Taxonomy of the Giant Horsetails



"Sine examine microscopico nulla scientia Equisetorum!" (Milde, 1867, p. 213)


    Plant taxonomists currently recognize two species of giant horsetail and a hybrid between them: Equisetum giganteum L., E. myriochaetum Schlecht. and Cham., and E . x schaffneri  Milde (E. giganteum x E. myriochaetum ) ( Hauke, 1963 ).  Prior to Hauke's ( 1963 ) work, taxonomists recognized a relatively large number of species and varieties of giant horsetails. For example, Milde, in his 1867 monograph of Equisetum , recognized seven species of tropical giant horsetails.  For a comprehensive synonymy for these and the other Equisetum species, please refer to the Checklist of World Ferns .

    The giant horsetails are some of the least known of the 15 species in the genus Equisetum.  All of the giant horsetails are members of the subgenus Hippochaete within the genus Equisetum .  The subgenus Hippochaete is characterized by species having sunken stomata (in contrast to the raised stomata of subg. Equisetum ).  Most of the seven members of the subgenus Hippochaete , including the giant horsetails and the familiar temperate "scouring rush" E. hyemale , have have tough evergreen stems.  However, the giant horsetails are the only Equisetum species that have stems that are both evergreen and regularly (i.e. radially symmetrically) branched.  Furthermore, the regular branching habit of the giant horsetails is  unique in the subgenus Hippochaete . (Hauke, 1963 ; 1978 ).
 (For more detailed background information on the genus Equisetum , I recommend the information and links provided on the website of the U.K. National Collection of Equisetum .)

    Because the three giant horsetails appear similar in overall habit, and because Equisetum species exhibit considerable morphological plasticity ( Hauke, 1963 ; Schaffner, 1928 ), more stable anatomical characters are used to distinguish between the species (see table below).  The most important diagnostic characters (branch ridge patterns, stomatal patterns, and endodermal patterns) can only be observed under high magnification and many of characters of E. x schaffneri overlap with its parent species ( Hauke, 1963 ).  Therefore, accurate identification of giant horsetails can be problematic.  As a result, both dried specimens in herbaria (Stolze, 1983 ; Husby, personal observation) and living specimens in botanical gardens (Moyroud, 1991 ; Husby, personal observation) are often misidentified (for further discussion of misidentfication, see Notes, Observations, and Ideas ).

    Below is a synopsis of the key characteristics useful for distinguishing between the giant horsetails.  The information is from Dr. Richard L. Hauke's ( 1963 ) monograph of Equisetum subgenus Hippochaete , unless otherwise referenced.  Dr. Hauke's work was based upon intenstive study and analysis of a large number of herbarium specimens.


E. giganteum
E. myriochaetum
E. x schaffneri
Type locality
Jamaica (or Hispaniola 1 )
Vera Cruz, Mexico
Orizaba, Mexico
Distribution 2
( more detailed distribution information )
Cuba, Jamaica, Hispaniola, Central American (Guatemala, El Salvador, Honduras, Nicaragua ( Luis Diego Gómez, 1985 ), Costa Rica, Panama), and South America(Venezuela, Colombia, Ecuador, Peru, Bolivia, Brazil, Paraguay, Uruguay, Chile, and Argentina)
Mexico, Central America, Colombia, Ecuador, and Peru
Mexico3 , Central America, Colombia, Venezuela 3 , Ecuador, Peru
Habitat
"...along rivers or in swampy places, usually shaded"
"In swampy places or along rivers and streams, ususally at the edge of or within forested areas..."
"...in springy, marshy places, or along rivers and streams."
Altitudinal range
150 - 2600 m
200 - 3000 m
500 - 3000 m
Maximum stem height 4
5 m
8 m
4.5 m
Maximum stem diameter 4
2.4 cm
1.8 cm
2.2 cm
Branch ridge pattern
"...square or flattened in profile, in Brazil, Peru, and southern South America tending to be irregular."
sawtooth pattern oriented apically
"...sawtooth to irregular."
Main stem stomatal pattern
"...in bands of 3-4 (rarely 2-3 or 4-5)"
"Stomata in one line on each side of the groove."
"...in bands of 1-2 (occassionally 2-3)"
Endodermal arrangement
"Cross section with separate endodermis around each vascular bundle (individual endodermises)"
"...double common..."
individual endodermises
Sheath teeth persistence
"mostly persistent" ( Tryon, R. M. and Tryon, A. F., 1982 )
"...usually thin and brown to white, drying and breaking off to produce a clipped appearance at the top of the sheath, or (especially in South America), the bases or much of the teeth persisting"
"...mostly smoothly shed."
Cones
have a "short (about 0.5 mm) but distinct apiculum"
"blunt, or the branch cones frequently with a slight apiculum"
"acute or with slight apiculum"



1.  Proctor ( 1985 ) and Lellinger ( 1989 ) gave the type locality as "presumably...Hispaniola", whereas Hauke (1963) gave the locality as Jamaica.

2.  Only two other Equisetum species reach the tropics in Latin America, E. hyemale (which reaches Guatemala) and E. bogotense (From Costa Rica to Chile and Argentina) ( Moran and Riba, 1995 ; Stolze, 1983 ).

3.  Interestingly, this hybrid is found in Mexico, where its parent, E. giganetum, is not known to be present, and in Venezuela, where its other parent E. myriochaetum is not known to be present.  Hauke (1963) hypothesized that this unexpected phenomenon may be due to occassional viable spores being produced by E. x schaffneri and the resulting plants persisting vegetatively.  Viable spores have been observed for other Equisetum subg. Hippochaete hybrids ( Krahulec et al., 1996 ), so this hypothesis appears plausible.   Rolla M. Tryon and Alice F. Tryon ( 1982 ) have suggested that E. x schaffneri may not be a hybrid at all, but rather an intermediate form of a single polymorphic horsetail species E. giganteum (with the currently recognized E. giganteum and E. myrichaetum representing extremes of this species). However, Hauke( 1963 ) notes in support of E. x schaffneri's hybrid status that its spores are not viable in contrast to those of the other two species.  The status of E. x schaffneri could probably be settled by attempting to synthesize E. x schaffneri from a cross of E. giganteum and E. myriochaetum in the laboratory, as has been done for other subgenus Hippochaete hybrids ( Ducket, 1979 ).  Molecular taxonomic analysis would likely shed additional light on this question.

4.  These numbers are based on collector's notes and measurements of herbarium specimens (See How large are giant horsetails? )


If you have any comments or questions, please contact the author, Chad Husby ( chad.husby@fiu.edu or husby.1@osu.edu )

© Chad E. Husby 2002

Last updated October 19, 2002

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