Sphenophyte
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Equisetidae is one of the four subclasses of
Polypodiopsida The ferns (Polypodiopsida or Polypodiophyta) are a group of vascular plants (plants with xylem and phloem) that reproduce via spores and have neither seeds nor flowers. They differ from mosses by being vascular, i.e., having specialized tissue ...
(ferns), a group of
vascular plant Vascular plants (), also called tracheophytes (, ) or collectively tracheophyta (; ), are plants that have lignin, lignified tissues (the xylem) for conducting water and minerals throughout the plant. They also have a specialized non-lignified Ti ...
s with a fossil record going back to the
Devonian The Devonian ( ) is a period (geology), geologic period and system (stratigraphy), system of the Paleozoic era (geology), era during the Phanerozoic eon (geology), eon, spanning 60.3 million years from the end of the preceding Silurian per ...
. They are commonly known as horsetails. They typically grow in wet areas, with whorls of needle-like branches radiating at regular intervals from a single vertical stem. The Equisetidae were formerly regarded as a separate division of spore plants and called Equisetophyta, Arthrophyta, Calamophyta or Sphenophyta. When treated as a class, the names Equisetopsida s.s. and Sphenopsida have also been used. They are now recognized as rather close relatives of the
fern The ferns (Polypodiopsida or Polypodiophyta) are a group of vascular plants (plants with xylem and phloem) that reproduce via spores and have neither seeds nor flowers. They differ from mosses by being vascular, i.e., having specialized tissue ...
s (Polypodiopsida) of which they form a specialized lineage. However, the division between the horsetails and the other ferns is so ancient that many botanists, especially paleobotanists, still regard this group as fundamentally separate at the higher level.


Description

The horsetails comprise photosynthesising, "segmented", hollow stems, sometimes filled with pith. At the junction ("node", see diagram) between each segment is a whorl of
leaves A leaf (: leaves) is a principal appendage of the stem of a vascular plant, usually borne laterally above ground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", while the leaves, stem, ...
. In the only extant genus ''Equisetum'', these are small leaves (
microphyll In plant anatomy and evolution a microphyll (or lycophyll) is a type of plant leaf with one single, unbranched leaf vein. Plants with microphyll leaves occur early in the fossil record, and few such plants exist today. In the classical concept of ...
s) with a singular vascular trace, fused into a sheath at each stem node. However, the leaves of ''Equisetum'' probably arose by the reduction of
megaphyll In plant anatomy and evolution a microphyll (or lycophyll) is a type of plant leaf with one single, unbranched leaf vein. Plants with microphyll leaves occur early in the fossil record, and few such plants exist today. In the classical concept of ...
s, as evidenced by early fossil forms such as ''
Sphenophyllum ''Sphenophyllum'' is a genus in the order Sphenophyllales. It has been placed in the family Sphenophyllaceae. Species Species that have been described include: *†''Sphenophyllum angustifolium'' *†''Sphenophyllum biarmicum'' Zalessky (1937) ...
'', in which the leaves are broad with branching veins. The
vascular bundle A vascular bundle is a part of the transport system in vascular plants. The transport itself happens in the stem, which exists in two forms: xylem and phloem. Both these tissues are present in a vascular bundle, which in addition will incl ...
s 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 new branches of their neighbours. The vascular system itself resembles that of the vascular plants'
eustele In a vascular plant, the stele is the central part of the root or stem containing the tissues derived from the procambium. These include vascular tissue, in some cases ground tissue (pith) and a pericycle, which, if present, defines the outermost b ...
, which evolved independently and convergently. 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 Xylem is one of the two types of transport tissue (biology), tissue in vascular plants, the other being phloem; both of these are part of the vascular bundle. The basic function of the xylem is to transport water upward from the roots to parts o ...
. Similar spaces, the vallecular canals are formed in the cortex. Due to the softer nature of the phloem, these are very rarely seen in fossil instances. In the
Calamitaceae Calamitaceae is an extinct Family (biology), family of Equisetales, equisetalean plants related to the modern horsetails, known from the Carboniferous and Permian periods. Some members of this family like ''Arthropitys'' attained tree-like statur ...
, secondary xylem (but not secondary
phloem Phloem (, ) is the living tissue in vascular plants that transports the soluble organic compounds made during photosynthesis and known as ''photosynthates'', in particular the sugar sucrose, to the rest of the plant. This transport process is ...
) was secreted as the
cambium A cambium (: cambiums or cambia), in plants, is a tissue layer that provides partially undifferentiated cells for plant growth. It is found in the area between xylem and phloem. A cambium can also be defined as a cellular plant tissue from whic ...
grew outwards, producing a woody stem, and allowing the plants to grow as high as 10m. All extant species of ''
Equisetum ''Equisetum'' (; horsetail) is the only living genus in Equisetaceae, a family of vascular plants that reproduce by spores rather than seeds. ''Equisetum'' is a "living fossil", the only living genus of the entire subclass Equisetidae, which ...
'' are herbaceous, and have lost the ability to produce secondary growth. The underground parts of the plants consist of jointed
rhizome In botany and dendrology, a rhizome ( ) is a modified subterranean plant stem that sends out roots and Shoot (botany), shoots from its Node (botany), nodes. Rhizomes are also called creeping rootstalks or just rootstalks. Rhizomes develop from ...
s, 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 most seed plants, which grow from an apical meristem - i.e. new growth comes only from growing tips (and widening of stems). Horsetails bear cones (technically ''
strobili A strobilus (: strobili) is a structure present on many land plant species consisting of sporangia-bearing structures densely aggregated along a stem. Strobili are often called cones, but some botanists restrict the use of the term cone to the woo ...
'', sing.
strobilus A strobilus (: strobili) is a structure present on many land plant species consisting of sporangia-bearing structures densely aggregated along a stem. Strobili are often called cones, but some botanists restrict the use of the term cone to the woo ...
) at the tips of some stems. These cones comprise spirally arranged
sporangiophore {{Short pages monitor thumb , right , A photomicrograph of a sporangiophore