Equisetopsida

Equisetopsida, or Sphenopsida, is a class of vascular plants with a fossil record

Equisetopsida
going back to the Devonian. They are commonly known as horsetails.[2][3]
Living horsetails are represented by about twenty herbaceous species in the single Temporal range:
genus Equisetum.[4] They typically grow in wet areas, with whorls of needle-like Late Devonian [1] to Recent
branches radiating at regular intervals from a single vertical stem.
PreЄ Є OS D C P T J K Pg N

The Equisetopsida were formerly regarded as a separate division of spore plants

and also called Equisetophyta, Arthrophyta or Sphenophyta; today they have
been recognized as rather close relatives of the typical ferns (Pteridopsida) and
form a specialized lineage of the Pteridophyta.[5] However, the division between
the Equisetopsids and the ferns is so ancient that many botanists, especially
paleobotanists, still regard this group as fundamentally separate at the higher
level.

Contents
Morphology
Fossil record
Systematics
References
External links Equisetum telmateia

Scientific classification
Kingdom: Plantae
Morphology
Division: Pteridophyta
The Sphenophytes comprise photosynthesising, "segmented", hollow stems,
sometimes filled with pith. At the junction ("node", see diagram) between each Class: Equisetopsida
segment is a whorl of leaves. In the only extant genus Equisetum, these are small C. Agardh
leaves (microphylls) with a singular vascular trace, fused into a sheath at each Orders
stem node. However, the leaves of Equisetum probably arose by the reduction of
megaphylls, as evidenced by early fossil forms such as Sphenophyllum, in which Equisetales
the leaves are broad with branching veins.[6]
† Archaeocalamitaceae
The vascular bundles trifurcate at the nodes, with the central branch becoming the
vein of a microphyll, and the other two moving left and right to merge with the
† Calamitaceae
new branches of their neighbours.[7] The vascular system itself resembles that of Equisetaceae
the vascular plants' eustele, which evolved independently and convergently.[7]
† Pseudoborniales
Very rapid internode elongation results in the formation of a pith cavity and a ring
of carinal canals formed by disruption of the primary xylem. Similar spaces, the † Sphenophyllales
vallecular canals are formed in the cortex.[7] Due to the softer nature of the Synonyms
phloem, these are very rarely seen in fossil instances. In the Calamitaceae,
secondary xylem (but not secondary phloem) was secreted as the cambium grew Sphenopsida
outwards, producing a woody stem, and allowing the plants to grow as high as
[7]
10m. All extant species ofEquisetum are herbaceous, and have lost the ability to produce secondary growth.
The underground parts of the plants consist of jointed rhizomes, from which roots and aerial axes emerge. The plants have intercalary
meristems in each segment of the stem and rhizome that grow as the plant gets taller. This contrasts with the seed plants, which grow
from an apical meristem - i.e. new growth comes only from growing tips (and widening of stems).

Sphenophytes bear cones (technically strobili, sing. strobilus) at the tips of some stems. These cones comprise spirally arranged
sporangiophores, which bear sporangia at their edges, and in extant sphenophytes cover the spores externally - like sacs hanging from
an umbrella, with its handle embedded in the axis of the cone. In extinct groups, further protection was afforded to the spores by the
presence of whorls ofbracts - big pointed microphylls protruding from the cone.

The extant horsetails are homosporous, but extinct heterosporous species such as Calamostachys casheana appear in the fossil
record.[8] The sporangia open by lateral dehiscence to release the spores. The spores bear characteristic elaters, distinctive spring-like
attachments which are hygroscopic: i.e. they change their configuration in the presence of water, helping the spores move and aiding
their dispersal.

Fossil record
The extant horsetails represent a tiny fraction of Sphenophyte diversity in the past. There were three
orders of Equisetopsid; the Pseudoborniales, which first appeared in the late Devonian.[1] Second, the
Sphenophyllales which were a dominant member of the Carboniferous understory, and prospered until
the mid and early Permian respectively. The Equisetales existed alongside the Sphenophyllales, but
diversified as that group disappeared into extinction, gradually dwindling in diversity to today's single
genus Equisetum.

The organisms first appear in the fossil record during the late Devonian,[9] a time when land plants were
undergoing a rapid diversification, with roots, seeds and leaves having only just evolved. (See
Evolutionary history of plants) However, plants had already been on the land for almost a hundred
million years, with the first evidence of land plants dating to475 million years ago.[10]

Systematics
The horsetails and their fossil relatives have long been recognized as distinct from other seedless
vascular plants.[11] Before the advent of modern molecular studies, the relationship of this group to
other living and fossil plants was considered problematic.[12] Because of their unclear relationships, the Vegetative stem:
rank botanists have assigned to the horsetails varies from order to division. When recognized as a N = node,
separate division, the literature uses many possible names, including Arthrophyta,[12] I = internode,
Sphenophyta,[1][13] or Equisetophyta. Other authors have regarded the same group as a class, either B = branch in whorl,
L = fused
within a division consisting of the vascular plants or, more recently, within an expanded fern group.
[14] or Sphenopsida.[7]
microphylls
When ranked as a class, the group has been termed the Equisetopsida

Recent phylogenetic analysis has produced evidence that this group of plants belongs firmly within the
fern clade of vascular plants.[15] A 2006 classification by Smith et al. places the class Equisetopsida within an unranked clade of
Marattiopsida and Polypodiopsida.[5]
broadly defined ferns, as a sister to two classes more traditionally called ferns,

[16]
The probable relationships within Equisetopsida are shown in the cladogram below:

†Pseudobornia ursina

†Sphenophyllales

†Archeocalamitaceae

†Calamitaceae
 Equisetaceae

References
1. Taylor, T.N.; Taylor., E.L. (1993). The biology and evolution of fossil
plants. Englewood Cliffs, NJ: Prentice Hall. pp. 303–305. ISBN 0-13-
651589-4.
2. "horsetail" (http://www.merriam-webster.com/cgi-bin/dictionary?book=Di
ctionary&va=horsetail). Merriam-Webster Online Dictionary.
Cross-section through a strobilus;
3. Allaby, A.; Allaby, M. (1999). "Sphenopsida." A Dictionary of Earth sporangiophores, with attached
Sciences. sporangia (spore capsules) full of
4. Judd, W.S.; Campbell, C.S.; Kellogg, E.A.; Stevens, P.F.; Donoghue, spores, can be discerned.
M.J. (2002). Plant systematics, a phylogenetic approach. Sunderland,
Massachusetts: Sinauer Associates, Inc. p. 193.ISBN 0878934030.
5. Smith, A.R.; Pryer, K.M.; Schuettpelz, E.; Korall, P.; Schneider, H.; Wolf,
=P.G. (2006). "A classification for extant ferns"(https://web.archive.org/
web/20080226232147/http://www.pryerlab.net/publication/fichier749.pdf)
(PDF). Taxon. 55 (3): 705–731. doi:10.2307/25065646 (https://doi.org/1
0.2307%2F25065646). JSTOR 25065646 (https://www.jstor.org/stable/2
5065646). Archived from the original (http://www.pryerlab.net/publicatio
n/fichier749.pdf) (PDF) on 2008-02-26.
6. Rutishauser, R. (1999). "Polymerous leaf whorls in vascular plants:
Developmental morphology and fuzziness of organ identities".
International Journal of Plant Sciences. 160 (6): 81–103.
doi:10.1086/314221 (https://doi.org/10.1086%2F314221).
JSTOR 10.1086/314221 (https://www.jstor.org/stable/10.1086/314221).
PMID 10572024 (https://www.ncbi.nlm.nih.gov/pubmed/10572024).
7. Stewart, W.N.; Rothwell, G.W. (1993). Paleobotany and the evolution of
plants (2nd ed.). Cambridge:Cambridge University Press. ISBN 0-521-
38294-7.
8. Bower, F.O. (1959) [1935]. The origin of the land flora, a theory based Strobilus of E. telmateia,
upon the facts of alternation. New York: Hafner publishing Co. p. 381. terminal on an unbranched
9. Taylor, T.N.; Taylor, E.L. (1993). The Biology and Evolution of fossil stem
plants. Prentice Hall. ISBN 978-0136515890.
10. Wellman, C.H.; Osterloff, P.L.; Mohiuddin, U. (2003)."Fragments of the
earliest land plants" (http://sherpa.leeds.ac.uk/archive/00000106/01/well
manch1.pdf) (PDF). Nature. 425 (6955): 282–285.
doi:10.1038/nature01884 (https://doi.org/10.1038%2Fnature01884).
PMID 13679913 (https://www.ncbi.nlm.nih.gov/pubmed/13679913).
11. Eames, A.J. (1936). Morphology of vascular plants (Lower Groups)
.
New York and London: McGraw-Hill Book Company. pp. 110–115.
12. Bold, H.C.; Alexopoulos, C.J.; Delevoryas, T
. (1987). Morphology of
Plants and Fungi (5th ed.). New York: Harper-Collins. pp. 371–387, 478,
506–514. ISBN 0-06-040839-1.
13. Gifford, E.M.; Foster, A.S. (1988). Morphology and evolution of vascular
plants (3rd ed.). New York: W. H. Freeman and Company. pp. 175–207.
ISBN 0-7167-1946-0.
14. Kenrick, P.; Crane, P.R. (1997). The Origin and early diversification of
land plants: A cladistic study. Washington, D. C.: Smithsonian Institution
Press. pp. 241–242. ISBN 1-56098-730-8.
15. Pryer, K.M.; Schneider, H.; Smith, A.R.; Cran, R.; Wolf, P.G.; Hunt, J.S.;
Sipes, S.D. (2001). "Horsetails and ferns are a monophyletic group and
the closest living relatives to seed plants"(https://web.archive.org/web/2
0090902222737/http://www.pryerlab.net/publication/fichier199.pdf)
(PDF). Nature. 409 (6820): 618–621. doi:10.1038/35054555 (https://doi.
org/10.1038%2F35054555). PMID 11214320 (https://www.ncbi.nlm.nih.
gov/pubmed/11214320). Archived from the original (http://www.pryerlab.
net/publication/fichier199.pdf)(PDF) on 2009-09-02.
16. "Introduction to the Sphenophyta"(http://www.ucmp.berkeley.edu/plants/
sphenophyta/sphenophyta.html). University of California Museum of
Paleontology. Retrieved 31 July 2011.

External links
Data related to Equisetopsida at Wikispecies

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