to New Mexico that are classified as Ligusticum porteri J.M. Coult. & Rose (= Osha or Chuchupate of western United States & northern Mexico) and Conioselinum scopulorum (A. Gray) J.M. Coult. & Rose (= Rocky Mountain hemlock-parsley of western United States) have been DNA sequenced and found to be members of a lineage informally named the Conioselinum chinense Clade (Downie, et al., 1998; Downie, et al., 2000a; Downie, et al., 2000b; Downie, et al., 2001; Downie, et al., 2010), together with species found elsewhere (Fernandez Prieto & Cires, 2014), such as C. chinense (L.) Britton, Sterns & Poggenb. (= Chinese hemlockparsley in China, Japan, Russia, and North America), Ligusticum canadense (L.) Britton (= Canadian Licorice Root of North America), and the Old Word genera Dethawia Endl. (only from the Pyrenees and Cantabrian Mountains of Europe), Meum Mill. (Europe & northern Africa), Mutellina Wolf (temperate regions in Europe & Asia), Rivasmartinezia Fern. Prieto & Cires (southern Europe on limestone in northwestern part of the Iberian Peninsula), and Trochiscanthes W.D.J. Koch (southern Europe, only from the western Alps and northern Apennines). Because the whole genus "Ligusticum" (as presently and traditionally defined) is so polyphyletic, both L. porteri (Osha) and L. canadense (Canadian Licorice Root) of the Conioselinum chinense Clade may actually be much less closely related to other species classified in the same genus, such as members of the Acronema Clade (Ligusticum scoticum L.), Pyramidoptereae (Ligusticum rhizomaticum Hartvig), Selineae (L. daucoides Franch., L. involucratum Franch., L. mucronatum (Schrenk) Leute, L. oliverianum (H. Boissieu) Shan, L. physospermifolium Albov, L. pteridophyllum Franch. ex Oliv., and L. scapiforme H. Wolff), and the Sinodielsia Clade (L. acuminatum Franch., L. sinense 'Chuanxiong' S.H. Qiu, Y.Q. Zeng, K.Y. Pan, Y.C. Tang & J.M. Xu, L. jeholense (Nakai & Kitag.) Nakai & Kitag., and L. tenuissimum (Nakai) Kitag.). However, both L. porteri (Osha) and its closer relative L. canadense are probably much more closely related to species classified in other genera (besides "Ligusticum"), such as Dethawia, Conioselinum Hoffm. (in part, if just including C. chinense and C. scopulorum), Meum, Mutellina, Rivasmartinezia, and Trochiscanthes.
    Within the Conioselinum chinense Clade (see Fernandez Prieto & Cires, 2014), L. porteri is apparently sister to C. scopulorum and both these species form a lineage sister to the species C. chinense. The lineage comprising the three last mentioned species form a lineage that is apparently nested within a broader lineage also including Dethawia splendens (Lapeyr.) Kerguelen, L. canadense, Meum athamanticum Jacq., and Mutellina purpurea (Poir.) Reduron, Charpin & Pimenov, and Trochiscanthes nodiflora (Vill.) Koch. This broader lineage is apparently sister to Rivasmartinezia Fern. Prieto & Cires. Finally, the Conioselinum chinense Clade appears to be sister to the Acronema Clade, which consists of the type species [namely L. scoticum L. (= Scottish licorice-root)] of the genus "Ligusticum" and L. scoticum L. ssp. hultenii (Fernald) Calder & Roy L. Taylor (= Hulten's licorice-root), together with (at least) the genus Acronema Falc. ex Edgew., species Angelica anomala Ave-Lall., the genus Halosciastrum Koidz., the genus Harrysmithia H. Wolff, the genus Kitagawia Pimenov, the genus Meeboldia H. Wolff, the genus Oreocomopsis Pimenov & Kljuykov, the species Ostericum grosseserratum (Maxim.) Kitag., the species Ostericum scaberulum (Franch.) C.Q. Yuan & R.H. Shan, the genus Pachypleurum Ledeb., the species Pimpinella brachycarpa Nakai, the species Pleurospermum hookeri C.B. Clarke, the species P. yunnanense Franch., the genus Pternopetalum Franch., the genus Pterygopleurum Kitag., the genus Rupiphila Pimenov & Lavrova, the genus Sinocarum H. Wolff ex R.H. Shan & F.T. Pu (including Apium ventricosum H. Boissieu), the genus Spuriopimpinella (H. Boissieu) Kitag., and the genus Tilingia Regel & Tiling.
    [Convergent uses: Within the Conioselinum chinense Clade, the species Dethawia splendens (= Ligusticum splendens Lapeyr.) is endemic to the Pyrenees and the Cantabrian Mountains and forms a strongly supported monophyletic group with the species Meum athamanticum (Fernandez Prieto & Cires, 2014). M. athamanticum (commonly called Spignel in England and probably very closely related to D. splendens) is native to the mountains from west and central Europe and has been cultivated in Scotland as a root vegetable (used like parsnips). Acording to Nicholas Culpeper (1616-1654), Spignel was used to improve both appetite and digestion and to treat excessive flatulence, belching, colic, and stomach aches. Culpeper recommended that the root also be used to regulate menstruation, ease childbirth and to promote the expulsion of the afterbirth. The leaves have been used as a condiment in soups or stews and medicinally (as a stomachic, diuretic, and emmenagogue) for stomach ailments and other conditions (Usher, 1974). The roots of Spignel are known in Europe to be similar in both flavor and medicinal effects to Angelica (= Angelica archangelica Linnaeus) and Lovage (= Levisticum officinale Koch). Trochiscanthes nodiflora (= Ligusticum nodiflorum Vill.) found only in the French Alps, Italian Apennines, and a small section of Switzerland has also been reported by Dragendorff to have been used as a carminative and emmenagogue (See posters from Panta Medica 55, 1989). All these medicinal uses together with the placement of both M. athamanticum and T. nodiflora on the family tree close to L. porteri (Osha) point to the presence of bioactive chemicals called phthalides, several of which have already been identified in both M. athamanticum and T. nodiflora. The species Mutellina purpurea [= Ligusticum mutellina (L.) Crantz; Meum mutellina (L.) Gaertner; Mutellina adonidifolia (J.Gay) Gutermann], which has been used as a stomachic (and the dry leaves as a substitute for parsley and for tea), can be found in the mountains of Central and Southern Europe. The bioactive phthalide called (Z)-Ligustilide has already been identified (Brandt & Schultze, 1995) in the essential oil of this species commonly called Mountain lovage or Alpine lovage. Mutellina caucasica (Sommier & Levier) Lavrova (= Ligusticum caucasicum Sommier & Levier) is a Caucasian endemic. Mutellina corsica (J. Gay) Reduron is endemic to the Corsican Mountains. The newly discoved Spanish species Rivasmartinezia vazquezii Fern. Prieto & Cires has as yet no reported medicinal uses.]
    Although the species C. chinense and C. scopulorum are probably closely related to each other and appear nested within the Conioselinum chinense Clade with L. porteri (Osha), the genus "Conioselinum" Hoffm. as a whole (as traditionally defined) is also polyphyletic with at least one other species (C. tataricum Hoffm.) in another lineage (besides the Conioselinum chinense Clade) called the Sinodielsia Clade (Downie, et al., 2010). Furthermore, the type species of the genus "Ligusticum" or "Conioselinum" is not part of the lineage (the Conioselinum chinense Clade) of the closely related species L. porteri and C. scopulorum. Nevertheless, the names of the polyphyletic "genera" "Ligusticum" and "Conioselinum" continue to be treated here as mostly by other authors in a traditional, conservative way until a more complete knowledge of phylogenetic relationships becomes available. However, the reader should be informed that these "genera" are artificial groups of species whose relationships are not completely understood and future scientific name changes are likely. In the case of "Conioselinum" and "Ligusticum", these "genera" (as currently defined) have various species within and outside the Apioid Superclade within the subfamily Apioideae. The Apioid Superclade (including such major lineages as Pyramidoptereae Boiss., Selineae Spreng., and the Sinodielsia Clade) is nested within an even larger (unnamed) lineage that includes some lineages outside the Apioid Superclade, such as the Acronema Clade and Conioselinum chinense Clade; and all these lineages together with some others are within the subfamily Apioideae.
    The type for the genus name "Ligusticum" L. is represented by a type specimen
    of the species named L. scoticum L. In other words, the genus name "Ligusticum" L.
    can be said to be based on the species named L. scoticum L. The type species of
    "Ligusticum", L. scoticum L., occurs in the lineage called the Acronema Clade.
    The type species for "Conioselinum", C. tataricum Hoffm., occurs in a lineage called
    the Sinodielsia Clade. However, not all species of "Ligusticum" are found in the
    Acronema Clade (see above); and not all species of "Conioselinum" are found in the Sinodielsia
    Clade. Furthermore, the type species of the genus "Ligusticum" or "Conioselinum" is not
    part of the lineage of the closely related L. porteri and C. scopulorum in New Mexico.
    One suggested possibility (Papini & Mosti, 2006) would be to place all species of
    Ligusticum and Conioselinum of the Conioselinum chinense Clade in a genus such as
    Trochiscanthes that presently includes only one European species T. nodiflora (All.) W.D.J. Koch.
    The primarily Arctic Ligusticum scoticum L. (= Scottish licorice-root) can be found in North America from western Greenland to New England. It is also found in northern Norway and the northern British Isles. The related L. scoticum L. ssp. hultenii (Fernald) Calder & Roy L. Taylor (= Hulten's licorice-root), also sometimes referred (by the synonym) as L. hultenii Fernald, can be found in Alaska and Western Canada. It is also found in Asia (Japan). Both L. scoticum and L. hultenii are placed in the lineage called the Acronema Clade; and (as stated above) L. scoticum is designated as the type species of the polyphyletic genus "Ligusticum".
    Other North American species of the genus "Ligusticum" (besides L. canadense, L. porteri, and L. scoticum or L. hultenii) with lineage membership apparently still unknown or uncertain and at least not DNA sequenced by Downie, et al. (2010) include L. apiifolium (Nutt.) A. Gray, L. californium J.M. Coult. & Rose, L. calderi Mathias & Constance, L. canbyi (J.M. Coult. & Rose) J.M. Coult. & Rose, L. filicinum S. Watson, L. grayi (J.M.Coult.) Rose, and L. tenuifolium S. Watson, and L. verticillatum (Hook.) Coult. & Rose ex Rose. Of these other North American species, L. calderi (Calder's lovage, Calder's licorice-root) can be found native as far north as Alaska (?) and Western Canada. Although L. canbyi, L. filicinum, L. grayi, and L. tenuifolium are found outside New Mexico, they are considered medicinal in their region and have often been referred to commercially as 'Osha' (Turi & Murch, 2010), which is a common name that most likely originated in the Upper Rio Grande Valley of New Mexico for the species L. porteri (see intro).
    In the below descriptions of Cicuta maculata Linnaeus, Conioselinum scopulorum, Conium maculatum Linnaeus, Levisticum officinale Koch, and Ligusticum porteri, the highlighted phrases do not necessarily represent shared, derived characters. These highlighted phrases simply represent easily observable, key characteristics that can help to distinguish Ligusticum porteri from the other (above mentioned) species in New Mexico. The species of Cicuta and Conium are also described in this section, because they can superficially resemble C. scopulorum and L. porteri.
    o In New Mexico, Ligusticum porteri has fibrous roots, with a thin, wrinkled, brown (like dark chocolate) outer, skin-like layer that covers the fibrous, yellowish-white, strongly scented (aromatic) inner tissue, and with root crown branched or unbranched and surrounded by numberous, persistent, uniquely reddish tinted leaf bases, the withered petiole (leaf stalk) bases surrounding root crown forming a collar of dead, distinctively hair-like material (remnant, often vertically oriented, stalk fibers); foliage and especially inner root tissue strongely aromatic [the smell very distinctive, due mostly to relatively high concentations of volatile chemicals called phthalides (Leon et al., 2017) similar to those found in some species of "Angelica", "Conioselinum" scopulorum, and Levisticum officinale, but also found in less amounts in celery (Apium graveolens)]; multiply divided (ternate-pinnately) compound leaves with leaflet margins serrate (notched like a saw with small, forwardly pointed, toothlike projections called teeth) or cleft (cut about half-way to middle) and with lateral veins not ending in the sinuses (grooves between the teeth), but directed toward the teeth tips, primary (large, divided) leaves at base of plant usually stalked (lower leaves with often long, well-developed petioles), upper leaf blades arising directly from dilated sheaths (without stalks); flower clusters (inflorescences) flat-topped to dome-shaped, compound umbels [each characterized as a larger (primary) umbrella shaped cluster of smaller (secondary) umbrella shaped flower clusters (sometimes called umbellets), with elongated stalks (called rays) arising more or less from a common point], terminal umbel occasionally solitary, often basally surrounded (subtended) by a lateral whorl of 3-8 (-12) umbels, or more often a pair of opposite umbels; primary (larger) umbel with involucre of one or a whorl of a few bracts, secondary (smaller) umbels subtended by two, rarely three bractlets in a whorl called the involucel, but both involuce and involucel can be absent; small flowers white, styles three (other plants in family often have only two styles), enlarged or expanded base of styles (stylopodium) collectively conical in shape; fruit small, dry, not compressed on back side, but more or less cylindrical and slightly compressed on lateral sides, with two parts (carpels) called mericarps that separate at maturity; the slender prolongation of flower receptacle called a carpophore forms a central axis between the carpels [the carpophore of Ligusticum porteri can split in two all the way to its base with each of the two mericarps (reddish, oblong- to cylindric-shaped ribbed, fruit segments) often still attached to either of its two separated apices].
    The reader should be informed that Conioselinum scopulorum (Rocky Mountain hemlock-parsley) and Ligusticum porteri (Osha) are most frequently confused with each other. They are both strongly aromatic with a similar scent; they have both been used medicinally by Native Americans for some of the same purposes, although they seen to have been considered distinct from one another; and they both may have some of the same medicinal properties due to a relatively close genetic relationship within the same lineage. One reason why C. scopulorum is not more commonly confused with L. porteri is that people tend to shy away from wet places (e.g., wet meadows and streamsides) where the former species often grows; and in such places this species is often unnoticed because it is often surrounded by tall grasses and sedges. Nevertheless, when viewed up close, the compound umbels of C. scopulorum are often beautiful clusters of tiny, bright white flowers; and (unlike L. porteri) the flower buds of the umbels are distinctively tinged with pink. However, both C. scopulorum and L. porteri can also potentially be confused with species of the poisonous, less aromatic genera Cicuta L. and Conium L. due to superfical similarity in appearance. The poisonous species Cicuta maculata L. and Conium maculatum L. are also found in the mountains in wet places, where C. scopulorum and sometimes even L. porteri can also be found. Therefore, when making plans to try to find the C. scopulorum or L. porteri in the wild, the reader should also know how to distinguish these plants (especially in wet places) from Cicuta and Conium (see below).
    o Conioselinum scopulorum, although aromatic like its close relative L. porteri, can be distinguished by a presence more commonly along streamsides, seeps, and wet meadows in mostly northern mountains (extending southward to the Sacramento Mountains, but unknown in extreme southwest New Mexico); crown of taproot usually unbranched (simple) with few if any persistent fibrous leaf bases [without persistent petiole (leaf stalk) bases at root crown or with only a few withered but weakly persistent leaf bases not distinctively fibrous]; ultimate segments of leaves not as conspicuously veined, irregularly dissected or divided into numerous segments, distinctive leaflets often not as clearly obvious, lower leaves pinnate or ternately-pinnate with stalks, upper ones pinnate or pinnately lobed (in a feather-like fashion) and without stalks or nearly without them on dilated sheaths; terminal umbel solitary or above alternate lateral umbels and usually never above a pair of opposite or a lateral whorl of umbels, stalks (rays) of compound umbels on average shorter, only one or few bracts of involucre or entirely absent, but with 3 or usually more (often 6), more conspicuous, relatively long (2-8 mm), linear, pointed bracklets of involucel, often projecting out beyond the flower clusters, individual flowers brighter white and flower buds distinctively tinged with pink; dorsally flattened mericarps (fruit) not cylindrical.
    o The introduced European species Levisticum officinale (= Lovage in gardens in New Mexico and elsewhere) has also sometimes been referred to as 'Osha' among the some Spanish speaking people in Colorado (Bye & Linares, 1986). Although in the same family and subfamily and with a similar aromatic scent, the genus Levisticum Hill is not found in the same lineage as the closely related Conioselinum scopulorum and Ligusticum porteri (see above). Levisticum officinale (often grown from seeds) is sometimes found to escape from nearby gardens. The leaflets are lobed or cleft in Levisticum but toothed or cleft in Ligusticum porteri. The involucre (whorl of bracts at base of primary umbel) is prominent in Levisticum and more or less absent in Ligusticum porteri. The flowers of Levisticum are yellow or light green but in the more closely related Conioselinum scopulorum and Ligusticum porteri they are white. All ribs of fruit of Levisticum and some (not all) fruits of Ligusticum porteri are winged (with thin and flat, bordering and extending margins); fruits are not much compressed in both of the more closely related species; and some plants of Ligusticum porteri have fruits without any wings or only scarcely winged.
    The reader that is interested in wild crafting should also learn to distinguish the medicinal and food or spice plants of the family Apiaceae from a few poisonous plants that may superficially resemble them.
    o The species Cicuta maculata (= Water-hemlock) can be distinguished as growing in mountains along streamsides, pond and lake margins, marches, canals, ditches, and wet meadows; perennials with thick rootstocts bearing a series of cross-partitions that separate cavities and partitions that can be observed close to ground level; not that aromatic; stems sometimes with purple stripes; lower leaves once-pinnate or ternately-pinnate compound with well-defined (wholly separate) leaflets; upper reduced leaves less dissected, once-pinnate or even simple (without leaflets), often without stalks on dilated sheaths; leaflet margins serrate [with only forwardly pointed, teeth-like margins, never cleft or entire (with smooth, uninterrupted margins)], lateral veins of the leaflets tending to end in the sinus (notches) between the teeth; involucre absent or one or a whorl of few bracts, involucels with usually 6 bractlets.
    o The species Conium maculatum (Poison hemlock) is much more common in New Mexico than the native, perennial Cicuta. Poison hemlock can be distinguished as a introduced, much-branched, biennial that is found growing along streams, rivers, moist roadsides, fence lines, ditches, moist (weedy) waste places, wet meadows, and wet low ground; stems spotted with brownish or reddish-brown to purple dots; not that aromatic; crown of rootstock with only a few or no cross-partitions that separate cavities; leaves irregularly dissected or divided into numerous segments (like Conioselinum or Wild carrot), distinctive leaflets often not that clearly obvious, lower leaves pinnate or ternately-pinnate, reduced upper leaves once pinnate or pinnatifid (lobed in a feather-like fashion) without stalks on dilated sheaths; many umbels (10-30 or more), bracts of involucre 2-6, bracklets of involucels 4-6.

  • Brandt JJ & Schultze W (1995) Composition of the essential oils of Ligusticum mutellina (L.) Crantz (Apiaceae). J Essent Oil Res 7:231
  • Downie, S.R., Ramanath, S., Katz-Downie, D.S. & Llanas, E. (1998) Molecular systematics of Apiaceae subfamily Apioideae: phylogenetic analyses of nuclear ribosomal DNA internal transcribed spacer and plastid rpoC1 sequences. Amer. J. Bot. 85: 563-591.
  • Downie, S.R., Katz-Downie, D.S. & Watson, M.F. (2000a) A phylogeny of the flowering plant family Apiaceae based on chloroplast DNA rpl16 and rpoC1 intron sequences: towards a suprageneric classification of subfamily Apioideae. Amer. J. Bot. 87: 273-292
  • Downie, S.R., Watson, M.F., Spalik, K. & Katz-Downie, D.S. (2000b) Molecular systematics of Old World Apioideae (Apiaceae): relationships among some members of tribe Peucedaneae sensu lato, the placement of several island-endemic species, and resolution within the apioid superclade. Can. J. Bot. 78: 506-528.
  • Downie, S.R., Plunkett, G.M., Watson, M.F., Spalik, K., Katz-Downie, D.S., Valiejo-Roman, C.M., Terentieva, E.I., Troitsky, A.V., Lee, B.-Y., Lahham, J. & El-Oqlah, A. (2001) Tribes and clades within Apiaceae subfamily Apioideae: the contribution of molecular data. Edinb. J. Bot. 58: 301-330.
  • Downie S.R., Spalik K., Katz-Downie D.S. & Reduron J.P. (2010) Major clades within Apiaceae subfamily Apioideae as inferred by phylogenetic analysis of nrDNA ITS sequences. Pl. Diversity Evol. 128: 111-136.
  • Fernandez Prieto, J.A., & Cires, E. (2014) Phylogenetic placement of Dethawia, Meum, and Rivasmartinezia (Apioideae, Apiaceae): Evidence from nuclear and plastid DNA sequences. Plant Biosystems-An International Journal Dealing with all Aspects of Plant Biology, 148(5), 975-987.
  • Leon, A., Del-Angel, M., Avila, J. L., & Delgado, G. (2017) Phthalides: distribution in nature, chemical reactivity, synthesis, and biological activity. In Progress in the Chemistry of Organic Natural Products 104 (pp. 127-246). Springer, Cham.
  • Papini, A., & Mosti, S. (2006) Notes on Trochiscanthes Koch (Apiaceae) on the basis of ITS rDNA sequence. Webbia, 61(2), 217-225.
  • Turi, C., & Murch, S.J. (2010) The genus Ligusticum in North America: An ethnobotanical review with special emphasis upon species commercially known as 'Osha'. Herbal Gram, 89, 40-51.

    Ligusticum porteri - Coulter & Rose - Species Name
    Family Apiaceae
    According to Bye & Linares (1986), Chuchupate is possibly based on the modification by early Spanish settlers of a Nahuatl term such as Chichipatli (meaning in English 'dog medicinal herb').
    In southern New Mexico, Chuchupate and Hierba de cochino can both apply to L. porteri.

  • Bye, R.A., & Linares, E. (1986) Ethnobotanical notes from the valley of San Luis, Colorado. Journal of Ethnobiology, 6, 289-306.

    Lithospermum onosmodium - J. Cohen - Species Name
    Family Boraginaceae
    Onosmodium molle Michaux (= Onosmodium occidentale Mackenzie) is now formally named Lithospermum onosmodium J. Cohen (also found wild in moist areas in northeastern plains of New Mexico), because according to molecular (DNA) evidence (Weigend et al. 2009; Cohen & Davis 2009), recognition of Onosmodium at the generic rank renders the genus Lithospermum paraphyletic. Even though plants traditionally placed in the genus Onosmodium form a monophyletic group sister to the southeastern North American species Lithospermum tuberosum Rugel, Onosmodium is deeply nested within the genus Lithospermum. The species Lithospermum onosmodium is reported to been used by Native Americans externally for swellings, lumbago, numb skin, and as a rubbing solution for horses (Moerman).

  • Cohen, J.I. and J.I. Davis (2009) Nomenclatural changes in Lithospermum (Boraginaceae) and related taxa following a reassessment of phylogenetic relationships. Brittonia 61:101-111.
  • Weigend, M., M. Gottschling, F. Selvi, and H.H. Hilger (2009) Marbleseeds are gromwells: systematics and evolution of Lithospermum and allies (Boraginaceae tribe Lithospermeae) based on molecular and morphological data. Molec. Phylogen. Evol. 52:755-768.

    Phytolaccaceae - R.Br. - Family Name
    Caryophyllales - Juss. ex Bercht. & J.Presl - Order Name
    The family Phytolaccaceae s.l., as often broadly defined traditionally to include the subfamily Rivinoideae Nowicke [= family Petiveriaceae Meissner (genera: Gallesia Casar., Hilleria Vellozo, Ledenbergia Moquin, Monococcus F. Mueller, Petiveria L., Rivina L., Schindleria H. Walter, Seguieria Loefling, and Trichostigma A. Richard)], as well as subfamily Phytolaccoideae Arnott [= Phytolaccaceae s.s. (genera: Anisomeria D. Don, Ercilla A. de Jussieu, Nowickea Martinez G. & McDonald, and Phytolacca L.)] and the subfamily Agdestidoideae Nowicke (1 genus: Agdestis Sesse & Mocino ex de Candolle of Florida, Texas, West Indies, Mexico, and Guatemala to Nicaragua), is almost certainly polyphyletic.
    The separately defined family Petiveriaceae (= Rivinoideae) may be more closely related to the family Nyctaginaceae Juss. and Rivinoideae should probably be removed from Phytolaccaceae. Therefore, the genus Rivina should more likely be placed in the separate family Petiveriaceae outside the family Phytolaccaceae and closer to the family Nyctaginaceae.
    Although less certain, it is possible on the basis of some DNA evidence that the genus Sarcobatus Nees (= Greasewood) should also be considered a member of Phytolaccaceae. This genus with 1 species Sarcobatus vermiculatus (Hooker) Torrey in New Mexico is often traditionally placed in the family Chenopodiaceae Vent. and more recently in the separate family Sarcobataceae Behnke.
    The genus Agdestis (1 species: A. clematidea Sesse & Mocino ex de Candolle) was recovered as sister to Sarcobatus with moderate support in the analyses of Cuenoud et al. (2002), Schaferhoff et al. (2009), and Brockington et al. (2011).
    References:
    o Brockington SF, Walker RH, Glover BJ, Soltis PS, Soltis DE. (2011) Complex
    pigment evolution in the Caryophyllales, New Phytol. 190(4):854-864.
    o Cuenoud P, Savolainen V, Chatrou LW, Powell M, Grayer RJ, Chase MW. (2002)
    Molecular phylogenetics of Caryophyllales based on nuclear 18S rDNA and
    plastid rbcL, atpB, and matK DNA sequences, Am J Bot. 89(1):132-144.
    o Schaferhoff B, Muller KF, Borsch T. (2009) Caryophyllales phylogenetics:
    disentangling Phytolaccaceae and Molluginaceae and description of
    Microteaceae as a new isolated family, Willdenowia 39(2):209-228.

    Pinnate leaves
    A pinnate leaf (or once-pinnate leaf) is considered a single compound leaf made up of smaller leaflets arranged on both sides of an axis in a feather-like fashion (like many fern fronds). Therefore, a bipinnate leaf is made up in a feather-like arrangment of smaller leaflets that are made up in the same arrangment of even smaller leaflets; and a ternately-pinnate leaf (a single, triple-pinnately compound or ternate-pinnately compound or tripinnate leaf) is made up in a feather-like arrangment of smaller leaflets, that are made up in the same arrangment of smaller leaflets, that are made up in the same arrangment of even smaller leaflets. The simple leaves that are not fully dissected into seperate leaflets can sometimes be lobed in a pinnate (feather-like) fashion. Such leaves are considered pinnatifid or simply pinnately lobed. Some leaves that have pinnately arranged lobes with incisions that extend almost up to the central axis (called the midrib) are considered pinnatisect.

    Plant Identification
    Identification is the process of matching the characters of an unknown organism with those of a known (previously described) one. In this matching process, it is the features that distinguish the unknown organism from all other organisms that lead to diagnosis or diagnostic characterization. Books (manuals) cataloging the plant species of a specific location called regional floras are most commonly used for the purpose of identification, because they provide all the required, possible descriptions and diagnostic features (including keys) for native or naturalized plants. Naturalized plants are those that are initially introduced from some other geographic region but now are found growing wild. In identification, it is important to at least have the capability to check all (or most) of the possibilities for a given region. Although there is probably no regional flora that is 100% complete, these manuals are extensive enough to be useful most of the time. However, when using a manual for a regional flora, it is important to be aware of the geographical range of this reference to be sure that it is likely to include the unknown plant. Many regional floras include only the native or naturalized vascular plants; some may rarely also include other wild land plants of the region, such as the bryophytes (liverworts, mosses, and hornworts), but usually never the lichens (a separate reference is usually needed for them). Therefore, it is important to determine whether the regional flora includes the major taxonomic group of the organism of interest to be identified. If complete identification (to the level of species or below) is not possible by using a regional flora, the unknown organism can often be at least placed in some higher level group, such as a family or genus. Once identification is pursued to the lowest possible level, it is important to recognize the reasons why the unknown organism does not closely fit any of the lower level descriptions. More commonly, the unknown may be an introduced species (not native or naturalized) in the geographical region of the organisms available for comparison. Only rarely will the unknown be found to be a native or wild species undiscovered or not yet included in the regional flora.
    For the purpose of identification of organisms, keys (usually included in manuals for regional floras) are constructed as a hierarchy of two or more contrasting sequences of characters, each sequence with two or more contrasting sequences of characters beneath it. Such keys are applied to identification by choosing one sequence of contrasting characters that best fits the organism versus other such sequences of characters and doing the same for sequences of contrasting characters beneath the one chosen. This is repeated until the combination of characters with the best fit in the hierarchy terminates with some named group of organisms.
    Plant descriptions
    The descriptions useful to plant identification emphasize easily observable characters like those found in the keys. However, the well-defined characters employed to help characterize natural groups (= lineages) may not always be that easy to observe in the wild (in the field). Although the easily observable morphology of flowers (= floral morphology, such as number and degree of fusion, including arrangement or number of whorls, of flower segments) is a difficult subject when used to characterize broader lineages, it, together with the characteristics of the subsequently developing fruit, can often be used to disclose broad trends in the evolution of the flower and in turn help to more or less distinguish lineages of flowering plants. The same can be said for the details of cone-like structures in non-flowering vascular plants. Therefore, these characters, even though sometimes less useful in defining broader lineages, will often be described in much detail, because, when present, they are easy to observe in the field. These are the characters most easily used to identify plants (like those found in the keys and descriptions of regional floras). However, such easily observable characters may or may not be that useful in diagnosing broader lineages. Often it is the less obvious and somewhat cryptic characters (as many shared, derived characters) that are in many cases more useful in distinguishing the broader lineages (see Cantino, et al., 2007). Nevertheless, it is the descriptive terminology for the more easily observable characters that is most often used in plant identification and description.
    The learning of this descriptive terminology of plants called phytography is like learning a new language and it enables the reader to create a word picture of the plants described. Once the reader becomes familiar with this terminology, it also makes it possible (for some people) to more or less mentally picture a plant without the use of a drawing or photograph. In addition to describing them more precisely, this descriptive terminology can also help one to observe plants in the field more critically, making possible more certain identification.

    Plastid
    A plastid (e.g., the chloroplast) is a distinct small structure (organelle) found in the cytoplasm of cells of eukaryotes called photoautotrophs; it is bound by at least a two membranes and functions in manufacture or storage of food or other important chemical compounds. Plastids in land plant include chloroplasts that act as centers within the cell for photosynthesis and closely related structures that can act as nonphotosynthetic sites for the storage of various materials. The major plastids in land plants include proplastids (undifferentiated plastids mostly present in the meristematic regions), which can differentiate into chloroplasts (for photosynthesis), etioplasts (immature chloroplasts, when non-pigmented, also considered leucoplasts, or chloroplasts not exposed to light), chromoplasts (for pigment synthesis and storage), and non-pigmented leucoplasts (for monoterpene synthesis or production of various compounds, such as tetrapyrroles like haem). The leucoplasts can differentiate into amyloplasts (for starch storage), statoliths (for detecting gravity), elaioplasts (for storing fat), and proteinoplasts (for storing and modifying protein). Chloroplast and leucoplasts (leukoplasts) can occur in algae, but amyloplasts, chromoplast, and etioplasts, although found in land plants, are absent in algae. Plastids can also be referred to as cyanelles (in the glaucophytes), rhodoplasts (in red algae), and chloroplasts (in green algae and land plants). Those plastids with two membranes (originating from the two membranes of gram-negative bacteria) are the result of primary endosymbiosis of cyanobacteria, while those choroplasts or rhodoplasts with more than two membranes originate from secondary endosymbiosis of mostly green algae and red algae, respectively. The two membranes surrounding a plastid is an indication that the endosymbiont was a prokaryote, while three or four membranes surrounding a plastids is an indication that the endosymbiont was a eukaryote already with its own plastids. The plasma membrane of the eukaryote endosymbiont often immediately surrounds the two inner membranes from the prokaryote. In obligate endosymbiosis, neither the endosymbiont nor the host can survive without the other. Check out definition for endosymbiosis, which (if symbiogenesis takes place) can be considered a type of mutation. In the alveolates of the proposed supergroup called Chromalveolata (now often consided the expanded Chromista supergroup), the so-called apicoplast of some apicomplexans is no longer capable of photosynthesis, but it is still an essential or functional structure within the cell. The kleptoplastid of some dinoflagellates (also alveolates) is a plastid of a ingested alga that is kept functional for the purpose of photosynthesis, but after some time this plastid is also digested. In the chromists (cryptophytes, haptophytes, and stramenopiles) of Chromista, the plastids have four bounding membranes, the outer one usually confluent with the envelope of the nucleus. In the alveolates of Chromista, the ciliates lack plastids and the rest, the apicomplexans and dinoflagellates, include both plastid-less and plastid-bearing members. In most of the dinoflagellates that possess plastids, peridinin is the major carotenoid pigment of the plastids that are generally enclosed by three membranes. The apicoplasts in the majority of the apicomplexans are exclusively non-photosynthetic and are bound by 2–4 membranes. Plastids related to the apicoplasts with four membranes have recently been identified in a photosynthetic relative of the apicomplexans. Unlike the chromists, the outermost membrane of alveolate plastids is not connected to the membrane of the nucleus. The plastids of chromists and most aveolates originate from red algae endosymbionts. In chlorarachniophytes, the plastids are surrounded by four membranes, including the two inner membranes arising from the inner and outer membranes of the cyanobacterial endosymbiont of some green alga, the third membrane from the engulfed green alga plasma membrane, and the outer membrane from the food vacuole of the final engulfing organism. The engulfed green algal eukaryote (that has become the plastid of chlorarachniophytes) still carries out photosynthesis with its plastids and still contains its own much reduced nucleus called a nucleomorph. In eugenids, the plastids are surrounded by three membranes and appear to have originated (like in chlorarachniophytes) from some green alga endosymbiont. The photosynthetic secondary plastids of these organisms are often generally also referred to as chloroplasts. The organisms with all plastids surrounded by two membranes comprise a proposed lineage called Archaeplastida (= Plantae sensu lato = Primoplantae). This lineage is supported by genetic studies indicating that plastids probably had a single origin. Chloroplasts assume different shapes in various groups of plants, including those that are discoid (few red algae, Chara, and land plants), plate-like (Mougeotia), stellate (some red alge and desmids), girdle-shaped, near cell periphery and folding inwardly (Ulothrix), spiral ribbon shaped (Spirogyra), and cup-shaped (Chlamydomonas). The discoid (disc-shaped) chloroplasts are somewhat unique, because they are the most common type found in land plants and Chara (in only a few red algae and rare elsewhere). Plastids only occur in certain ancestrally biflagellate (bikont) lineages. See also more on endosymbiosis (involving eukaryotic phagocytosis or possibly prokaryotic cell fusion), and first eukaryotic photoautotrophs.

    'Polysporangiophytes'
    'Polysporangiophytes' (as the name implies) are land plants with sporophytes bearing multiple sporangia. Until recently, it was widely assumed that liverworts are sister to mosses + the rest of the land plants (hornworts + ('polysporangiophytes' (including stem group 'polysporangiophytes' + vascular plants))). Therefore, the 'polysporangiophytes' have been assumed to include a paraphyletic series (grade) of stem group 'polysporangiophytes' (with or without a well developed water and food conducting tissue) basal to the lineage of presently living vascular plants (with a well developed water and food conducting tissue). Although the 'polysporangiophyte' character described as 'branched sporophyte plants with multiple spore sacs' (correlated with eventual 'independence of sporophyte in early development from the gametophyte') appears easy enough to understand, this is actually a 'cryptic' character, because (among other reasons) there is presently no way of telling for sure whether the 'unbranched condition' with a single spore sac (sporangium) of liverworts, mosses, and hornworts led to the branched condition with multiple spore sacs (sporangia) of 'polysporangiophytes' or the other way around. This is where not yet discovered or re-interpretation of known fossil evidence could be helpful. Puttick et al. (2018) provides a few further comments on some of these issues. Although these authors narrowed down the currently best supported, alternative relationships between liverworts, mosses, hornworts, and vascular plants, it was still uncertain from their results whether bryophytes are monophyletic or paraphyletic. The reasonably well-supported results included (1) hornworts sister to setaphytes (liverworts + mosses), rendering bryophytes s.l. monophyletic and sister to vascular plants, (2) setaphytes (liverworts + mosses) sister to hornworts + vascular plants, and (3) hornworts sister to setaphytes (liverworts + mosses) + vascular plants. The correct placement of hornwort (whether they are sister to liverworts + mosses or whether they are sister to the rest of the land plants or sister to the vascular plants) remained an open question. Nevertheless, (1) hornworts sister to setaphytes (rendering bryophytes monophyletic and sister to vascular plants) was one of the best-supported results of the molecular (transcriptomic) analyses of these authors. Furthermore, all the analyses of disclosed a well supported lineage, referred to by them as setaphyta, including liverworts sister to mosses, and which appears to possess, as a possible shared, derived character, a similar, 'unbranched' stalk-like structure (seta) that in the spore bearing plant (sporophyte) helps to elevate a solitary spore capsule (sporangium) above the dominant gametophyte for dispersal of the spores by mostly wind or rarely by water. This lineage was probably first proposed on the basis of phylogenetic analysis of the development of male gametes (sperm) by Garbary, Renzaglia, & Duckett (1993). The common name 'setophytes' for this lineage was already introduced by Renzaglia & Garbary (2001). Furthermore, one summary phylogenetic tree of Renzaglia & Garbary (2001) supported bryophytes as monophyletic and sister to vascular plants. A few additional references supporting monophyletic bryophytes sister to vascular plants are also listed below.
    Is it possible that some ancestral line of early 'polysporangiophytes' may have given rise to bryophytes (today collectively represented by liverworts, mosses, and hornworts) as well as vascular plants (today collectively represented by lycophytes and euphyllophytes)? If so, the bryophytes may have become secondarily reduced or more simple in biochemistry and structure [already suggested by Raven (2000)]. In this case, the persistent dependence on the mother gametophyte (obligate matrotrophy) of a reduced sporophyte (with a single sporangium) could be a derived innovation unique to bryophytes. According to Cox (2018), hornworts sister to liverworts + mosses, rendering bryophytes monophyletic and sister to vascular plants, is the better supported hypothesis, with the next in line, less likely hypothesis involving liverworts + mosses branching first within land plants and hornworts sister to vascular plants. If the first hypothesis continues to gain support in the future, it is possible that the ancestor of both bryophytes and vascular plants may have possessed alternation of generations involving an equally dominant (isomorphic) sporophyte and gametophyte of similar size and appearance and even a relatively well developed water and food conducting (vascular) tissue. In comparison to such a common ancestor, the gametophyte of bryophytes may have successively become larger and more complex, while the sporophyte may have successively become more reduced in size and complexity. The opposite trend in the lineage that led to vascular plants could have resulted in successively smaller, less complex gametophytes and larger, more complex sporophytes. If bryophytes are indeed monophyletic, the possiblity arises that 'polysporangiophytes' may not all form a monophyletic group that together with vascular plants is sister to bryophytes. Therefore, the placement of some early 'polysporangiophytes' (including known fossils) on the tree of life becomes uncertain. Some early (now extinct) 'polysporangiophytes' could be paraphyletic with respect to bryophytes, vascular plants, or both. However, since the currently best supported genealogical trees on the basis of genetic (molecular) evidence now feature monophyletic bryophytes s.l. sister to monophyletic vascular plants, then the alternation of generations of the common ancestor of all embryophytes cannot currently be (for sure) reconstructed, because the life cycles that involve gametophyte dominance and sporophyte ­dominance, as well as more or less equal dominance of gametophyte and sporophyte, are all possible ancestral states (Gitzendanner et al., 2018; Kenrick, 2017). Although the earliest, undisputed body fossils (other than spores) of land dwelling plants appear to have branched sporophytes, this is not to say that a dominant gametophyte bearing an unbranched sporophyte could not be representative of the earliest character state in embryophytes. The reconstructed genealogy on the basis of form and structure (morphology) of the earliest fossils of 'polysporangiophytes' even suggests that early sporophytes were unbranched with a single, terminal sporangia like those of mosses (Szovenyi, Waller, & Kirbis, 2018). However, the sporophyte (whether unbranched or branched) of the common ancestor of embryophytes could have been much more complex in morphology than that of the present day bryophytes.

  • Budke, J. M., Bernard, E. C., Gray, D. J., Huttunen, S., Piechulla, B., & Trigiano, R. N. (2018) Introduction to the Special Issue on Bryophytes. Critical Reviews in Plant Sciences, 1-11.
  • Cox, C. J. (2018) Land plant molecular phylogenetics: a review with comments on evaluating incongruence among phylogenies. Critical Reviews in Plant Sciences, 1-15.
  • de Sousa, F., Foster, P. G., Donoghue, P. C., Schneider, H., & Cox, C. J. (2018) Nuclear protein phylogenies support the monophyly of the three bryophyte groups (Bryophyta Schimp.). New Phytologist.
  • Garbary, D. J., Renzaglia, K. S., & Duckett, J. G. (1993) The phylogeny of land plants: a cladistic analysis based on male gametogenesis. Plant Systematics and Evolution, 188(3-4), 237-269.
  • Gitzendanner, M. A., Soltis, P. S., Wong, G. K.-S., Ruhfel, B. R., & Soltis, D. E. (2018) Plastid phylogenomic analysis of green plants: A billion years of evolutionary history. American J. Bot. 105: 291-301.
  • Kenrick, P. (2017) Changing expressions: A hypothesis for the origin of the vascular plant life cycle. Phil. Trans. Royal Soc. B 373:20170149.
  • Morris, J.L., Puttick, M.N., Clark, J.W., Edwards, D., Kenrick, P., Pressel, S., Wellman, C.H., Yang, Z., Schneider, H. & Donoghue, P.C.J. (2018) The timescale of early land plant evolution. Proceedings of the National Academy of Sciences, USA 115: E2274-E2283.
  • Puttick, M. N., Morris, J. L., Williams, T A., Cox, C.J., Edwards, D., Kenrick, P., Pressel, S., Wellman, C. H., Schneider, H., Pisani, D., & Donoghue, P. C. J. (2018) The interrelationships of land plants and the nature of the ancestral embryophyte. Curr. Biol. 28: 733-745
  • Raven, J.A. (2000) Land plant biochemistry. Philosoph. Trans. Roy. Soc. London B 355: 833-846.
  • Rensing, S.A. (2018) Plant evolution: phylogenetic relationships between the earliest land plants. Current Biology 28: R210-R213.
  • Renzaglia, K.S. & Garbary, D.J. (2001) Motile gametes of land plants: diversity, development, and evolution. Critical Reviews in Plant Sciences 20: 107-213.
  • Renzaglia, K.S., Aguilar, J.C.V., & Garbary, D.J. (2018) Morphology supports the setaphyte hypothesis: mosses plus liverworts form a natural group. Bryophyte Diversity and Evolution, 40(2), 11-17.
  • Szovenyi, P., Waller, M., & Kirbis, A. (2018) Evolution of the plant body plan. Current topics in developmental biology, 131, 1-34.
  • Wickett, N. J., Mirarab, S., Nguyen, N., Warnow, T., Carpenter, E., Matasci, N., ... & Ruhfel, B. R. (2014) Phylotranscriptomic analysis of the origin and early diversification of land plants. Proceedings of the National Academy of Sciences, 111(45), E4859-E4868.

    Seed
    The seed (in terms of its most important features) is simply a propagule (a structure with the capacity to give rise to a new individual and used as a means of dispersal (spread of a species to a new location) with an embryo packaged along with stored food within at least one resistant coat (called the seed coat or testa, developing from the one or two integuments after fertilization). The seed coat is often considered synonomous with the testa. However, in some flowering seed plants, the part of the seed coat that develops from the outer integument is more strictly called the testa, while the part that develops from the inner integument (if present) is called the tegmen.
    The ovule (unfertilized pre-seed) comprises an indehiscent megasporangium (= non-splitting, relatively large, 'female' sporangium, more appropriately called a megasporangium) surrounded by one or two sheath-like structure(s) called integument(s) that in presently living seed plants possesses only at a single small sperm or pollen tube receptive opening called the micropyle. Furthermore, these sheathed (integumented), indehiscent (non-splitting) megasporangia called ovules each retain only one functional, relatively large 'female' spore (more commonly called a megaspore), which contains within its spore-like wall an internally developing 'female' gametophyte (more commonly called a megagametophyte). Upon contact with pollen, it is the ovules that can develop into seeds. Aside from the very rare exception of pre-fertilization dispersal, the seed is a matured ovule that (prior to its dispersal) contains an internally developed megagametophyte that has already been (via pollen generated sperm) fertilized, and some embryo development has already taken place within the megagametophyte tissue.
    Some basic points of distinction between free-sporing and seed plant reproduction need to be emphasized before the sometimes subtle nature of the seed can be further clarified. Since spores germinate to develop directly into gametophytes, this process can be referred to as spore development (spore germination + gametophyte development). Free-sporing plants include those that shed their spores before or even after this development as long as the gametophytes are independent of the sporophytes prior to fertilization. Even in plants with sporophytes attached and dependent on a more dominant gametophyte, new and separate gametophytes result from spore germination at usually some distance from the combined sporophyte + gametophyte of the previous generation. Even zygospores of "algae" (spore-like zygotes that could be considered analogous to sporophytes) germinate and liberate their spores (usually separate from the mother gametophyte); and the spores germinate independently of each other to produce separate gametophytes. All plants with homosporous reproduction are free-sporing, because they shed their spores from the zygote or sporangium sporophyte before spore development, so the gametophytes are always free-living or independent of the zygote or sporophyte prior to the fertilization event. Only certain free-sporing plants with heterosporous reproduction are known to shed spores after their development but before the ferilization event. This is possible in these plants, because gemetophyte development takes place within the spores before they are shed from the sporangium with fertilization taking place after spore release. There have been claims in some text books that certain species of Selaginella (spikemosses) can simulate an early step in the evolution of the seed by liberating their spores from megasporangia after fertilization and development of the embryo. These claims come from the observation that in these species (e.g., Selaginella rupestris), young plants can emerge from the female regions of sporangia bearing structures. It has been thought that the microspores can get caught between leaves that bear (at the base of their upper surface) megasporangia, allowing fetilization of an egg cell to take place before the megaspore is shed. However, these claims have been shown to be invalid, because the species under consideration have been found to be apogamous, meaning that the megaspores can produce embryos directly without fertilization. It is true that certain heterosporous, non-seed plant species can retain the megaspore within the megasporanium for a long enough time to allow the internal (endosporic) development of the megagametophytes within the megaspore walls. However, as far as it is known, the living land plants, except for the seed plants, all eventually shed their spores prior to fertilization irrespective of whether each spore gives rise to a gametophyte prior to or after its release. Because the spores can be said to be free-living before fertilation takes place, the gametophytes can also be said to be free-living; and, therefore, the plants involved are free-sporing. However, seed plants (also with heterosporous reproduction) are not free-sporing, because although the matured microspores (pollen grains) with internal microgametophytes are released from the microsporangia, the megaspores (also with internal but more elaborate megagametophytes) are not released and fertilzation takes place within megasporangia. This because seed plants possess a non-splitting megasporangium that retains a single functional megaspore. The basic distinctions outlined so far do not completely solve the mystery of the seed, but they help get somewhat beyond the free-sporing plant stage. Another point that should be added is that microgametophytes tend to become more dependent on megaspore bearing sporophyte tissue prior to their fertilization by sperm of the egg of the megagametophyte. For example, nourishment for the developing microgametophyte can be derived from its digestion of tissues of the megasporangium (tissues that are produced by the sporophyte). What comes next is the appreciation that after fertilization, the zygote develops into a differentiating embryo not only within the reproductive organ of the female gametophyte but also within the megasporangium of the sporophyte. Nutrition for the developing embryo can now come not only from the megagametophyte but also from the megasporangium. Since the embryo is very important (like an unborn baby), an additional (sheath-like and protective) integument (with, don't forget, a small opening or pollen tube receptive micropyle) has already (prior to fertilization) developed from the sporophyte (besides the more interior and also protective spore wall) as an outer envelope-like structure around the megasporangium. The baby-like embryo (a new sporophyte) inside the mother-like gametophyte is found inside the wall of a single megaspore, which is enclosed within a sheathed megasporangium! This entire structure (baby-like embryo inside mother-like gametophyte inside wall of spore inside sporangium inside integument) develops into the seed. Therefore, the seed can be described as a sheathed (integumented) megasporangium enclosing a single megaspore with internal development that is already fertilized with a differentiated embryo. When ready, it is this entire structure that is shed and not just the unfertilized megaspore with some internal development, as in some free-sporing, heterosporous plants. From this discussion, it should be easier to understand that there is a big difference between free-sporing and seed plant reproduction.

    Seed plants (= spermatophytes)
    Seed plants generate via meiosis many 'male' spores (more appropriately called microspores) that germinate to produce internally (within their walls) tiny, partially developed 'male' gametophytes (more appropriately called microgametophytes). These tiny spore-like containers of partially developed 'male' gametophytes made up of only a few cells are referred to as pollen, which can be shed from an initially enclosing pollen sac ('male' sporangium, more appropriately called a microsporangium) of the sporophyte. In a different location, seed plants generate via meiosis only one, somewhat larger, functional, 'female' spore (more appropriately called a megaspore) that germinates to produce internally (within its wall) a 'female' gametophyte (more appropriately called a megagametophyte). The single spore-like container of a 'female' gametophyte is enclosed in capsule-like tissue of the sporophyte referred to as the nucellus ('female' sporangium, more appropriately called a megasporangium), which is sheathed by enclosing structures of the sporophyte called integument(s) that possess an opening (tiny hole) called the micropyle to recieve sperm generated by the 'male' gametophytes transported by the pollen. Therefore, in seed plants, the 'female' gametophytes no longer become completely separated from the sporophyte, because they are each enclosed and live within sporophytic tissue. In most seed plants, the microgametophyte emerges from transported pollen as a sperm containing, narrow, filamentous, tube-like structures called the pollen tube that passes through the micropyle, nucellus, and megaspore wall to fertilize the egg containing megagametophyte. Therefore, in seed plants, the megagametophyte [housed within the nucellus almost completely surrounded by the integument(s) and often more directly surrounded by a megaspore wall] has become dependent for protection and survival on the sporophyte. The entire sheathed (integumented) megasporangium (nucellus) is the structure of the sporophyte that generates by meiosis the megaspores (with usually only one functional megaspore out of four able to eventually germinate to produce by mitosis an internal megagametophyte). This entire structure also called the ovule is attached to the rest of the sporophyte and becomes the seed only after the egg cell of the internal megagametophyte is fertilized by the sperm of a microgametophyte and development of the fertilized egg (zygote) via mitosis into an embryo takes place. Therefore, with development of the 'female' gametophyte, pollination, fertilization, and development of an embryo, the ovule becomes the seed. In essentially all presently living seed plants, the dispersal of the megaspore usually occurs only after development of the seed. However, in very rare cases, dispersal of the megaspore as part of ovules can occur prior to seed development, because some ovules of the non-flowering seed plant species Ginkgo biloba can be pollinated prior to dispersal and fertilized after dispersal.
    The lineage of seed plants, including angiosperms (flowering plants), as well as other gymnosperms (non-flowering seed plants), can be distinguished from clubmosses, ferns, horsetails, and other more primitive plants by the presence of ovules, which upon fertilization of egg cells within them, become the seeds. The ovule of presently living seed plants can be defined as consisting of a spore producing structure called the nucellus (= seed plant megasporangium), which is enclosed by a sheath called the integument (forerunner of seed coat) with integumentary micropyle [a small canal in the integument through which the fertile contents (sperm) produced from the germinated pollen grain can penetrate and fuse with the egg]). [The so-called pre-ovules of early proto-seed plant fossils did not have an entire integument (= integument lobes completely fused together and almost completely enclosing the nucellus) and a true micropyle (= integumentary micropyle)]. There is megasporangium indehiscence (= female spore producing part (nucellus or megasporangium) of the ovule is indehiscent (does not break open to liberate spores). After meiosis within the ovule, there remains only a single function female spore (= megaspore), which germinates to form within the megaspore wall a minute egg producing plant called endosporic female gametophyte (= endosporic megagametophyte). As stated elsewhere, the land plants, except for the seed plants, all have a free-living independent gametophyte. Nevertheless, in all embryophytes (even in the highly derived flowering plants), the gametophyte can still be said to nurture (at least initially) the zygote, embryo, or young sporophyte. In seed plants, the dividing (germinating) megaspore within an integumented sporangium receives its nutrition from the parental sporophyte, whereas in other heterosporous land plants (with the possible exception of only some extinct arborescent lycopsids), the megaspore is free-living; and although a certain amount of megaspore development can take place within a non-integumented megasporangium with resulting endosporic gametophyte development (within the megasore wall), the megaspore is eventually shed prior to fertilization (as in spikemoss called Selaginella). Therefore, even in non-seed plant, heterosporous embryophytes, megaspore development and the resulting gametophyte can be said to be independent of the sporophyte. See free-sporing versus seed plant reproduction. The rare and somewhat questionable exceptions to this in only some extinct arborescent lycopsids can help to clarify the essential characters that in combination can be used to define other seed-like modes of reproduction. The seed plant pollen grains (as mature seed plant microspores) internally form minute sperm producing plants called the male gametophytes (= microgametophytes). Fertilization (fusion of the sperm with the egg) takes place within the ovule, either before or (in certain very rare cases) after it is shed. An ovule becomes a seed if it contains an egg that has been fertilized with at least some development into an embryo. Such a propagule (dispersal unit that spreads a species from one place to another) can often undergo a period of dormancy (a resting stage) before it germinates to produce a seedling (young seed plant sporophyte).

    Spermatophyta Britton & Brown (= spermatophytes s.s.)
    Spermatophyta is the lineage comprising the seed plants (spermatophytes) other than medullosans or the "hydraspermans" and earlier plants with dispersed (shed), fossil seed-like propagules that can be called 'pre-ovules' and sometimes 'pre-seeds' or 'proto-seeds'. Spermatophyta (as define here) strictly corresponds to the lineage that includes the fossil platysperms plus all presently living seed plants.
    The shared, derived characters (acording to Cantino, et al. 2007) of spermatophytes s.s. include:
    (1) endarch primary xylem in the stem.
    (2) both male meiospores and microgametophytes occurring together as individual units called pollen grains with distal aperture or sulcus.
    (3) a linear tetrad of megaspores (four megaspores arranged in a straight line).
    (4) platyspermic ovules with presence of some signs of bilateral or biradial symmetry (such as two vascular bundles); platyspermic ovules are sometimes but not necessarily flattened or disc-like in shape.
    Although the broader seed plant lineage called Apo-Spermatophyta P.D. Cantino & M.J. Donoghue (= Spermatophytata Kenrick & Crane) is often considered the same as Spermatophyta, the former lineage is defined solely by the presence of ovules (Cantino, et al. 2007), which can be defined as an integumented, indehiscent (unsplitting) megasporangia, each containing a single (functional) megaspore and its internal (endosporic) megagametophyte. There were early seed plants known only from fossils that possessed an ovule (as a shared, derived character) but that lacked the above four shared, derived characters of Spermatophyta. Apparently, Spermatophyta is a lineage nested within the broader lineage called Apo-Spermatophyta that also includes all known seed plant fossils that are not platyosperms.
    o The Late Carboniferous "seed fern" fossil Callistophyton and the living cycad genus Cycas can be said to have platyspermic ovules that are bilaterally symmetrical, while all other living cycads have secondarily spherical (radiospermic) ovules. The secondarily radiospermic seeds of cycads are apparently more closely related to their platyspermic progenitors than to the radiospermic (radially symmetrical) seeds of Paleozoic "seed ferns", including those of medullosans, of the broader seed plant lineage referred to as Apo-Spermatophyta. It can probably be said that all presently living seed plants had a platyspermic ancestry.
    o Usually, the threshold distinguishing the ovule from the seed is fertilization mostly associated with some development of an embryo prior to dispersal [in other words, the ovule (as an immature seed not yet fertilized) is often distinguished from the seed (as a mature ovule already pollinated and fertilized, containing a partially developed embryo, prior to dispersal)]. This can often be used to not only distinguish an ovule from a seed but also distinguish a seed from a megaspore of "free-sporing plants", provided that the ovule or its resulting seed possesses megasporangium indehiscence. However, although very rare in modern seed plants, the pre-fertilization dispersal of some of the ovules of Ginkgo makes this distinction between ovules and seeds appear somewhat fuzzy.
    o A linear tetrad of megaspores results from two divisions of meiosis accompanied by cell wall formation. This results in the development of a well-defined linear row or tetrad of four megaspores. Most commonly, the three upper megaspores nearest the micropyle degenerate and the lower surviving megaspore enlarges to become the only functional megaspore.
    o The dependent megaspore that receives nutrition from the parent sporophyte might be considered a shared, derived character of seed plants. This condition certainly differs from the independent megaspores of other heterosporous land plants.
    o In tracheary element evolution, the proportion of the decay-resistant part within secondary wall thickenings increases while that of the nonresistant part decreases and finally becomes more or less absent in living seed plants. This more consistently lignified and resistant secondary wall of tracheary elements of living seed plants certainly differs from the core of nonresistant material in the tracheary elements of monilophytes and lycophytes. This high proportion of decay-resistant material may be considered a potential shared, derived character of fossil and living seed plants.
    o Megasporangium indehiscence (a megasporangium that never opens or splits) is the most distinctive feature of the ovule. Even the earliest 'proto-seed' plants had this feature that is usually correlated with a single megaspore per megasporangium and the presence of an integument and often (but not always) a cupule surrounding the ovule. The integument of most of these so-called 'proto-seeds' was lobed and never almost completely enclosing (as in modern seeds) the nucellus. Therefore, in these early plants, there was no need for a integumentary micropyle, which was absent.
    o The earliest known 'proto-seed' plants of the Middle Devonian had integument lobes only partially enclosing the nucellus and a lobed cupule partially surrounding the ovules.
    o Seed-like reproduction and seed plant reproducion can apparently be distinguish on the basis of whether the megasporangia are indehiscent (non-splitting) or dehiscent (splitting). For an example, a quick overview of seed-like reproduction in unrelated lycophytes is included below. However, in "hydraspermans", the distinction between the ovule and seed becomes a fuzzy issue. Therefore, the dispersed (shed), fossil seed-like propagules (apparently with megasporangium indehiscence) of "hydraspermans", including some medullosans) of the Upper Devonian and Carboniferous or earlier plants of the Middle Devonian are often referred to as 'pre-ovules' and sometimes 'pre-seeds' or 'proto-seeds'.
    o Apparently, pollination of 'pre-ovules' occurred prior to dispersal. This is called in situ pollination. However, it is uncertain whether fertilization and some initial development of the embryo occurred before dispersal. Most modern seed plants have both in situ pollination and in situ fertilization (pollination and fertilization of ovules prior to dispersal). However, some of the ovules of the still living plant of the genus Ginkgo can undergo pre-fertilization dispersal.
    o Some of Upper Devonian and Carboniferous fossil "seed ferns" had a mesarch primary xylem, radiospermic ovules, and pollen with an absence of a sulcus. Several of these Paleozoic "seed ferns" with hydrasperman reproduction and the medullosans with or without this mode of reproduction apparently had the less derived radiospermic (radially symmetrical) ovules, while the remainder of the seed plants called platysperms can be characterized by the more derived platyspermic seeds that were bilaterally or biradially symmetrical.
    o Another possible lineage (usually not formally named) includes all the medullosans and platysperms. In some studies, the medullosans are weakly supported as monophyletic and could be sister to the platysperms.
    o In a lineage far removed from that of seed plants, the partial splitting (partial dehiscence) of megasporangium of arborescent lycopsids, like the fossil Lepidodendron seed-like structures called Lepidocarpon, left an opening that enabled sperm to swim and reach the archegonia of the endosporic megagametophyte. Such seed-like structures can be distinguished from true seeds by their partial dehiscence (splitting).

  • Cantino, Philip D., James A. Doyle, Sean W. Graham, Walter S. Judd, Richard G. Olmstead, Douglas E. Soltis, Pamela S. Soltis & Michael J. Donoghue (2007) Towards a phylogenetic nomenclature of Tracheophyta, Taxon 56 (3) E1-E44.

    Spores of embryophytes
    Embryophytes produce haploid spores (meiospores) by meiosis of diploid spore mother cells (sporocytes) located in multicellular sporangia of diploid sporophytes. In contrast, the fresh water "streptophyte algae" that are the closest living relatives to all embryophytes do not develop from the zygote an embryo and spore producting sporophyte, but instead they produce haploid meiospores directly from meiosis of the diploid zygote that functions like a single spore mother cell of the land plant sporophyte. Because spores of plants on land compared to those of submerged ('under water') algae become exposed to increased solar UV-B radiation levels, desiccation, and likelihood of mechanical damage, an essential prerequisite for plant colonization of dry land with exposure to subaeral ('under air') conditions is the ability to protect the ungerminated haploid gametophyte (spore). The key to understanding how this has been achieved centers on the ability of land plants and their freshwater ancestors to protect zygotes and spores by producing the mysterious molecule called sporopollenin. This substance appears to be a huge co-polymer composed of an aromatic and aliphatic component of many organic chemical units (monomers) linked by ether and ester chemical bonds in a complex, matrix-like fashion. Sporopollenin is referred to as a co-polymer, because it appears to be composed of two (aromatic and aliphatic) components. The organic compounds (monomers) of the aromatic component apparently comprise phenylpropanoids (e.g., p-coumaric acid, caffeic acid, ferrulic acid), while those of aliphatic component can include medium- to long-carbon chain fatty acid (e.g., lauric acid) derivatives with in-chain or end-of-chain hydroxyl (-OH) groups (Dobritsa et al., 2009; Li et al., 2010; Morant et al., 2007). The phenylpropanoid compounds are considered aromatic, because they each contain a phenyl group (benzene ring), while the fatty acids are considered aliphatic, because they each contain open carbon chains that do not form into rings. The two or more different types of fatty acids with different carbon chain lengths and hydroxyl groups in different positions on their carbon chains, each containing a carboxyl (-COOH) group, provide higher level fatty acid to fatty acid and fatty acid to phenylpropanoid cross-linking, while the two or more different types of phenylpropanoids with hydroxyl groups in different positions on their phenyl group (benzene ring), each containing a shorter open carbon chain also with a carboxyl (-COOH) group, provide higher level phenylpropanoid to phenylpropanoid and phenylpropanoid to fatty acid cross-linking. It is the handle-like -OH groups of any two monomers with the release of water (HOH) that make it possible for the cross-linking. However, the -OH of a -COOH group of one monomer can combine with a simple -OH group of another monomer with the release of HOH to form an ester linkage (-COO-), while the simple -OH groups of different monomers with the release of HOH can combine to form an ether linkage (-O-). The complexity of this cross-linking has apparently contributed to the remarkable strength and resistence of this copolymer. Among other possible functions, it is the phenylpropanoids that protect the ungerminated gametophyte (spore) from exposure to higher levels of UV-B radiation from sunlight. Although the complex chemical configuration of sporopollenin is still not totally understood, there is much evidence that it has been conserved from "streptophyte algae" and cryptospore microfossils (see below) to embryophytes (de Leeuw et al., 2006; Fraser et al., 2012). The protective sporopollenin rich covering of the haploid cells called meiospores resulting from meiosis of presently living plants on land should provide one of the most important clues to how plants were both first able to undergo terrestrialization [the colonization of land from an aquatic (water inhabiting) ancestry] and become sustained and dominant in the subaerial ('under air') realm. Although a sporopollenin-like covering in an inner layer of the zygote wall is typical of some "streptophyte algae" (Delwiche et al., 1989; Graham 1993; Graham & Gray, 2001), the spores produced by these algae via meiosis lack an outer sporopolleninan-impragnated wall that is found in the spores of embryophytes. However, the possible land colonizing progenitors of embryophytes known only from spore-like microfossils called cryptospores had a life cycle that involved sporopollenin-impregnated walls on the surface of spores possibly (?) from zygotes that directly underwent meiosis (Strother, 2010; Strother et al., 2015). It is thought that sporopollenin-impregnated walls of early cryptospores originated prior to the origin of sporophytes. On the basis of evidence from presently living bryophytes, this was probably due simply to a modification of the timing of meiosis and the deposition of sporopollenin (Brown and Lemmon, 2011). Such a change would significantly increase the viability of spores, protecting the ungerminated gametophytes (spores) from the increased solar UV-B radiation levels, extreme temperatures, and increased likelihood of desiccation or mechanical damage of the subaerial realm. The deposition of sporopollenin on meiospores rather than the zygote, resulting in a sporopollenin-rich outer layer (exine) of the spore wall (sporoderm), could possibly be considered a character unique to the spores of subaerial progenitors that was eventually (in the coarse evolution) transfered via an intervening sporophyte generation to the spores of the embryophyte descendants. Therefore, this sporopollenin covering of meiospores only found in the subaerial realm could be considered a shared, derived (synapomorphic) character unique and common to both embryophytes and their possibly more immediate progenitors that are only known from the microfossils called cryptospores. The newly produced meiospores of embryophytes first appear in groups of 4 (tetrads) that soon separate into individual spores called monads. The individual spores possess distinctive surface features (markings) providing evidence that they were initially associated with other members in tetrads before they became dispersed. Such characteristic markings [often referred to as haptotypic mark(s) or laesura(e)] can in fossils provide evidence for the occurrence of meiosis. However, the cryptospores, although containing a sporopollenin covering and presumably representing an intermediate stage between aquatic "strepotophyte algae" and embryophytes, are spore-like bodies that lack the haptotypic characters (e.g., trilete mark) typical of embryophyte spores.
    Embryophytes presently include either "free-sporing plants" [e.g., liverworts, mosses, hornworts, lycophytes or lycopsids (e.g., clubmosses, spikemosses, and quillworts), monilophytes (e.g., horsetails, whisk ferns, and other ferns)] or seed plants (gymnosperms and angiosperms). Land plant spores are produced in the sporangia of sporophytes via meiosis of specialized cells called spore mother cells. The liverworts, mosses, and hornworts produce only one type of spore in one type of sporangium and, therefore, are said to be homosporous. Among the lycophytes, the clubmosses are all homosporous, while the spikemosses and quillworts produce two types of spores that can be called male and female spores in two corresponding types of male and female sporangia and, therefore, are said to be heterosporous. The monilophytes are mostly homosporous and rarely heterosporous (e.g., the water ferns). The seed plants are all heterosporous. In contast to free-sporing plants (sometimes called spore plants), seed plants disperse their male spores as pollen and female spores as part of the seed. In the "free-sporing plants" (those that do not produce seeds), the homosporous condition is sometimes defined as the production of functional (viable) spores of only one type (usually similar in size) that germinate to produce gametophytes each with both male and female parts [e.g., a male sex organ called the antheridium (composed of a jacket of multiple cells surrounding the sperm producing tissue) and a female sex organ called the archegonium (flask shaped structure composed of multiple cells, with often a slender neck-like projection and a widened, often rounded, egg containing base)], while heterosporous condition is sometimes defined as the production of two types of distinctively male and female spores, with each viable (usually smaller) male spore germinating to produce a male gametophyte that bears only male sex organs and each viable (usually larger) female spore germinating to produce a female gametophyte that bears only female sex organs. However, the distinction between the homosporous and heterosporous condition in "free-sporing plants" on this basis alone can sometimes become fussy and even inadequite, because some "free-sporing plants" that are considered homosporous can produce functional (viable) isospores [all alike (indistinguishable) in form, structure, size, and development] that upon germination yield distinct male and female gametophytes. Therefore, in an attempt to distinguish these two conditions, it is best to consider the homosporous condition as transitional with different degrees of the heterosporous condition. In homosporous land plants with all spores identical (isospores), both male and female sex organs are often located on the same (bisexual or monoecious) gametophyte, but sometimes, depending on the species, the male and female organs occur on separate (unisexual or dioecious) gametophytes. Although bisexual (monoecious) gametophytes (bearing both male and female sex organs) produced by isospores of homosporous sporophytes may ultimately turn out to be the primitive type, all of the fossil gametophytes thus far described for pre-vascular and early vascular 'polysporangiophytes' from the Rhynie chert (Early Devonian chert deposits of Aberdeenshire, Scotland) appear to be unisexual (producing two sexually different gametophytes). [In some of the living homosporous freshwater relatives of the land plant ancestor, the spores of the "algae" in the order Coleochaetales produce bisexual (monoecious) gametophytes, while the spores of those in the order Charales (depending on the species) produce either bisexual (monoecious) or unisexual (dioecious) gametophytes.] The homosporous land plants called mosses also have some species with spores that develop into monoecious gametophytes and other species with spores that develop into dioecious gametophytes. In the homosporous horsetail genus Equisetum subgenus Hippochaete smaller gametophytes bear only antheridia, while larger gametophytes initially bear archegonia but later undergo a transition to bear antheridia. A similar situation occurs in the fresh water inhabiting, homosporous fern species Ceratopteris thalictroides, although there is a slight difference in spore size with smaller spores producing gametophytes with only antheridia and the larger ones producing the transitional archegonia-antheridia types. Therefore, to be sure that a certain plant is heterosporous, the production of distinct male and female (dioecious) gametophytes is necessary but not sufficient. However, if dioecious gametophytes can be coupled with one or more the different degrees of the heterosporous condition, such as dimorphic spores (e.g., smaller male and larger female spores) associated with distinct male and female sporangia, this becomes a more useful way of distinguishing heterosporous plants from those that are homosporous. The different degrees of the heterosporous condition (according to Bateman and DiMichele, 1994) are covered in a little more detail in the discussion on reduction of the gametophyte in embryophytes.
    When meiosis takes place in each spore generating cell (sporocyte) of the sporangia, there is often formed a group of four (frequently genetically dissimilar) spores, each attached centrally in often a tetrahedral (or linear) arrangement called a tetrad. Spores free from each other rather than attached in groups are called monads. The proximal face of a monads is considered the original contact face of the spore while still in the tetrad stage. The distal face is on the side of the spore furthest away from the original contact face. When the four spores become detached from each other, a triradiate scar or trilete mark can remain on the proximal face of each monad that marks the place of original attachment in the tetrad. Detached trilete spores (those with well defined triradiate scars or trilete marks on their original contact faces) are characteristic spore types in polysporangiophytes (including early vascular plants or their stem groups). The relative abundance of trilete spores in the fossil record has been generally considered an indication of the appoximate date or time period of the emergence of these lineages. Germination of spores in ferns is proximal or at the site of the trilete mark. The deep cleavage (trilete aperture) at the site of the trilete mark often can be observed in fossils, indicating proximal germination.
    Pollen is seed plant microspores shed only after they become mature with internal microgametophytes. If pollen has a distinctive trilete mark that indicates proximal germination, as was the case in the earlier (fossil) seed plants, this is what is referred to as pre-pollen. Such pre-pollen can release internal microgametophyte produced motile sperm from a proximal trilete aperture. In contrast to pre-pollen, the germination of pollen in later conifers and flowering plants is at the distal face of the microspore. In most seed plants, distal germination of the pollen grain is indicated by the development of a distal germiation groove or furrow (= distal aperature) also called the sulcus to facilitate the emergence from the upper end of the microspore wall of a pollen tube, which can be defined as a tubular extension of the microgametophyte that eventially grows into (or penetrates) the megasporangium. If a sulcus is present in fossil pollen, it is assumed that a pollen tube was also produced. This is a valuable association, because pollen tubes are rarely preserved in fossils. Furthermore, when the germination groove called the sulcus develops, there is a tendancy to loose the trilete mark. There are fossils with both a trilete mark (tetrad scar) and a sulcus. However, this mixed type is relatively rare, for most fossils referred to as the Paleozoic cordaites and conifers had only a sulcus, suggesting a strong tendency for loss of the trilete mark once a sulcus is present. Therefore, the fossil trilete microspores without a sulcus indicate (as a maker) the occurrence of proximal germination typical of what is referred to as pre-pollen of the earlier seed plants. Actually, during the late Paleozoic period, fossil pre-pollen appears to have been the rule rather than the exception in most seed plant lineages.

  • Bateman, R.M. and W.A. DiMichele (1994) Heterospory: the most iterative key innovation in the evolutionary history of the plant kingdom, Biological Reviews 69: 345-417.
  • Brown, RC. and BE. Lemmon (2011) Spores before sporophytes: Hypothesizing the origin of sporogenesis at the algal-plant transition. New Phytol. 190: 875-881.
  • de Leeuw, J. W., Versteegh, G. J., & van Bergen, P. F. (2005) Biomacromolecules of algae and plants and their fossil analogues. In Plants and Climate Change (pp. 209-233). Springer, Dordrecht.
  • Delwiche CF, Graham LE, Thomson N. (1989) Lignin-like compounds and sporopollenin in Coleochaete, an algal model for land plant ancestry. Science 245: 399-401.
  • Dobritsa, A.A., Shrestha, J., Morant, M., Pinot, F., Matsuno, M., Swanson, R., Moller, B.L. and Preuss, D. (2009) CYP704B1 is a long-chain fatty acid omega-hydroxylase essential for sporopollenin synthesis in pollen of Arabidopsis. Plant Physiol. 151, 574-589
  • Graham, LE. (1993) Origin of land plants. New York: Wiley.
  • Graham, LE., & J. Gray (2001) The origin, morphology, and ecophysiology of early embryophytes: neontological and paleontological perspectives. Plants invade the land: evolutionary and environmental perspectives, 140-158.
  • Fraser, W. T., Scott, A. C., Forbes, A. E. S., Glasspool, I. J., Plotnick, R. E., Kenig, F., & Lomax, B. H. (2012) Evolutionary stasis of sporopollenin biochemistry revealed by unaltered Pennsylvanian spores. New Phytologist, 196(2), 397-401.
  • Li, H., Pinot, F., Sauveplane, V. et al. (2010) Cytochrome P450 family member CYP704B2 catalyzes the {omega}-hydroxylation of fatty acids and is required for anther cutin biosynthesis and pollen exine formation in rice. Plant Cell, 22, 173-190.
  • Morant, M., K. Jorgensen, H. Schaller, F. Pinot, BL. Moller, D. Werck-Reichhart, and S. Bak (2007) CYP703 Is an Ancient Cytochrome P450 in Land Plants Catalyzing in-Chain Hydroxylation of Lauric Acid to Provide Building Blocks for Sporopollenin Synthesis in Pollen, The Plant Cell, Vol. 19: 1473-1487.
  • Strother, PK. (2010) Thalloid carbonaceous incrustations and the asynchronous evolution of embryophyte characters during the Early Paleozoic. Int. J. Coal Geol. 83, 154-161.
  • Strother, PK., A. Traverse, M. Vecoli (2015) Cryptospores from the Hanadir Shale Member of the Qasim Formation, Ordovician (Darriwilian) of Saudi Arabia: taxonomy and systematics, Rev. Palaeobot. Palynol. 212, 97-110.

    Sporophyte and gametophyte of embryophytes
    It may surprise the reader that in sexual reproduction of land plants called embryophytes, individuals of any given species include a multicellular organism called the sporophyte or plant that first develops as the embryo from the fertilized egg cell also called the zygote, at maturity produces the spores, and from a germinated spore, another multicellular organism develops called the gametophyte or plant that at maturity produces the gametes or sex cells, the sperm and egg. This relationship of two organisms, one producing the spores and the other producing the gametes, is found in all embryophytes from the earliest, least derived forms to the most derived vascular plants, including the presently living lycophytes and euphyllophytes. As in all eukaryotes with sexual reproduction, two types of cell division are involved in the sexual life cycle of land plants. The type of cell division called mitosis usually involves exact cell duplication, maintaining the same number of chromosomes in the nucleus as the precursor cell. The type of cell division called meiosis reduces the number of chromosomes to half that of the precursor cell called (in land plants) the spore mother cell or sporocyte that divides twice to produce four spores. The sporophyte develops from the type of cell division called mitosis of the fertilized egg cell (zygote) to become the multicellular embryo and the embryo develops into a multicellular organism that at maturity produces by further mitosis the multicellular housing (sporangium) for the generation of spores, while the gametophyte develops from mitosis of the germinating spore to become a multicellular organism that at maturity produces by further mitosis the multicellular housing (sex organs) for the gametes. Within the sex organs, the gametes themselves are also produced by mitosis. The spores themselves are produced by the other type of cell division called meiosis that reduces the number of chromosomes in the nucleus of each cell from 2n (the diploid number) to n (the haploid number), where 2n represents twice the number as n. During fertilization (union the egg cell and the sperm cell) to produce the fertilized egg cell (zygote), the the chromosome number is restored to 2n. Therefore, the life cycle of land plants can be characterized as the alternation of generations of the sporophyte with the gametophyte, which has become part of the alternation of meiosis with fertilization found in all eukaryotes with sexual reproduction. In the land plants called liverworts, mosses, and hornworts, the often smaller sporophyte (with the diploid chromosome number 2n) remains attached to and dependent on the often larger (dominant) gametophyte (with the haploid chromosome number n), while in the land plants called vascular plants, the often (eventually) larger (dominant) sporophyte (with 2n chromosomes) soon becomes separate (independent) from the often smaller gametophyte (with n chromosomes). In liverworts, mosses, and hornworts (collectively called bryophytes), the single axis of the sporophyte plant, perminately attached to the larger mother gametophyte, remains unbranched bearing a single sporangium, while in vascular plants, the main axis of the sporophyte plant, only temporarily attached to the gametophyte and becoming independent in early development, grows into a usually larger, often highly branched organism bearing multiple sporangia.

    Thelesperma - Less. - Genus Name
    Family Asteraceae
    From analysis of DNA of the tribe Coreopsideae Turner & Powell [Crawford, et al. (2009)], the distinctive genus Thelesperma Less. is a well defined lineage (monophyletic group) that appears to be derived from within a broader lineage comprising mostly eastern North American (mostly US) species of the polyphyletic genus "Coreopsis" Linnaeus and certain north temperate species of the polyphyletic genus "Bidens" Linnaeus.
    The broader, highly derived, currently unnamed, but apparently well-supported and probably terminal lineage within which the genus Thelesperma is nested within the tribe Coreopsideae includes:
    The sublineage informally called Coreopsis-6 +
    [the sublineage informally called Bidens-3 +
    [the species Bidens tripartita Linnaeus [= Bidens comosa (A. Gray) Wiegand] +
    [the sublineage informally called Coreopsis-7 + the genus Thelesperma]]].
    The following includes species of the lineages (or sublineages) of the polyphyletic genera "Coreopsis" and "Bidens" that appear most closely related to Thelesperma on the basis of the DNA analysis of Crawford, et al. (2009):
    o Coreopsis-6 includes eastern North American species of Coreopsis section Gyrophyllum Nuttall (e.g., C. delphiniifolia Lamarck, C. major Walter, C. palmata Nuttall, C. pulchra F. E. Boynton, C. tripteris Linnaeus, and C. verticillata Linnaeus) and section Silphidium (Torrey & A. Gray) A. Gray (e.g., C. latifolia Michaux).
    o Bidens-3 includes north temperate species of Bidens section Bidens (e.g., B. cernua L. and B. frondosa L.) and section Hydrocarpaea A. Gray (e.g., B. beckii Torr.).
    o Bidens tripartita Linnaeus [= Bidens comosa (A. Gray) Wiegand] is a north temperate species native to North America (also New Mexico), Europe, Asia, and north Africa [introduced elsewhere (e.g., Pacific Islands and Australia)].
    o Coreopsis-7 includes mostly eastern North American species of Coreopsis section Calliopsis (e.g., Coreopsis leavenworthii Torrey & A. Gray and Coreopsis tinctoria Nuttall), section Coreopsis (e.g., Coreopsis auriculata Linnaeus, Coreopsis basalis (A. Dietrich) S.F. Blake, Coreopsis grandiflora Hogg ex Sweet, Coreopsis intermedia Sherff, Coreopsis lanceolata Linnaeus, Coreopsis nuecensis A. Heller, and Coreopsis pubescens Elliott), and section Eublepharis Nuttall (e.g., Coreopsis gladiata Walter, Coreopsis integrifolia Poiret, Coreopsis nudata Nuttall, and Coreopsis rosea Nuttall).
    Although relationships of plants within the above broader lineage are not yet that well supported and more DNA work is required, it appears that the species mentioned here (especially those in the sections Calliopsis, Coreopsis, and Eublepharis of the genus Coreopsis) are the closest relatives of the species of Thelesperma.
    Therefore, the genus Thelesperma appears from analysis of DNA to be nested within a lineage including mostly eastern North American species of "Coreopsis" and some north temperate species of "Bidens."
    Although no species of Coreopsis section Eublepharis is found wild as far west as New Mexico, Coreopsis tinctoria of section Calliopsis, as well as Coreopsis grandiflora and Coreopsis lanceolata [possibly (?) also Coreopsis basalis (A. Dietrich) S.F. Blake var. wrightii (Gray) S.F. Blake] of section Coreopsis, can be found wild in some parts of the state.
    Species of Thelesperma are used widely in southwestern United States as herbal beverages or for substitutes of tea or coffee, as well as for various medicinal purposes. Since Coreopsis tinctoria, as well as Coreopsis grandiflora and Coreopsis lanceolata, are members of two of the mostly eastern North American sections of "Coreopsis" apparently very closely related to Thelesperma according to DNA analysis, it is expected that in New Mexico these species of "Coreopsis" and the species of Thelesperma should possibly have similar chemical and medicinal properties with similar potential uses as beverages or medicinal agents. Supportive of this is documented evidence from New Mexico that at least the species Coreopsis tinctoria of section Calliopsis has been prepared and used as a beverage and medicinal agent in many of the same ways as species of Thelesperma. For example, New Mexican Native Americans have been reported to have made a beverage (as a substitute for coffee or tea) from Coreopsis tinctoria by folding the dried plant stems (including attached leaves and flowering tops) into bundels and steeping the bundels in hot, boiled water. Like Thelesperma, Coreopsis tinctoria has also been reported to have been used as a medicinal agent to treat such health conditions as diarrhea and type 2 diabetis. See further discussions by following the links in the dictionary under the polyphyletic genus "Coreopsis".
    Although Coreopsis tinctoria is considered an annual, sometimes (if the weather remains warm or mildly cool) it will bloom for two or three years. This species has distinctive (bicolored) radiate flower heads with red-brown to purplish centers extending various distances toward the brilliant yellow margins of the ray flowers. Some of these plants have ray and disk flowers that are red-brown to purplish throughout. Some other annual species only found in Texas (endemics), including Thelesperma burridgeanum (Regel) S. F. Blake with ray flower laminae red-brown to purplish throughout or ray flower laminae mostly red-brown with more peripheral margins yellow to orange and disk flowers red-brown at least toward their tips and Thelesperma nuecense B. L. Turner with ray flower laminae yellow to golden but sometimes suffused with red-brown or with a basal red-brown spot or band and disk flowers red-brown at least apically, have radiate flower heads that can appear very similar to those of Coreopsis tinctoria. Like the annual species Thelesperma flavodiscum (Shinners) B. L. Turner (also only found in Texas) with ray flower laminae all yellow and disk flowers yellow, Thelesperma burridgeanum and Thelesperma nuecense are closely related to and possibly indistinguishable from the more widespread annual species Thelesperma filifolium with ray flower laminae yellow to golden yellow but sometimes basally red-brown and disk flowers red-brown or yellow with red-brown nerves (viens). Although no reports on use could be found for Thelesperma burridgeanum, Thelesperma nuecense, and Thelesperma flavodiscum, these Texas endemic plants (possibly only varients of Thelesperma filifolium) are likely (almost certainly) to have chemical and medicinal properties similar to those of Coreopsis tinctoria, Thelesperma filifolium, and other species of the genus Thelesperma. The more widespread annual Thelesperma filifolium can be found as far southeast as Mississippi and Louisiana, extending into Texas and Nuevo León in Mexico, as well as into New Mexico, Colorado, Wyoming, Montana, and other parts of the Great Plains. The annual Coreopsis tinctoria has a similar but much more widespread distribution as far north as almost all of southern Canada, including northeastern United States (e.g., as far north as Maine), southeastern United States (e.g., as far south as Mississippi and Louisiana) where it is commonly called Calliopsis, western United States (e.g., as far west as California, Oregon, and Washington), the Great Plains (especially), Arizona, New Mexico, Texas, and northeastern Mexico (Coahuila, Nuevo León, Tamaulipas). This species has also been widely introduced to other parts of the world (e.g., traditionally used for type 2 diabetes in Portugal and as a daily tea beverage and medicinal agent for diabetes and diarrhea in China).
    The leaves of species of Thelesperma are mostly opposite (pairs of leaves even with each other at each leaf bearing node on opposite sides of stem) or upper (distal) leaves rarely alternate (only one leaf at each node); blades of leaves (or their lobes) usually not that hairy [often glabrous (hairless)] and simple (never compound with separate leaflets) but pinnately lobed (becoming lobed from different points along an axis like sections of a feather). The ultimate lobes of the leaves are often much longer than wide to very thin (filiform). If most of the leaves are not opposite, the plant is almost certainly not a member of the genus Thelesperma. However, in the aster family, the feature mostly opposite leaves (like in Thelesperma) is also characteristic of most of the complex of tribes (the higher level lineage called the Heliantheae Alliance) from which the sunflowers (genus Helianthus) are derived, so more distinctive features are needed to identify the genus Thelesperma.
    The bracts geneally appear as small, sometimes modified, often reduced, leaf-like structures associated with individual flowers or an entire cluster of flowers called an inflorescence. For example, in Thelesperma and relatives, some bracts can be thin, dry, scale-like structures called paleae (or chaff) associated the individual flowers that are borne on a receptacle or they can be modified leaf-like structures associated with the entire flower cluster (inflorescence) as the one or two or more series (whorls) of bracts called phyllaries that collectively make up what is called the involucre, a structure which directly subtends (extends immediately below) the condensed cluster (head) of small flowers (florets), entirely encloses them in initial development, and occurs in nearly all members of the family Asteraceae. Go forward to discussion on radiate or discoid heads of family Asteraceae or see description of the of the involucre and calyculus of most members of the tribe Coreopsideae.
    The involucres in Thelesperma are usually campanulate (bell-shaped) to hemispheric (half spherical). See Thelesperma ID.
    The seed-like fruit (ripened inferior ovary called the achene or cypsela) of Thelesperma is compressed from back to front with margins (sides) sometimes slightly winged and with a pappus (modified calyx) absent or persistent at the top of the fruit as 2 unbarbed (ciliate) scales or short awns (small, sharp pointed, apical appendages).
    The genus Thelesperma can be distinguished in Coreopsideae from related genera by bracts (phyllaries) of involucre [those immediately surrounding the base of the compact flower cluster (head)] strongly united (inner phyllaries fused 1/5 to 7/8 their lengths). In most members of Coreopsideae, the outer, green, sometimes reflexed (bent backwards), and more leaf-like bracts are immediately below but appear distinct from the inner, erect, often membranaceous phyllaries of the involucre. (It is the inner phyllaries that completely surround the compact flower cluster in early development.) These inner involucral bracts have scarious (paper-thin) margins and are often striated [bearing thin resin ducts that appear as few to many, brownish-orange (or sometimes grey) parallel lines (vertical striations or striae)], with also a few brownish-orange striae found on the paleae [chaffy (dry) bracts (often collectively called chaff) more closely associated with the individual small flowers (florets)] on the flower bearing axis called the receptacle. If the inner phyllaries are partially but strongely united (as described above) and if the outer bracts (often collectively called the calyculus) are equal to at least 1/2 (sometimes surpassing) the lengths of the inner phyllaries, this narrows it down to two species both with ray flowers, including the perennials or subshrubs T. ambiguum [that live for three or more years with a well developed horizonal underground stem or rootstock (rhizome)] and the annuals T. filifolium [that germinate from seed, produce flowers and new seeds, and (without a perennial rootstock) die in the same year]. If outer bracts are less than 1/2 the lengths of the inner phyllaries, this applies to four species, including the rest of the species of the genus in New Mexico, two species with ray flowers absent (T. megapotamicum and T. longipes) and two with ray flowers usually present (T. simplicifolium and T. subnudum).
    Among the last mentioned four species, the lobes of the corollas of the disk flowers of T. megapotamicum are longer than the throats, a pappus of two smooth awns distinctly crowns each seed-like fruit, and ray flowers are absent. In the remaining 3 species in New Mexico, the lobes of the corollas of the disk flowers are shorter than the throats and the pappus is mostly absent or rarely present and less than 1/2 mm in length. The stem leaves are scattered along the stem (leaves scattered over the proximal 3/4 or more of the plant's height) in T. simplicifolium with usually 8 ray florets (rarely absent) with yellow laminae and yellow disk florets with red-brown nerves. In the remaining two species including T. longipes and T. subnudum, the leaves are crowded toward the base for 1/2 or less the length of the stems (with long nearly or completely leafless stalks that bear the flowers). The leaf lobes of T. longipes are thinner (1 mm or less), ray flowers are absent, and disk florets yellow. In contrast, the leaf lobes of T. subnudum can sometimes widen toward the tip and can range between 2 and 5 mm in width, with usually 8 ray florets bearing yellow laminae and yellow disk florets sometimes with red-brown nerves.
    o Daniel J. Crawford, Mesfin Tadesse, Mark E. Mort, Rebecca T. Kimball and Christopher P. Randle (2009) Coreopsideae. In V. A. Funk, A. Susanna, T. F. Stuessy, R. J. Bayer (eds.), Systematics, Evolution, and Biogeography of Compositae. Pp. 713-730. International Association for Plant Taxonomy, Vienna.

    Thelesperma ambiguum - Gray - Species Name
    Slender perennials or bushy subshrubs from rhizomes bearing masses of showy yellow flower heads in March-June that can be used together with young leaves to make a herbal tea.
    See Thelesperma ID.
    There is some interest in researching the medicinal properties and chemistry of the T. megapotamicum found wild in Argentina, South America (Barboza, et al., 2009). In this country, T. megapotamicum is commonly referred to as Té pampa (Pampa tea) or Té indio (Indian tea). The leaf is said to be digestive, antispasmodic, and employed for kidney diseases. The aerial parts are considered antibacterial and antioxidant. The entire plant contains the flavone chemicals called cynaroside and luteoline; the dried aerial parts contain the monoterpene chemical called thymohydroquinone dimethyl ether; and the dried root contains the phenylpropanoid chemical called 4-0-iso-butyryl-3-0-methyl-7-8-epoxy-coniferyl acetate. According to Ateya et al. (1982) and Dr. Duke's Phytochemical and Ethnobotanical Database, the flavone luteolin has been used as an inhibitor of the enzyme aldose reductase and has been shown to possess (among other properties) antiHIV, anticataract, anticomplementary, antidermatic, antifeedant, antiherpetic, antihistaminic, and antiinflammatory activity. Certain fractions of this species have been found to exert growth inhibition in cultured (in vitro) human breast cancer MCF-7 cells more effectively than the commonly used chemotherapeutic agent, paclitaxel (Figueroa, et al., 2012). Acetylenes from T. megapotamicum and T. filifolium have shown antibiotic action against the yeast Saccharomyces cerevisiae (Bohlmann, et al., 1973).
    o Figueroa, A. C., Soria, E. A., Cantero, J. J., Sanchez, M. S., & Goleniowski, M. E. (2012) Cytotoxic activity of Thelesperma megapotamicum organic fractions against MCF-7 human breast cancer cell line. Journal of Cancer Therapy, 3(01), 103.
    o Crawford, D. J., & Stuessy, T. F. (1981) The taxonomic significance of anthochlors in the subtribe Coreopsidinae (Compositae, Heliantheae). American Journal of Botany, 68(1), 107-117.
    o Pathak, V., F. Bohlmann, R. King and H. Robinson (1987) Chemotaxonomy of the Genus Thelesperma, Revista Latinoamericana de Quimica, Vol. 18, pp. 28-29.

    Tracheophyta Sinnott (= eutracheophytes)
    The vascular land plants are distinguished from other land plants (e.g., bryophytes) by characteristic thickenings that are deposited in a wide variety of patterns (such as rings, spirals, reticulate networks, or pitted regions) in the secondary walls surrounding water conducting cells. Although some bryophytes can have water conducting tissues, the usually thin cell walls of these tissues lack the secondary wall thickenings characteristic of vascular plants. Because the cell walls of most bryophytes are generally thin, primary or secondary walls can not be clearly distinguished. Bryophytes with such thin cell walls can easily absorb and transport water via pressure changes and simple diffusion. Therefore, they do not require a well developed water conducting system. Vascular plants can be distinguished as land plants that uniquely possess tracheids, defined as water conducting cells with secondary wall thickening; and a water conducting system with tracheids (and/or their derivatives) is referred to as the xylem. Both the water conducting cells of some bryophytes (certain mosses) and the tracheids of vascular plants are elongate cells that loose their living material at maturity, becoming empty cells consisting only of walls. However, unlike tracheids, the water conducting cells of mosses do not have an inner decay-resistent layer in their cell walls. It is the inner decay-resistent layer in the secondary wall and the unique secondary wall thickenings of the water conducting cells that characterize vascular plants and distinguish them from all others. Tracheids can also be elaborated as derivatives called vessel members that become attached at their ends to form vessels. The two possible types of water conducting cells (tracheids and vessel members) unique to vascular plants are both called tracheary elements. Although the xylem is defined as water conducting tissue that contains tracheids and/or their derivative vessel members as its most diagnotic feature, distinguishing it from the water conducting systems of non-vascular plants, the xylem also contains other types of cells that associate with the tracheids or vessel members. Furthermore, the xylem, as a rule, is always formed in association with the food conducting tissue called the phloem, composed of sieve elements or their derivatives (united into sieve tubes) as well as other associated cells. Vascular land plants are those with dominant sporophytes possessing water and food conducting tissue called xylem and phloem, respectively. However, even though the vascular system (unique to vascular plants) is defined as composed of both xylem and phloem, the enduring rigid cell walls with characteristic thickenings make the xylem much more conspicuous than the phloem. Therefore, it is the xylem, rather than the phloem, that mostly serves for the identification of vascular plants from the fossil record. The key substance responsible for the rigidity and endurance of the xylem is the complex molecule called lignin that is also exclusively found in vascular plants. It is lignin that is responsible for the rigidity and endurance of the decay-resistent layer in the secondary wall of tracheary elements of the xylem. In the development of all presently living vascular plants, there is a differenation of the first formed xylem into two parts called the protoxylem followed by the metaxylem. All fossil or living vascular plants with this shared, derived character are called eutracheophytes, which can be distinguished from other stem (fossil) group vascular plants by this differention in the first formed xylem called the primary xylem. A secondary xylem and phloem (or a secondary vascular system) is commonly found together with a primary xylem and phloem (or a primary vascular system) in certain (more derived) members within this lineage, but the secondary vascular system is not an ancestral condition common to all members. Therefore, the two shared, derived characters that are ancestral and define this lineage of vascular plants are (1) differentiation of protoxylem and metaxylem of the primary vascular system and (2) water-conducting cells called tracheids that possess thick, lignified, differentially thickened secondary walls with an inner decay-resistant layer. In order to gain a more thorough understanding of these lineage defining characters, check out the links on cells with walls, lignin, primary xylem, meristems, protomeristems, procambium, and protoxylem and metaxylem.
    It should be noted that three parts are commonly recognized in vascular plant cell walls: the glue-like substance between the cells or middle lamella (made up of sugar units linked together to form an adhesive gel producing pectin compound that can combine with calcium or can be made more rigid in woody plants by the incorporation of a substance called lignin), the primary wall (made up of cellulose, related substances, and some pectin, a combination that is sometimes made more rigid by a incorporation of some lignin), and the secondary wall (made up mainly of cellulose or varying mixtures of cellulose and related substances, a high cellulose combination that also can be modified through the deposition of lignin and other substances). The middle lamella can act like a 'cement' between the primary walls of two adjacent cells, and the secondary wall is laid over the primary, that is, next to the central cavity of the cell. The primary wall is the first wall proper formed in a developing cell. Of course, the secondary wall follows the primary in the order of appearance. The secondary wall is often considered a supplementary wall whose principle function is mechanical (primarily for structural support). It is most characteristic of cells that are highly specialized (differentiated) and undergo irreversible changes in their development. Some plant cells can become so specialized that they no longer need living material, becoming reduced to highly functional empty walls. This is especially true of the unliving water conducting cells called tracheids and a few other cell types of vascular plants, which (even though functional and important) can be thought of as mere structural components like empty shells that once contained living material. These thick walled, empty cells are the result of a process called programmed cell death that is almost exclusively found in vascular plants. Also unique and common to all vascular plants are the differentially thickened secondary walls with an inner decay-resistant layer.
    As land plant eating predators gradually became more common, plants gradually became tougher (more difficult to chew) and woodier due to the eventual origin of lignin, one of the most important plant cell wall substances. Lignin has been studied over one hundred years, but its complex chemistry is still not fully understood. The incorporation of lignin into the cellulose fibers that make up the cell wall is referred to as lignification, i.e., the cell wall can become lignified. Lignin apparently originated with the first emergence on land of the vascular plants. Although lignin functions as an important structural material likely always associated with the decay-resistent part in the secondary wall of water conducting cells of all fossil and living vascular plants, this complex chemical structure has not been generally associated directly with any medicinal properties other than the health promoting laxative effects of lignified cellulose or indigestable plant material (e.g., dietary fiber). Insoluble fiber consists of mostly cellulose and possibly a small amount of lignin, which comes from the indigestible cell walls of fruits and vegetables, while soluable fiber (without any lignin or cellulose) comes from water soluable pectins and gums, which can absorb water and add bulk to the large intestine. Both insoluble and soluable fiber are beneficial in maintaining a healthy colon, particularly in older adults with diverticulosis. The simple chemical building blocks of lignin are the same as those resulting in less complex chemicals called lignans, some of which can have many interesting and beneficial medicinal properties. However, unless one enjoys eating wood pulp and rope, insoluble fiber with a lot of lignin is not recommended for medicinal use. Although lignin can be important to plants in providing mechanical support, its indigestability can also protect cells in seeds and other plant parts from animal consumers.
    Since the water conducting xylem is the most obvious distinguishing feature of vascular plants, its initial development is described here in more detail. However, before describing this initial development of the xylem, an introduction to the initial development of internal anatomy of the main plant axis (including the shoot and root) plus the lateral shoots and roots is briefly outlined (see meristems, protomeristems, and procambium). During this development, the cells involved do not only increase in number by mitosis but also undergo elongation followed by differentiation, a process by which cells become specialized in their structure or take on particular functions.
    As the embryo sporophyte of a vascular plant enlarges and develops into an independent and dominant plant, the addition of new cells by mitosis is gradually restricted to certain perpetually young (embryonic) portions called meristems (regions on a plant of actively dividing meristematic cells that give rise to other so-called derivative cells), while other portions become specialized (through cell differentiation) for more mature functions other than the juvenile process of growth. Growth and development includes not only addition of new cells by mitosis but it can also involve cell growth (e.g., elongation) and differentiation. With much more specialization of cells than in more non-vascular plants (like bryophytes with cells essentially all alike), the function of cell division in vascular plants becomes more and more confined to meristems and their immediate derivatives (those cells immediately derived from previous cells of the meristems through division and sometimes only a slight amount of differentiation). Although cells other than those of the meristems can produce new cells, only the undifferentiated cells of the meristems remain active throughout the life of the plant, for they perpetuate themselves, retaining some products of division that do not differentiate into adult cells but continue to carry on the activity of division. The first meristems to arise in the embryo are called primary meristems that result in primary growth, most commonly involving elongation or increase in length. The primary meristems are called apical meristems, because they are at the apices or tips of main and lateral shoots plus main and lateral roots. In some vascular plants (and hornworts), there can also exist primary meristems called intercalary meristems, because they are inserted between two more adult regions of cells. However, it is the potentially immortal apical meristems at opposite ends of an axis that results in the basic (primary) structure of most presently living vascular plants. This results in what is referred to as bipolar primary growth, with both shoot and primary root present in embryo and causing the shoot and root to grow in opposite directions.
    [Bipolar primary growth contrasts with the unipolar primary growth of bryophytes and the earliest (known) 'polysporangiophytes' (including stem groups with or without well developed vascular systems), where the the apical meristem called the primary shoot meristem is present but the apical meristem called the primary root meristem is absent in the embryo. Unipolar growth is only briefly mentioned here, because it is mainly known only from bryophytes and some fossils of possible stem groups of the lineage of vascular plants. In more derived vascular plants, there are also meristems called lateral meristems, because they result in lateral increase in width or thickness of an plant axis. These meristems are only briefly mentioned here, because they were not present in the earliest (ancestral) eutracheophytes (much less the earlier, stem group vascular plants) that only had primary growth. Consequently, lateral meristems cannot be considered an ancestral condition that is common to all vascular plants. In certain (more woody) vascular plants, the lateral meristems become active usually only after some completion of primary growth. They are located within shoots or roots as often cylindical shaped regions (called the vascular cambium and cork cambium) and result in what is called secondary growth. Since primary growth is ancestral to all vascular plants, it is outlined here in more detail. This is done so that the reader can understand better what is meant by the differentiation of the primary xylem that is uniquely found as a shared, derived character in all presently living vascular plants and their acestors with a well developed vascular system. Secondary growth will be covered in a little more detail elsewhere.]
    It is the apical meristems involved in primary growth of the shoot and root that give rise to slightly more mature (slightly more differentiated) but still actively dividing regions called protomeristems. These protomeristems include the protoderm (yielding the outer surface dermal system), procambium (yielding the primary vascular system), and ground meristem (yielding the fundamental or ground tissue system). The procambium gives rise to the primary vascular system. The ground meristem gives rise to the fundamental or ground tissue system, which includes all tissues other than the dermal and the vascular system. The fundamental or ground tissue of shoots and roots is found in the cortex (between the dermal and vascular system) and the pith (when present, at the center surrounded by the vascular system). If the relatively mature, above ground, primary shoot is cut across at a right angle to its longitudinal axis, the shoot can be observed as a series internal areas with more or less concentric circular borders. From the circumference to the center of this roughly circular transverse section, the most peripheral area includes the epidermis (including dermal tissue) followed by the cortex (including ground tissue) followed by the vascular system (including vascular tissue) followed, if present, by a central pith (including ground tissue).
    It is the meristem called the procambium (or provascular system) that gives rise to the primary xylem and phloem or the primary vascular system. The procambium can give rise to one or more often slender cylindrical vascular strands. A vascular strand (also called a vascular bundle) is simply a strand of xylem and phloem with associated tissues. As the procambium arises from an apical meristem and gives rise to the vascular tissue, it may appear in the stem or root as a single central cylindrical strand or it may appear as a cylindrical ring of individual cylindrical strands surrounding the center. Vascular tissue can initially arise in a single ring of vascular strands that may remain separate from each other or other strands may be added to create a more or less continous cylinder. In other cases, a single ring of vascular strands may initially develop but other strands may be added so that the strands appear to become 'scattered' throughout the stem or root. In any case, the protoderm surrounds the ground meristem, and the provascular (procambial) strands are embedded in it. Vascular tissues and associated ground tissues of the stem or root is generally referred to as the vascular cylinder (or stele).
    Like most cells involved in primary growth and development other than the meristematic cells, most primary xylem cells not only increase in number by mitosis and subsequently become elongated, but also undergo cellular differentiation. The first formed primary xylem of vascular plants is called the protoxylem, while the later formed primary xylem is called the metaxylem. The differentiation of protoxylem and metaxylem is unique to all living vascular plants and considered a shared, derived character of the lineage called Tracheophyta Sinnott. Protoxylem cells are generally smaller in diameter than those of the metaxylem. Aften initiation of cell division (mitosis) by the apical meristem, the cells of the shoot (stem), leaf, or root undergo a period of elongation. The protoxylem often matures before intensive elongation, while the metaxylem, appearing after the protoxylem, develops while the shoot is elongating, and matures after elongation is completed. Of considerable importance in the classification of fossil vascular plants is the direction of development of successive cells of the procambium as the often well preserved primary xylem matures. In exarch primary xylem maturation, the first protoxylem cells to acquire secondary walls occur at the outermost edge of an often cylinderic procambium. The earliest developing conductive cells (the protoxylem) are external to those developing later (the metaxylem), with maturation from the outside toward the inside. The term 'exarch' refers to maturation beginning from the outside. This is found in the roots of all vascular plants and common in the shoots (stems) of fossils or their living descendants in the lineage called the lycophytes. In some of the oldest euphyllophyte fossils with xylem in only a single strand without a pith, there is centrarch primary xylem maturation, where the earliest developing conductive cells (the protoxylem) are central and surrounded by or embedded in those developing later (the metaxylem), with maturation from the inside toward the outside. In more derived (advanced) members of the euphyllophyte group, there can also be mesarch xylem maturation, with development starting at the first mature xylem elements and progressing both inwardly and outwardly. In this type of xylem maturation, the groups of protoxylem cells become surrounded by the metaxylem. The terms 'centrarch' and 'mesarch' refer to maturation beginning in the center or middle. In a centrarch xylem, the earliest maturing protoxylem is surrounded by the later maturing metaxylem. In a mesarch xylem, the protoxylem is flanked on two sides by metaxylem. In euphyllophytes, there is usually never or only very rarely exarch xylem maturation. However, lycophytes xylem maturation is almost exclusively exarch. In the euphyllophyte group (e.g., among horsetails and the now more common seed plants), there is also endarch xylem maturation, where the earliest developing conductive cells are internal to those developing later, with maturation solely from the inside toward the outside. The term 'endarch' refers to maturation beginning from the inside. This is often believed to be the most highly advanced type of xylem maturation. Although centrarch and endarch both involve centrifugal (inside to outside) maturation, centrarch applies when xylem forms only one strand in the stem with no pith. Fossil evidence supports that either centrarch or exarch may be the type of primary xylem maturation of the earliest (oldest) vascular plants. The centrarch xylem is probably the oldest, although some botanists consider exarch the oldest (Gifford and Foster, 1989).

  • Gifford, E. M. and A. S. Foster (1989) Morphology and Evolution of Vascular Plants, Third Edition. W. H. Freeman and Company. New York.

    Vascular plants (= tracheophytes)
    As the lineage leading to vascular plants (with a well developed water and food conducting tissue, respectively comprising the xylem and phloem) continued to diversify (change through evolution), it is more certain that the young (immature) sporophytes remained at least initially dependent on and nurtured by the gametophytes, but as the sporophytes matured, they could soon become free-living or independent of the gametophytes, allowing them to become larger and more branched with more sporangia. With the eventual independence of the sporophytes following an initial, short-lived dependence upon gametophytes, an evolutionary trend became established. As the sporophytes became independent earlier in the life cycle, they were free to grow larger and more branched, often bearing more and more sporangia, while the gametophytes could become smaller and smaller, because they were needed only in the short, initial stages of sporophyte development. Therefore, this eventual early independence of sporophytes from gametophytes led to gradually larger, more dominant, much branched sporophytes, bearing multiple sporangia, in contrast to the gametophytes becoming smaller and in certain heterosporous plants, eventually becoming so minute that they could develop within separate male and female spores (microspores and megaspores, respectively). The evolutionary trend of progressively larger sporophytes and progressively smaller gametophytes is most evident in the later evolution of seed plants, where male and female gametophytes have become minute and more and more dependent upon the sporophytes. These gametophytes can become so small that they can easily become sheltered within spores inside often multiple, heterosporous (male and female) sporangia produced by dominant, sometimes huge, and much branched sporophytes.
    However, in all currently living seed plants, the male gametophyte becomes only partially developed within the pollen just prior to dispersal. After the pollen reaches its appropriate destination, the male gametophyte can emerge from the pollen grain as a narrow, filamentous structure called the pollen tube that can extend considerably in length (sometimes penetrating or growing through tissue) in order to transport the sperm contained within it and fertilize the egg cell of the female gametophyte.
    The evolutionary trend of progressive reduction of the gametophyte in vascular plants can be related to the progressive origins of different degrees of the heterosporous condition. For example, the germination of isospores of homosporous plants that develop into distinct male and female (dioecious) gametophytes can be thought of as transitional to different (also transitional) degrees of the heterosporous condition. According to Bateman and DiMichele (1994), this includes the production of dioecious gametophytes progressively followed by the apportionment of two different (dimorphic) male and female spore types (often smaller microspores and larger megaspores, respectively) within two different (dimorphic) male and female sporangia (the heterosporangiate condition), the retention of the gametophyte within the spore wall (the endosporous condition), the retention of the megaspore(s) within the female sporangia (the endomegasporangiate condition), and the reduction of the contents of the female sporangium to a single functional (viable) megaspore (the monomegasporous condition). With the more derived development of the endosporous condition, the male and female gametophytes (respective microgametophytes and megagametophytes) can become so minute that they at least temporarily can remain enclosed within their respective male and female spore walls that at least temporarily remain attached within their respective male and female sporangia (respective microsporangium and megasporangium). This can results in male and female reproductive organs being borne on the same sporophyte (the monoecious condition) or on separate sporophytes (the dioecious condition). The basic trends in presently living seed plants and extinct plants with seed-like reproduction appear to be (1) heterosporous development, (2) reduction of the number of megaspores to one, and (3) enclosure of the megasporangia in a protective layer of tissue. In this summerized process, the megagametophytes becomes successively reduced with the smallest ones of all found in flowering seed plants.
    The trend in the reduction of the megagametophyte culminates in certain seed plants (including Gnetum and Welwitschia plus all angiosperms) with the loss of the archegonium; and such a reduced megagametophyte is called an embryo sac.
    It was until very recently (since 2018) widely assumed that the gametophyte dependent 'unbranched sporophyte' type with a single terminal sporangium likely predated the relatively independent branched sporophyte type with multiple sporangia. Because the evolution of the transition from one type to the other is still unclear, it is not yet possible to judge whether branching with multiple sporangia and the independence of the sporophyte occurred together or were separate processes, and if separate, the order of evolution of these characters remains unclarified. Although proposed to be unique to 'polysporangiophytes' (meaning plants with many sporangia), the definition of the features, such as an independent sporophyte with branching and multiple sporangia, remains ambiguous, because no transitional fossils are known. Consequently, these features could be considered an example of 'cryptic' characters, even though they are widely used in identification, especially for presently living land plants, and are easily observed in the wild or in the fossil record. Furthermore, in the past the comparison of so-called 'unbranched' and 'branched' sporophyte types may not have always been interpreted correctly. The entire 'unbranched sporophyte' of liverworts, mosses, and hornworts has often been described as composed of three parts: the foot at the base of the sporophyte that attaches it to the gametophyte, the terminal capsule (sporangium), and the seta, when present, that appears as an unramified or straight stem-like projection between the foot and the capsule. In the past, the seta bearing a single capsule was thought to be representative of descendants of the earliest land plants, which was often been assumed to have been further elaborated as the highly branched stem, bearing multiple sporangia, of the more complex land plants. However, the sporophyte of liverworts, mosses, and hornworts could be alternatively interpreted as a stem-less footed sporangium (the sporogonium) with or without a seta (the stalk of the sporangium) between a slightly swollen attachment to the gametophyte (the foot) and a single terminal capsule (the sporangium). The seta, if present, may be no more than a sporangial stalk, and the foot an embryonic transfer tissue at the base of the sporangium. In other words, the stalk-like seta (when present) may not be an unbranched stem, but it may merely be part of a stalked sporangium embedded in the gametophyte via nutrient (or growth factor) transfer tissue called the foot. If bryophytes are assumed to be monophyletic with hornworts first branching followed by liverworts sister to mosses, the seta becomes a possible shared derived character of a setaphyte lineage comprising liverworts and mosses. If the assumed basal (first branching) possition of hornworts within bryophytes is correct, it could be expected that hornworts would resemble the common ancestor of bryophytes more than the common ancestor of liverworts and mosses. It has been known for quite some time that hornworts, although with an unbranched sporophyte attached to a gametophyte, possess certain similarities to vascular plants that are not found in liverworts and mosses. These similarities may be due not to the possibility that hornworts share a direct common ancestor with vascular plants but due to the possibility that the common ancestor of bryophytes had greater resemblance to the common ancestor of vascular plants than to the common ancestor of liverworts and mosses. The evolutionary trend in bryophytes of the gametophyte would be expected to involve succesively greater elaboration, while that of the sporophyte would be expected to involve succesively greater reduction. On the other hand, the evolutionary trend in vascular plants of the gametophyte would be expected to involve succesively greater reduction, while that of the sporophyte would be expected to involve succesively greater elaboration. Less like setophytes (liverworts and mosses) and more like vascular plants, the sporophyte of hornworts, although unbranced, is more elaborated and somewhat less dependent on a more reduced gametophyte. The hornwort gametophyte itself is a simple, flattened (doriventral) disc-like body not differentiated into root, stem or leaves called a thallus that is more similar to many gametophytes of vascular plants than the more elaborated gametophytes of mosses and some liverworts. If bryophytes are indeed monophyletic and sister to vascular plants, all that can currently be said is that the ancestor of embryophytes might have had a more complex sporophyte phase than would be expected from the past assumption that liverworts are sister to the rest of embryophytes; however, this results in more life cycle possibilities for the common ancestor of embryophytes, including dimorphic (dissimilar) sporophyte and gametophyte phases with gametophyte dominance, dimorphic phases with sporophyte dominace, and dimorphic or isomorphic (similar) phases with co-dominance. Also, the earliest fossils, which have been assumed to represent stem vascular plants, might be stem group representatives of a more complex embryophyte ancestor or even a more complex bryophyte ancestor.

    Verbascum - L. - Genus Name
    Family Scrophulariaceae
    Only Verbascum thapsus is reported to be used as a medicinal in New Mexico. In the Old World, the less common species Verbascum blattaria (Moth mullein) and Verbascum virgatum (Wand mullein or Large flowered mullein) are said to have similar medicinal properties. All these species can be naturalized in North America and all originate from the Old World. Although less common than Verbascum thapsus, the species Verbascum blattaria and Verbascum virgatum can also be found in New Mexico.

    Verbascum thapsus - L. - Species Name
    Family Scrophulariaceae
    Native Americans adopted the European introduced plant for coughs. With a growth form superficially similar to native tabacco, this mullein is reported to have been smoked in the New World by some Spanish people and certain indigenous tribes. It has been smoked by itself as a tabacco substitute or combined with tabacco in a smoking mixture. Spanish New Mexicans claim that, besides being pleasurable, the inhaled smoke of the dryed leaves of Verbascum thapsus in corn husk cigarettes is good for asthma. Spanish in the USA and Mexico commonly use this species both externally and internally in pulmonary diseases.
    In Europe under the name of Cow's or Bullock's Lungwort, Verbascum thapsus was once used as a remedy for various cattle diseases, especially those of the lungs. In the first century AD, Dioscorides administered the roots for human diseases of the lungs, while Pliny gave its leaves to broken-winded horses.
    This species is traditionally used originally in Europe to treat asthma, bronchitis, and kidney infections. The thick woolly covering of trichomes ('hairs') is reported to contain an arsenal of defensive (somewhat toxic) chemicals, including coumarin and the insecticide rotenone. Although cited in several references (e.g., Duke & Foster's Medicinal plant feild guide; Duke's Green Pharmacy; and a series of books called 'The Wealth of India'), the presence of rotenone in mullein probably needs further varification. There is no mention of its presence in the review by Tatli & Akdemir (2004). Although the presence of rotenone is questionable (likely false), mullein seeds are known to stun fish because they are very high in saponins (more specifically, oleanane type triterpene glycosides with a soap-like action). The reported skin irritation from leaves is likely due to the coarse trichomes rather than toxic chemicals.
    o Tatli & Akdemir (2004) Chemical Constituents of
    Verbascum L. Species, FABAD J. Pharm. Sci., 29, 93-107.
    See also Verbascum thapsus as tobacco.
    There is a striking similarity in folk uses for respiratory complaints of Verbascum (commonly called Gordolobo) and Anaphalis, Gnaphalium, and Pseudognaphalium (commonly called Gordolobo Mejicano).
    Although the latter three genera together with Antennaria are closely related within the family Asteraceae and tribe Gnaphalieae and, therefore, expected to be used for similar purposes, they are unrelated to Verbascum.
    Verbascum thapsus L. appears in the writings of Dioscorides, Pliny, and Galen as used for many health conditions, especially for conditions of the chest or lungs. After its introduction from Europe, many Native Americans began using it in areas of North America where it became established in the wild. Dried leaves were reported (Herrick, 1977) to have been smoked by the Iroquois for catarrh, asthma, and bad hiccoughs. The leaf was reported (Smith, 1923) to have been gathered and smoked by the Menominee as an Indian tobacco. The dried leaves were reported (Smith, 1933) to have been smoked in pipe for asthma and smudged leaves inhaled for catarrh by the Potawatomi. The Thompson were reported (Turner, et al., 1990) to have used the leaf for smoking. In the southwest USA, it is reported to have been smoked by Mexican Americans of Colorado and New Mexico, as well as the Hopi, Isleta, and Navajo. In New Mexico, the Spanish were known by Native Americans to have gathered the leaves of this species (commonly known as Mullein, Punchon, Tobaco Cimarron, or Gordolobo) to smoke for the relief of asthma and bronchitis. The Isleta were reported (Jones, 1931) to have used this species as a ceremonial tobacco. The dried leaves of Verbascum thapsus have been reported (Vestal,1952) to have been smoked in corn husks as a psychological aid by Native Americans in New Mexico to clear a lost mind. It has also been claimed that smoking Nicotiana spp. (species of the same genus as commericial tobacco) is 'a practice likely to help one afflicted with fits' [Oklahoma folk lore cited by Curtin (1965)]. Nicotiana spp. sometimes combined with other plants have been widely reported to have been smoked by Native Americans to clear the mind or treat mental problems and even convulsive disorders. Some of these other plants include Verbascum thapsus and Macromeria viridiflora DC. used by themselves or combined with a different species. For example, the leaves of Verbascum thapsus were reported (Whiting, 1939) to have been smoked by the Hopi to treat mental illness. These leaves were reported to have been smoked with those of Macromeria viridiflora DC. (Gianttrumpets in family Boraginaceae) as an anticonvulsive for 'fits' or for craziness and witchcraft. Macromeria viridiflora (= Onosmodium thurberi A. Gray) is also reported to have been ceremonially smoked with wild Nicotiana spp.
    In Spain, the name Gordolobo was first applied to Verbascum thapsus (= Punchon or Tobaco cimarron in New Mexico). Although this plant is unrelated to Anaphalis and some of the aromatic species of Antennaria, Gnaphalium, and Pseudognaphalium in the tribe Gnaphalieae (Cass.) Lecoq & Juillet of the family Asteraceae, the name Gordolobo Mejicano was applied by Francisco Hernandez in Mexico to certain plants of the genus Pseudognaphalium used by the Aztecs for some of the same purposes as the Spanish or Old World genus Verbascum. Species of these above genera of Gnaphalieae known to grow wild in New Mexico are reported in other parts of North America north of Mexico to have been smoked like a tobacco by people of various Native American tribes often for treatment of respiratory conditions, including allergic rhinitis (= hay fever), asthma, and dry cough or used as a smudge or fumigant (Pennacchio et al., 2010). In southeastern North America north of Mexico, the common name Rabbit tobacco in English for species of the related genera Gnaphalium, Pseudognaphalium, and Gamochaeta "indirectly records the indigenous practice of mixing the leaves with tobacco (Nicotiana spp. inserted) for smoking" (Austin, 2006). If according to this author "virtually all who live near" these plants have used them, why are there little or no reports on the their use among Spanish New Mexicans? The species Anaphalis margaritacea is called Gordolobo in Baja California Sur; and people of recent Mexican descent in southern New Mexico and Arizona have used it as a brewed tea sweetened with honey for a raspy voice or sweetened with sugar for dry cough (Kay, 1996). Various species of the related genus Pseudognaphalium are commonly used for similar purposes right across the border by various indigenous people in Northern Mexico. One of these species Pseudognaphalium leucocephalum (Gray) A. Anderberg (= White rabbit-tobacco, also commonly called Everlasting, Gordolobo, Lemon cudweed, Lampaquate, or Manzanilla del rio) is also found in southwestern New Mexico. Although not found in New Mexico, Pseudognaphalium viscosum (Kunth) Anderberg (one of the closest relative to Pseudognaphalium leucocephalum) is commonly used for respiratory complaints in Mexico and among some people of recent Mexican descent in Texas. In this genus, the most common and widely distributed species Pseudognaphalium macounii (Greene) Kartesz (= Macoun's rabbit-tobacco) in New Mexico is also reported to be commonly used by indigenous people right across the border. From a global comparison of closely related plants, it has become apparent that knowledge of the uses of species of the genera Gnaphalium or Pseudognaphalium can be extended to the application of other closely related plants, such as Anaphalis and aromatic species of Antennaria for lung or respiratory conditions. One plains tribe north of Mexico even used the dried powdered flowers of Anaphalis margaritacea as a strengthening medicine placed on the soles of horse's hoofs and blown between the horse's ears to make them long-winded, enduring, and spirited. This is mentioned because there have been reports that the flowering tops of Pseudognaphalium obtusifolium (Rabbit tobacco) of the eastern USA have been smoked or even inserted in pillows to ease the breathing of sleeping Anglos suffering from asthma. See also similar European uses of the unrelated species Verbascum thapsus for broken-winded horses. Anaphalis margaritacea (L.) Benth. & Hook. f. (= Western pearlyeverlasting), which may be one of the first herbs from North America to have been cultivated in Europe, where it is now naturalized in Britain, was used for smoke therapies in Suffolk, Britain, where the leaves were smoked for treating coughs and headaches (Jobson, 1967). According to Gerarde (1633), the smoke and fumes of dried herbs were inhaled for coughs, for headaches, and for cleansing 'inward parts.' Aside from related Native American uses of species of Antennaria in the prairies of the North American Great Plains, Antennaria neglecta Greene (= Field pussytoes) and Antennaria plantaginifolia (L.) Richards. (= Woman's tobacco) were highly valued by Eclectic doctors for chest complaints. Constantine Rafinesque wrote in the Medicinal Flora of the United States that these two species are "pectoral, used in coughs, fevers, bruises, inflammations, debility; ..." Certain undesignated species of Antennaria are also reported to have been used by some Native Americans as cough or cold remedies (Steedman, 1928; Swanton, 1928; Taylor, 1940).
    The leaves of the aromatic species Antennaria rosea (D.C. Eaton) Greene is reported to have been used by Native Americans outside New Mexico as a mixture with tobacco (Johnston, 1987), kinnikinnick (Nickerson and Gifford, 1966), or as a fumigent (using dried, powdered roots) on hot coals (Turner, Bouchard, and Kennedy, 1980). This is a species with three intergrading subspecies that can be found in the northern mountains of New Mexico. See also similar uses of the unrelated species Verbascum thapsus as a tobacco.

  • Austin DF. (2006) Florida Ethnobotany, CRC Press, Boca Raton, London, New York, Washington, D.C.
  • Gerarde, John (1633) Herbal or General History of Plants (John Gerard's The herball or Generall historie of plantes).
  • Jobson, A. (1967) In Suff olk borders. London: Robert Hale.

    See also:

  • Castetter, Edward F. and Willis H. Bell (1942) Pima and Papago
    Indian Agriculture, Albuquerque: University of New Mexico Press,
    First Edition.
  • Chamberlin, Ralph V. (1911) The Ethno-Botany of the Gosiute
    Indians of Utah, Memoirs of the American Anthropological
    Association 2(5):331-405.
  • Curtin, L.S.M. (1965) Healing herbs of the upper Rio Grande,
    Southwest Museum, Los Angeles, California.
  • Grinnell, George Bird (1972) The Cheyenne Indians - Their History
    and Ways of Life Vol.2, Lincoln, University of Nebraska Press.
  • Hart, Jeffrey A. (1981) The Ethnobotany of the Northern Cheyenne
    Indians of Montana, Journal of Ethnopharmacology 4:1-55.
  • Herrick, James William (1977) Iroquois Medical Botany. State
    University of New York, Albany, PhD Thesis
  • Johnston, Alex (1987) Plants and the Blackfoot, Lethbridge, Alberta,
    Lethbridge Historical Society.
  • Jones, Volney H. (1931) The Ethnobotany of the Isleta Indians,
    University of New Mexico, M.A. Thesis.
  • Palmer, Gary (1975) Shuswap Indian Ethnobotany, Syesis 8:29-51.
  • Pandey, S. P., P. Shahi, K. Gase, and I. T. Baldwin (2008)
    Herbivory-induced changes in small-RNA transcriptome and
    phytohormone signaling in Nicotiana attenuata, Proceedings of the
    National Academy of Sciences 105(12): 4559-4564.
  • Pennacchio, M., Jefferson, L., & Havens, K. (2010) Uses and abuses of
    plant-derived smoke: Its ethnobotany as hallucinogen, perfume, incense,
    and medicine. Oxford University Press.
  • Skibbe, M., N. Qu, I. Galis, and I. T. Baldwin (2008) Induced plant
    defenses in the natural environment: Nicotiana attenuata WRKY3
    and WRKY6 coordinate responses to herbivory, The Plant Cell 20:
    1984-2000.
  • Smith, Huron H. (1923) Ethnobotany of the Menomini Indians,
    Bulletin of the Public Museum of the City of Milwaukee 4:1-174
  • Smith, Huron H. (1933) Ethnobotany of the Forest Potawatomi
    Indians, Bulletin of the Public Museum of the City of Milwaukee
    7:1-230.
  • Turner, Nancy J., Laurence C. Thompson and M. Terry Thompson,
    et al. (1990) Thompson Ethnobotany: Knowledge and Usage of
    Plants by the Thompson Indians of British Columbia, Victoria,
    Royal British Columbia Museum
  • Wyman, Leland C. & Stuart K. Harris (1941) Navajo Indian
    Medical Ethnobotany, The University of New Mexico Bulletin,
    Anthropological Series, Vol. 3, No. 5, June.
  • Vestal, Paul A. (1952) The Ethnobotany of the Ramah Navaho,
    Papers of the Peabody Museum of American Archaeology and
    Ethnology 40(4):1-94.
  • Whiting, Alfred F. (1939) Ethnobotany of the Hopi, Museum
    of Northern Arizona Bulletin #15.

    Verbenaceae - Jaume St-Hil. - Famiy Name
    Many of its genera [e.g., Tetraclea Gray & Vitex L.]
    have been transferred to Lamiaceae Martynov, reducing
    Verbenaceae to former subfamily Verbenoideae.
    Although traditionally placed in family Verbenaceae
    [as in Allred (2010)], the genus Tetraclea [= Clerodendrum L.],
    including species Tetraclea coulteri A. Gray
    [= Clerodendrum coulteri (A. Gray) Govaerts],
    is likely in subfamily Ajugoideae Kosteletzky of the
    mint family (Lamiaceae).
    Furthermore, Tetraclea should probably remain separate
    from the redefined genera Clerodendrum sensu stricto
    and Volkameria L. (including Huxleya),
    with North American Tetraclea more closely related to
    tropical American genera Aegiphila and Amasonia in a
    separate New World lineage (Steane, et al., 2004).
    See additional info under Clerodendrum.
    See also: Steane, D. A., de Kok, R. P., & Olmstead, R. G. (2004)
    Phylogenetic relationships between Clerodendrum (Lamiaceae)
    and other Ajugoid genera inferred from nuclear and chloroplast
    DNA sequence data. Molecular Phylogenetics and Evolution,
    32(1), 39-45.

    Verbesina encelioides - (Cavanilles) Bentham & Hooker f. ex Gray - Species Name
    Family Asteraceae
    Significant antibacterial, antifungal, antiviral, antitumor, hypoglycaemic and
    anti-implantation activities have been demonstrated from various fractions of
    these plants.

    References:

  • Ateya, Okarter, T., Knapp, J., Schiff, P. Jr. and Slatkin D. (1982) Flavonoids of Thelesperma megapotamicum, Planta Medica, Vol. 45, No. 8, pp. 247-248.
  • Barboza, G. E., Cantero, J. J., Nunez, C., Pacciaroni, A., & Ariza Espinar, L. (2009) Medicinal plants: A general review and a phytochemical and ethnopharmacological screening of the native Argentine Flora. Kurtziana, 34(1-2), 7-365.
  • Bohlmann, F., Burkhardt, T., and Zdaro, C. (1973) Naturally occurring acetylenes, Academic Press, London.
  • Bohlmann, F., M. Ahmed, M. Grenz, R.M. King, and H. Robinson (1983) Bisabolene derivatives and other constituents from Coreopsis species, Phytochemistry 22: 2858-2859.
  • Bown, D. (1995) Encyclopaedia of Herbs and their Uses, Dorling Kindersley, London.
  • Burlage, Henry M. (1968) Index of plants of Texas with reputed medicinal and poisonous properties, Austin Texas.
  • Chevallier, A. (1996) The Encyclopedia of Medicinal Plants, Dorling Kindersley, London.
  • Coffey, T. (1993) The History and Folklore of North American Wild Flowers.
  • Crowe, A. (1990) Native Edible Plants of New Zealand, Hodder and Stoughton.
  • Duke, J. A. & Ayensu, E. S. (1985) Medicinal Plants of China Reference Publications, Inc.
  • Elmore, Francis H. (1944) Ethnobotany of the Navajo. Sante Fe, NM. School of American Research.
  • Facciola, S. (1990) Cornucopia - A Source Book of Edible Plants, Kampong Publications.
  • Foster. S. & Duke. J. A (1990) A Field Guide to Medicinal Plants. Eastern and Central N. America. Houghton Mifflin Co.
  • Funk, VA, et al. (2005) Everywhere but Antarctica: Using a supertree to understand the diversity and distribution of the Compositae. Biol. Skr. 55: 343-374.
  • Grieve, M. (1981, 1984) A Modern Herbal, Penguin
  • Hamel, Paul B. and Mary U. Chiltoskey (1975) Cherokee Plants and Their Uses -- A 400 Year History. Sylva, N.C. Herald Publishing Co.
  • Hansen, H.V. (1991) Phylogenetic studies in Compositae tribe Mutisieae, Opera Bot. 109, 1-50.
  • Hocking, George M. (1956) Some Plant Materials Used Medicinally and Otherwise by the Navaho Indians in the Chaco Canyon, New Mexico. El Palacio 56:146-165
  • Hsu, H.-Y. (1986) Oriental Materia Medica. Keats, New Canaan, CT.
  • Huang, Y.-P., & Ling, Y.-R., (1996) Economic Compositae in China. In P.D.S. Caligari & D.J.N. Hind (eds). Compositae: Biology & Utilization. Proceedings of the International Compositae Conference, Kew, 1994. (D.J.N. Hind, Editor-in-Chief), vol. 2. pp. 431-451. Royal Botanic Gardens, Kew.
  • Kay, Margarita Artschwager (1996) Healing with plants in the American and Mexican West, University of Arizona Press, Tucson.
  • Kim, K.-J., et al., Choi, K.-S., & Jansen, R. K. (2005) Two chloroplast DNA inversions originated simultaneously during the early evolution of the sunflower family (Asteraceae). Mol. Biol. Evol. 22: 1783-1792.
  • Kunkel. G. (1984) Plants for Human Consumption. Koeltz Scientific Books
  • Lewis, W.H. and M.P.F. Elvin-Lewis (2003) Medical botany, plants affecting human health, 2nd edition, John Wiley & Sons, Inc.
  • Linares, E. & Bye, R. (1987) A study of four medicinal plant complexes of Mexico and adjacent United States, Journal of Ethnopharmacology 19: 153-183.
  • Martinez, M. (1959) Las plantas medicinales de Mexico, 4th ed., Botas, Mexico City.
  • Mills, S (1988) Dictionary of Modern Herbalism.
  • Panero, J. L., and V. A. Funk (2008) The value of sampling anomalous taxa in phylogenetic studies: major clades of the Asteraceae revealed. Mol. Phylogenet. Evol. 47: 757-782.
  • Reagan, Albert B. (1929) Plants Used by the White Mountain Apache Indians of Arizona. Wisconsin Archeologist 8:143-61.
  • Reichling, J., and U. Thron (1989) Comparative study on the production and accumulation of unusual phenylpropanoids in plants and in vitro cultures of Coreopsis tinctoria and C. lanceolata, Pharmaceutisch Weekblad 11: 83-86.
  • Rogers, Dilwyn J (1980) Lakota Names and Traditional Uses of Native Plants by Sicangu (Brule) People in the Rosebud Area, South Dakota. St. Francis, SD. Rosebud Educational Scoiety
  • Singh, Dr., G. & Kachroo, Prof., Dr., P. (1976) Forest Flora of Srinagar, Bishen Singh Mahendra Pal Singh.
  • Smith, Huron H. (1928) Ethnobotany of the Meskwaki Indians. Bulletin of the Public Museum of the City of Milwaukee 4: 175-326.
  • Spellenberg, R., L. McIntosh, & L. Brouillet (1993) New records of angiosperms from southern New Mexico, Phytologia 75:224-230.
  • Stevenson, Matilda Coxe (1915) Ethnobotany of the Zuni Indians. SI-BAE Annual Report #30.
  • Sturtevant, William (1954) The Mikasuki Seminole: Medical Beliefs and Practices. Yale University, PhD Thesis.
  • Swank, George R. (1932) The Ethnobotany of the Acoma and Laguna Indians, University of New Mexico, M.A. Thesis.
  • Usher, G. A (1974) Dictionary of Plants Used by Man, Constable.
  • Vestal, Paul A (1940) Notes on a Collection of Plants from the Hopi Indian Region of Arizona Made by J. G. Owens in 1891. Botanical Museum Leaflets (Harvard University) 8(8):153-168
  • Vestal, Paul A. (1952) The Ethnobotany of the Ramah Navaho, Papers of the Peabody Museum of American Archaeology and Ethnology 40(4):1-94.
  • Vestal, Paul A. and Richard Evens Schultes (1939) The economic botany of the Kiowa Indians. Cambridge, Mass.: Botanical Museum.
  • Watson, Linda E. (1996) Molecular systematics of Tribe Anthemideae (Asteraceae). Pp. 341-348, In: Proceedings of the 1994 Kew International Compositae Conference, C. N. Hind (ed.). Royal Botanic Gardens, Kew.
  • Watson, Linda E., Timothy M. Evans, and Tatiana Boluarte (2000) Molecular phylogeny of Tribe Anthemideae (Asteraceae), based on chloroplast gene ndhF. Molecular Phylogenetics and Evolution. 15: 59-69.
  • Wyman, Leland C. and Stuart K. Harris (1941) Navajo Indian Medical Ethnobotany, The University of New Mexico Bulletin, Anthropological Series. Volume 3. No. 5.
  • Zigmond, Maurice L. (1981) Kawaiisu Ethnobotany, Salt Lake City, University of Utah Press.

    .

    .

    .

    .

    .

    .

    .

    .

    .

    .

    .

    .

    .

    .

    .

    .

    .

    .

    .

    .

    .

    .

    .

    .

    .

    .

    .

    .

    .

    .

    .

    .

    .

    .