Isoetales
Encyclopedia
Isoetales, also written Isoëtales, is an order
Order (biology)
In scientific classification used in biology, the order is# a taxonomic rank used in the classification of organisms. Other well-known ranks are life, domain, kingdom, phylum, class, family, genus, and species, with order fitting in between class and family...

 of plants in the class Isoetopsida
Isoetopsida
The Isoetopsida is a class of the Lycopodiophyta. All living plants belong to the genus Selaginella in the Selaginellales or to Isoetes in the order Isoetales. In the past, members of this group have sometimes been placed in the class Isoetopsida, sometimes in the Selaginellopsida or Lycopodiopsida...

. There are about 140-150 living species, all classified in the genus Isoëtes (quillworts), with a cosmopolitan distribution
Cosmopolitan distribution
In biogeography, a taxon is said to have a cosmopolitan distribution if its range extends across all or most of the world in appropriate habitats. For instance, the killer whale has a cosmopolitan distribution, extending over most of the world's oceans. Other examples include humans, the lichen...

 but often scarce to rare. Living species are mostly aquatic or semi-aquatic in clear ponds and slow-moving streams. Each leaf
Leaf
A leaf is an organ of a vascular plant, as defined in botanical terms, and in particular in plant morphology. Foliage is a mass noun that refers to leaves as a feature of plants....

 is slender and broadens downward to a swollen base up to 5 mm wide where the leaves attach in clusters to a bulb-like, underground corm
Corm
A corm is a short, vertical, swollen underground plant stem that serves as a storage organ used by some plants to survive winter or other adverse conditions such as summer drought and heat ....

 characteristic of most quillwort species. This swollen base also contains male and female sporangia
Sporangium
A sporangium is an enclosure in which spores are formed. It can be composed of a single cell or can be multicellular. All plants, fungi, and many other lineages form sporangia at some point in their life cycle...

, protected by a thin, transparent covering (velum
Velum
Velum may refer to:* Superior medullary velum, part of the nervous system that stretches between parts of the brain* Veil , the veil-like membrane of immature mushrooms extending from the margin of the cap to the stem and torn by growth...

), which is used diagnostically to help identify quillwort species. Quillwort species are very difficult to distinguish by general appearance. The best way to identify them is by examining the megaspores under a microscope.

All quillworts and their extinct relatives are heterosporous
Spore
In biology, a spore is a reproductive structure that is adapted for dispersal and surviving for extended periods of time in unfavorable conditions. Spores form part of the life cycles of many bacteria, plants, algae, fungi and some protozoa. According to scientist Dr...

. Some fossil species are very well known, with many stages of development and the life cycle preserved. Two of the best known are the Carboniferous
Carboniferous
The Carboniferous is a geologic period and system that extends from the end of the Devonian Period, about 359.2 ± 2.5 Mya , to the beginning of the Permian Period, about 299.0 ± 0.8 Mya . The name is derived from the Latin word for coal, carbo. Carboniferous means "coal-bearing"...

 Chaloneria and Cretaceous
Cretaceous
The Cretaceous , derived from the Latin "creta" , usually abbreviated K for its German translation Kreide , is a geologic period and system from circa to million years ago. In the geologic timescale, the Cretaceous follows the Jurassic period and is followed by the Paleogene period of the...

 Nathorstiana.

Fossilised specimens of Isoetes beestonii have been found in rocks dating to the early Triassic
Triassic
The Triassic is a geologic period and system that extends from about 250 to 200 Mya . As the first period of the Mesozoic Era, the Triassic follows the Permian and is followed by the Jurassic. Both the start and end of the Triassic are marked by major extinction events...

. Quillworts are considered by some to be the last remnant of the fossil
Fossil
Fossils are the preserved remains or traces of animals , plants, and other organisms from the remote past...

 tree Lepidodendron
Lepidodendron
Lepidodendron is an extinct genus of primitive, vascular, arborescent plant related to the Lycopsids . It was part of the coal forest flora. They sometimes reached heights of over , and the trunks were often over in diameter, and thrived during the Carboniferous period...

with which they share some unusual features including the development of both wood and bark, a modified shoot system acting as roots, bipolar growth, and an upright stance.
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