Isoëtes

''Isoetes'', commonly known as the quillworts, is the only extant genus of plants in the family Isoetaceae, which is in the class of lycopods. There are currently 192 recognized species, with a cosmopolitan distribution but with the individual species often scarce to rare. Some botanists split the genus, separating two South American species into the genus ''Stylites'', although molecular data place these species among other species of ''Isoetes'', so that ''Stylites'' does not warrant taxonomic recognition. Species of ''Isoetes'' virtually identical to modern forms have existed since the Jurassic epoch.

The name of the genus may also be spelled ''Isoëtes''. The diaeresis (two dots over the e) indicates that the o and the e are to be pronounced in two distinct syllables. Including this in print is optional; either spelling (''Isoetes'' or ''Isoëtes'') is correct.

Description

Quillworts are mostly aquatic or semi-aquatic in clear ponds and slow-moving streams, though several (e.g. '' I. butleri'', '' I. histrix'' and '' I. nuttallii'') grow on wet ground that dries out in the summer. The quillworts are spore-producing plants and highly reliant on water dispersion. Quillworts have different ways to spread their spores based on the environment. Quillwort leaves are hollow and quill-like, with a minute ligule at the base of the upper surface. arising from a central corm. Each leaf is narrow, long (exceptionally up to ) and wide; they can be either evergreen, winter deciduous, or dry-season deciduous. Only 4% of total biomass, the tips of the leaves, is chlorophyllous.Tropical Alpine Environments: Plant Form and Function
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The roots broaden to a swollen base up to wide where they attach in clusters to a bulb-like, underground rhizome characteristic of most quillwort species, though a few (e.g. '' I. tegetiformans'') form spreading mats. This swollen base also contains male and female sporangia, protected by a thin, transparent covering (''velum''), which is used diagnostically to help identify quillwort species. They are heterosporous. Quillwort species are very difficult to distinguish by general appearance. The best way to identify them is by examining their megaspores under a microscope. Moreover, habitat, texture, spore size, and velum provide features that distinguish Isoëtes taxa. They also possess a vestigial form of secondary growth in the basal portions of its cormlike stem, an indication that they evolved from larger ancestors.

Biochemistry and genetics

Quillworts use Crassulacean acid metabolism (CAM) for carbon fixation. Some aquatic species don't have stomata and the leaves have a thick cuticle which prevents CO₂ uptake, a task that is performed by their hollow roots instead, which absorb CO₂ from the sediment. This has been studied extensively in ''Isoetes andicola''. CAM is normally considered an adaptation to life in arid environments to prevent water loss with the plants opening their stomata at night rather than in the heat of the day. This allows CO₂ to enter and minimises water loss. As mostly submerged aquatic plants, quillworts do not lack water and the use of CAM is considered to avoid competition with other aquatic plants for CO₂ during daytime.

The first detailed quillwort genome sequence, of '' I. taiwanensis'', showed that there were differences from CAM in terrestrial plants. CAM involves the enzyme phosphoenolpyruvate carboxylase (PEPC) and plants have two forms of the enzyme. One is normally involved in photosynthesis and the other in central metabolism. From the genome sequence, it appears that in quillworts, both forms are involved in photosynthesis. In addition, the time of day of the peak abundance of some of the components of CAM was different from terrestrial plants. These fundamental differences in biochemistry suggests that CAM in quillworts is probably another example of convergent evolution of CAM during the more than 300 million years since the genus diverged from other plants. However, they may also be because of differences between life in water and in the air. The genome sequence also provided two insights into its structure. First, genes and repeated non-coding regions were fairly evenly distributed across all the chromosomes. This is similar to genomes of other non-seed plants, but different from the seed plants (angiosperms) where there are distinctly more genes at the ends of chromosomes. Secondly, there was also evidence that the whole genome had been duplicated in the ancient past.

Taxonomy

Compared to other genera, ''Isoetes'' is poorly known. The first critical monograph on their taxonomy was published in 1922 and remained a standard reference for decades. Even after studies with cytology, scanning electron microscopy, and chromatography, species are difficult to identify and their phylogeny is disputed. Vegetative characteristics commonly used to distinguish other genera, such as leaf length, rigidity, color, or shape are variable and depend on the habitat. Most classification systems for ''Isoetes'' rely on spore characteristics, which make species identification nearly impossible without microscopy.

Reproduction

Overview

Like all land plants, ''Isoetes'' undergoes an alternation of generations between a diploid sporophyte stage and a sexual haploid gametophyte stage. However, the dominance of one stage over the other has shifted over time. The development of vascular tissue and subsequent diversification of land plants coincides with the increased dominance of the sporophyte and reduction of the gametophyte. ''Isoetes'', as members of the Lycopodiopsida class, are part of the oldest extant lineage that reflects this shift to a sporophyte dominant lifecycle. In closely related lineages, such as the extinct ''Lepidodendron'', spores were dispersed by the sporophyte through large collections of sporangia called strobili for wind-based spore dispersal. However, ''Isoetes'' are small heterosporous semi-aquatic plants, with different reproductive needs and challenges than large tree-like land plants.

Description

Like the rest of the Lycopodiopsida class, ''Isoetes'' reproduces with spores. Among the lycophytes, both ''Isoetes'' and the Selaginellaceae (spikemosses) are heterosporous, while the remaining lycophyte family Lycopodiaceae (clubmosses) is homosporous. As heterosporous plants, fertile Isoetes sporophytes produce megaspores and microspores, which develop in the megasporangia and microsporangia. These spores are highly ornate and are the primary way by which species are identified, although no one functional purpose of the intricate surface patterns is agreed upon. The megasporangia occur within the outermost microphylls (single-veined leaves) of the plant while the microsporangia are found in the innermost microphylls. This pattern of development is hypothesized to improve the dispersal of the heavier megaspore. These spores then germinate and divide into mega- and micro- gametophytes. The microgametophytes have antheridia, which in turn produce sperm. The megagametophytes have archegonia, which produce egg cells. Fertilization takes place when the motile sperm from a microgametophyte locates the archegonia of a megagametophyte and swims inside to fertilize the egg.

Outside of heterospory, a distinguishing feature of ''Isoetes'' (and ''Selaginella'') from other pteridophytes, is that their gametophytes grow inside the spores. This means that the gametophytes never leave the protection of the spore that disperses them, cracking the perispore (the outer layer of the spore) just enough to allow the passage of gametes. This is fundamentally different from ferns, where the gametophyte is a photosynthetic plant exposed to the elements of its environment. However, containment creates a separate problem for Isoetes, which is that the gametophytes have no way to acquire energy on their own. ''Isoetes'' sporophytes solve this problem by provisioning starches and other nutrients to the spores as an energy reserve for the eventual gametophytes. Although not a homologous process, this provisioning is somewhat analogous to other modes of offspring resource investment in seed-plants, such as fruits and seeds. The extent to which resources provisioned to the megaspore also support the growth of the new sporophyte is unknown in ''Isoetes''.

Dispersal

Spore dispersal occurs primarily in water ( hydrochory) but may also occur via adherence to animals (zoochory) and as a result of ingestion (endozoochory). These are among the reasons suggested for the ornamentations of the spore, with some authors demonstrating that certain patterns seem well-adapted for sticking to relevant animals like waterfowl. Another critical element of dispersal is the observation that in some species of ''Isoetes'', the outer coat of megaspores have pockets that trap microspores, a condition known as synaptospory. Typically, heterospory means that colonization and long-dispersal are more difficult due to the fact that a single spore cannot grow a bisexual gametophyte and thus cannot establish a new population from a single spore as can happen in homosporous ferns. ''Isoetes'' may mitigate this issue via microspores stuck to megaspores, greatly increasing the possibility of successful fertilization upon dispersal.

Species

, ''Plants of the World Online'' accepted the following extant species:

*'' Isoetes abyssinica'' Chiov.
*'' Isoetes acadiensis'' Kott
*'' Isoetes aemulans'' J.P.Roux
*'' Isoetes aequinoctialis'' Welw. ex A.Br.
*'' Isoetes alcalophila'' S.Halloy
*'' Isoetes alpina'' Kirk – New Zealand quillwort
*''Isoetes alstonii'' C.F.Reed & Verdc.
* ''Isoetes'' × ''altonharvillii'' Musselman
*'' Isoetes amazonica'' A.Br.
*'' Isoetes anatolica'' Prada & Rolleri
*'' Isoetes andicola'' (Amstutz) L.D.Gómez
*'' Isoetes andina'' Spruce ex Hook.
*'' Isoetes appalachiana'' D.F.Brunt. & D.M.Britton – Appalachian quillwort
*'' Isoetes araucaniana'' Macluf & Hickey
*'' Isoetes asiatica'' (Makino) Makino
*'' Isoetes attenuata'' C.R.Marsden & Chinnock
*'' Isoetes australis'' S.Williams – Australian quillwort
*'' Isoetes azorica'' Durieu
*'' Isoetes baculata'' Hickey & H.P.Fuchs
*''Isoetes biafrana'' Alston
*'' Isoetes bischlerae'' H.P.Fuchs
*'' Isoetes bolanderi'' Engelm. – Bolander's quillwort
*'' Isoetes boliviensis'' U.Weber
*'' Isoetes boomii'' Luebke – Boom's quillwort; southeast US
*'' Isoetes boryana'' Durieu
*'' Isoetes boyacensis'' H.P.Fuchs
*'' Isoetes bradei'' Herter
*'' Isoetes brasiliensis'' H.P.Fuchs
*'' Isoetes brevicula'' E.R.L.Johnson
* ''Isoetes'' × ''brittonii'' D.F.Brunt. & W.C.Taylor
* ''Isoetes'' × ''bruntonii'' Knepper & Musselman
*'' Isoetes butleri'' Engelm. – Butler's quillwort
*'' Isoetes cangae'' J.B.S.Pereira, Salino & Stützel
*'' Isoetes capensis''
* ''Isoetes'' × ''carltaylorii'' Musselman
*'' Isoetes caroli'' E.R.L.Johnson – Brazil
*'' Isoetes caroliniana'' (A.A.Eaton) Luebke is regarded by ''Plants of the World Online'' as a synonym of '' Isoetes valida'', but other sources treat it as a valid species
*'' Isoetes chubutiana'' Hickey, Macluf & W.C.Taylor
*''Isoetes coreana'' Y.H.Chung & H.K.Choi is a synonym of '' Isoetes sinensis'' var. ''coreana''
*'' Isoetes coromandelina'' L.f.
*'' Isoetes creussensis'' Lazare & S.Riba
*'' Isoetes cristata'' C.R.Marsden & Chinnock
*'' Isoetes cubana'' Engelm.
*'' Isoetes delilei'' (Bory) Rothm.
*'' Isoetes dispora'' Hickey
*'' Isoetes dixitii'' Shende
* ''Isoetes'' × ''dodgei'' A.A.Eaton
*'' Isoetes drummondii'' A.Braun – Drummond's quillwort
*'' Isoetes durieui'' Bory – Durieu's quillwort
* ''Isoetes'' × ''eatonii'' R.Dodge – Eaton's quillwort
*''Isoetes echinospora'' Durieu
* ''Isoetes'' × ''echtuckerii'' D.F.Brunt. & D.M.Britton
*'' Isoetes ecuadoriensis'' Aspl.
*'' Isoetes ekmanii'' U.Weber
*'' Isoetes elatior'' A.Braun
*'' Isoetes eludens'' J.P.Roux, Hopper & Rhian J.Sm. – elusive quillwort
*'' Isoetes engelmannii'' A.Braun – Engelmann's quillwort
*'' Isoetes escondidensis'' S.Halloy
*'' Isoetes eshbaughii'' Hickey & H.P.Fuchs
* ''Isoetes'' × ''fairbrothersii'' J.D.Montgom. & W.C.Taylor
*'' Isoetes flaccida'' Shuttlew. – southern quillwort
*'' Isoetes fluitans'' M.I.Romero
* ''Isoetes'' × ''foveolata'' A.A.Eaton
*'' Isoetes fuliginosa'' R.L.Small & Hickey
*'' Isoetes fuscomarginata'' H.P.Fuchs
*'' Isoetes gardneriana'' Kunze
*'' Isoetes georgiana'' Luebke – Georgia quillwort; southeast US
*'' Isoetes giessii'' Launert
*'' Isoetes gigantea'' U.Weber
* ''Isoetes'' × ''gopalkrishnae'' S.K.Singh, P.K.Shukla & N.K.Dubey
*'' Isoetes graniticola'' D.F.Brunt.
*'' Isoetes gunnii'' A.Braun
*''Isoetes gymnocarpa'' (Gennari) A.Braun
*'' Isoetes habbemensis'' Alston
*'' Isoetes hallasanensis'' H.K.Choi, Ch.Kim & J.Jung – Korea
* ''Isoetes'' × ''harveyi'' A.A.Eaton (syn. ''Isoetes'' × ''heterospora'' Eaton)
*'' Isoetes haussknechtii'' Troìa & Greuter
*'' Isoetes hawaiiensis'' W.C.Taylor & W.H.Wagner
*'' Isoetes heldreichii'' Wettst.
*'' Isoetes hemivelata'' R.L.Small & Hickey
* ''Isoetes'' × ''herb-wagneri'' W.C.Taylor
*'' Isoetes herzogii'' U.Weber
*'' Isoetes hewitsonii'' Hickey
* ''Isoetes'' × ''hickeyi'' W.C.Taylor & Luebke
*'' Isoetes hieronymi'' U.Weber
*'' Isoetes histrix'' Bory – land quillwort; Channel Islands
*'' Isoetes hopei'' J.R.Croft
*'' Isoetes howellii'' Engelm. – Howell's quillwort
*'' Isoetes humilior'' A.Braun
*''Isoetes hyemalis'' D.F.Brunt. is a synonym of ''Isoetes riparia'' var. ''amesii''
*'' Isoetes hypsophila'' Hand.-Mazz.
*'' Isoetes inflata'' E.R.L.Johnson
*'' Isoetes jaegeri'' Pitot
*'' Isoetes jamaicensis'' Hickey
*'' Isoetes japonica'' A.Braun – Japan
* ''Isoetes'' × ''jeffreyi'' D.M.Britton & D.F.Brunt.
*'' Isoetes jejuensis'' H.K.Choi, Ch.Kim & J.Jung
*'' Isoetes junciformis'' D.F.Brunt. & D.M.Britton – rush quillwort; southeast US
*'' Isoetes karstenii'' A.Braun
*'' Isoetes killipii'' C.V.Morton
*'' Isoetes kirkii'' A.Braun
*''Isoetes labri-draconis'' N.R.Crouch
*'' Isoetes lacustris'' L. – lake quillwort
*'' Isoetes laosiensis'' C.Kim & H.K.Choi
*'' Isoetes lechleri'' Mett.
*''Isoetes libanotica'' Musselman, Bolin & R.D.Bray
*'' Isoetes lithophila'' N.Pfeiff.
*'' Isoetes longissima'' Bory
*'' Isoetes louisianensis'' Thieret – Louisiana quillwort
*'' Isoetes luetzelburgii'' U.Weber
*''Isoetes macrospora''
*'' Isoetes malinverniana'' Ces. & De Not.
* ''Isoetes'' × ''marensis'' D.M.Britton & D.F.Brunt.
*'' Isoetes maritima'' Underw. – maritime quillwort
*'' Isoetes martii'' A.Braun
*'' Isoetes mattaponica'' Musselman & W.C.Taylor
*'' Isoetes maxima'' Hickey, Macluf & Link-Pérez
*'' Isoetes melanopoda'' J.Gay & Durieu – black-footed quillwort
*'' Isoetes melanospora'' Engelm. – black-spored quillwort
*'' Isoetes melanotheca'' Alston
*'' Isoetes mexicana'' Underw. (syn. ''Isoetes montezumae'' A.A.Eaton)
* ''Isoetes'' × ''michinokuana'' M.Takamiya, Mits.Watan. & K.Ono
*'' Isoetes microvela'' D.F.Brunt.
*'' Isoetes minima'' A.A.Eaton
*'' Isoetes mississippiensis'' S.W.Leonard, W.C.Taylor, Musselman & R.D.Bray
*'' Isoetes mongerensis'' E.R.L.Johnson – Monger quillwort
*'' Isoetes montana'' U.Weber
*'' Isoetes mourabaptistae'' J.B.S.Pereira, P.G.Windisch, Lorscheitt. & Labiak
*'' Isoetes muelleri'' A.Braun – Mueller's quillwort
*'' Isoetes naipiana'' P.G.Windisch, Lorscheitt. & Nervo
*'' Isoetes nana'' J.B.S.Pereira
*''Isoetes neoguineensis''
*'' Isoetes nigritiana'' A.Br.
*'' Isoetes nigroreticulata'' Verdc.
* ''Isoetes'' × ''novae-angliae'' D.F.Brunt. & D.M.Britton
*'' Isoetes novogranadensis'' H.P.Fuchs
*'' Isoetes nuttallii'' A.Braun – Nuttall's quillwort
*'' Isoetes occidentalis'' L.F.Hend.
*'' Isoetes olympica'' A.Br.
*'' Isoetes orcuttii'' A.A.Eaton - Orcutt's quillwort
*'' Isoetes organensis'' U.Weber
*'' Isoetes orientalis'' Hong Liu & Q.F.Wang
*'' Isoetes ovata'' N.Pfeiff.
*'' Isoetes pallida'' Hickey
*'' Isoetes palmeri'' H.P.Fuchs
*'' Isoetes panamensis'' Maxon & C.V.Morton
* ''Isoetes'' × ''paratunica'' D.F.Brunt., Mochalova & A.A.Bobrov
*'' Isoetes parvula'' Hickey
*'' Isoetes pedersenii'' H.P.Fuchs ex E.I.Meza & Macluf
*'' Isoetes perralderiana'' Durieu & Letourn. ex Milde
*'' Isoetes perrieriana'' Iversen
*'' Isoetes philippinensis'' Merr. & L.M.Perry
*'' Isoetes phrygia'' Hausskn.
*'' Isoetes piedmontana'' (N.Pfeiff.) C.F.Reed – Piedmont quillwort
*'' Isoetes pitotii'' Alston
*'' Isoetes precocia'' R.L.Small & Hickey
*'' Isoetes pringlei'' Underw.
*'' Isoetes prototypus'' D.M.Britton & Goltz
*'' Isoetes pseudojaponica'' M.Takamiya, Mits.Watan. & K.Ono
* ''Isoetes'' × ''pseudotruncata'' D.M.Britton & D.F.Brunt.
*''Isoetes pusilla'' C.R.Marsden & Chinnock
*'' Isoetes quiririensis'' J.B.S.Pereira & Labiak
*''Isoetes ramboi'' Herter
*''Isoetes riparia'' Engelm. ex A.Braun
*'' Isoetes sabatina'' Troìa & Azzella
*'' Isoetes saccharata'' Engelm.
*'' Isoetes sahyadrii'' Mahab.
*'' Isoetes saracochensis'' Hickey
*'' Isoetes savatieri'' Franch.
*''Isoetes schweinfurthii'' A.Br.
*'' Isoetes sehnemii'' H.P.Fuchs
*'' Isoetes septentrionalis'' D.F.Brunt.
*'' Isoetes serracarajensis'' J.B.S.Pereira, Salino & Stützel
*'' Isoetes setacea'' Lam.
*'' Isoetes sinensis'' T.C.Palmer – China
*'' Isoetes smithii'' H.P.Fuchs
*'' Isoetes spannagelii'' H.P.Fuchs
*'' Isoetes spinulospora'' C.Jermy & Schelpe
*'' Isoetes stellenbossiensis'' A.V.Duthie
*'' Isoetes stephanseniae'' A.V.Duthie
*'' Isoetes stevensii'' J.R.Croft
*'' Isoetes storkii'' T.C.Palmer
*''Isoetes taiwanensis'' De Vol – Taiwan
*'' Isoetes tamaulipana'' Mora-Olivo, A.Mend. & Mart.-Aval. – Tamaulipas, México
*''Isoetes tegetiformans'' Rury – mat-forming quillwort
*'' Isoetes tenella'' Léman ex Desv. – spiny-spore quillwort
*'' Isoetes tennesseensis'' Luebke & Budke – Tennessee quillwort; southeast US
*'' Isoetes tenuifolia'' Jermy
*'' Isoetes tenuissima'' Boreau – French quillwort
*'' Isoetes texana'' Singhurst, Rushing & W.C.Holmes
*'' Isoetes todaroana'' Troìa & Raimondo
*'' Isoetes toximontana'' Musselman & J.P.Roux - green-spored quillwort; South Africa
*'' Isoetes transvaalensis'' C.Jermy & Schelpe
*'' Isoetes triangula'' U.Weber
*'' Isoetes tripus'' A.Braun
*'' Isoetes truncata'' Clute
*'' Isoetes tuckermanii'' A.Braun ex Engelm.
*'' Isoetes tuerckheimii'' Brause
*'' Isoetes udupiensis'' P.K.Shukla, G.K.Srivast., S.K.Shukla & P.K.Rajagopal
*'' Isoetes ulei'' U.Weber
*'' Isoetes valida'' Clute – Carolina or strong quillwort
*'' Isoetes vanensis'' M.Keskin & G.Zare
*'' Isoetes vermiculata'' Hickey
*''Isoetes virginica'' N.Pfeiff. is a synonym of '' Isoetes melanopoda'' subsp. ''melanopoda''
*'' Isoetes viridimontana'' M.A.Rosenthal & W.C.Taylor – Green Mountain quillwort
*'' Isoetes weberi'' Herter
*'' Isoetes welwitschii'' A.Br. ex Kuhn
*'' Isoetes wormaldii'' Sim
*'' Isoetes yunguiensis'' Q.F.Wang & W.C.Taylor

Many species, such as the Louisiana quillwort and the mat-forming quillwort, are endangered species. Several species of ''Isoetes'' are commonly called Merlin's grass, especially '' I. lacustris'', but also the endangered species '' I. tegetiformans''.

Evolution

Fossilised specimens of '' I. beestonii'' have been found in rocks dating to the latest Permian. Quillworts are considered to be the closest extant relatives of the fossil tree ''Lepidodendron'', 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. Studies indicates that the Isoetes crown group which exist today evolved from a single lineage in the early Cenozoic.Phylogenomics indicates the “living fossil” Isoetes diversified in the Cenozoic
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References

External links

* ttps://gobotany.nativeplanttrust.org/species/isoetes/echinospora/. Distribution and classification list for world isoetes

{{Authority control

Lycophyte genera