Marattiaceae Bercht. & J. Presl
EOL Text
The Marattiaceae are represented by about 260 extant species and many fossil representatives. The extant members are placed in six genera: Angiopteris, Christensenia, Danaea, Eupodium, Marattia and Ptisana.
This early fern family is one of the oldest extant lineages of vascular plants that have changed little over periods of millions of years. They have a relatively good fossil record from the early Carboniferous (300 Ma) to the present day.
They can be easily distinguished from other ferns by their complex polycyclic stelar structure, often large, starchy rhizomes with fleshy or papery stipule-like outgrowths on each side of the petioles, presence of swollen nodes on rachises and often also on petioles, and eusporangia that are often fused and ligneous. All genera, except Christensenia, have free venation.
Marattiaceae are found in tropical or subtropical forests where the temperature and air humidity remain high throughout the year. While Angiopteris and Christensenia prefer lowland rain forest and grow in partly open habitats, Marattia has a preference for high elevation cloud forest and semi-deciduous forest in the subtropics. Danaea grows mostly in closed canopy, lowland and mountain rain forest and other shady, humid places, along stream banks, in ravines, and in sinkholes or near waterfalls, quite often on rather steep slopes.
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| Rights holder/Author | Maarten Christenhusz, Tree of Life web project |
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Large terrestrial, perennial ferns. Rhizome erect, large, roots fleshy covered in fleshy persistent outgrows. Stipe with persistent fleshy stipules and a basal swelling (pulvinus). Fronds large, numerous, tufted, 2-pinnate, basiscopically developed; secondary rhachis with a basal pulvinus; veins simple or forked, free. Indumentum composed of simple uniseriate or branched hairs, and short-stalked, slender scales occurring along the axes. Sporangia large, clustered in double rows near vein endings on underside of frond, free or fused into elongate bivalved group of synangia; without an indusium, homosporous.
The Marattiaceae are a distinct group of (almost always) terrestrial ferns with large, fleshy, erect or creeping rhizomes, with a polycyclic dictyostele (Sporne 1962, Smith et al. 2006), and mucilage canals in the roots, rhizomes and leaves. Young plants usually contain a mycorrhizal fungus - the oomycete Stigeosporium marattiacearum C. West - within the cortex (Sporne 1962). Other notable features are the starchy, leathery or papery stipule-like paired auricles at the base of each petiole, which are unique among ferns. Many species produce proliferous buds on the stipules or on the leaves (Uffelen 1994). The leaves are generally large and leathery (except in some “filmy” Danaea), and 1-3-pinnate (simple in some Danaea, or palmate in Christensenia). The petioles and rachises have polycyclic xylem and swollen pulvinae at the base of each internode. The sporangia are usually fused in round or elongate synangia, where a thin part of the sporangium dries and shrinks to form a pore through which the spores fall, except in Angiopteris where the sporangia are almost free. Each sporangium encloses very large numbers of spores (from about 1000 in Angiopteris to 7000 in Christensenia); these are usually monolete, bilateral or ellipsoid, and echinate (Sporne 1962, Smith et al. 2006).
The spores germinate rapidly, within a few days of being shed, and develop into large, monoecious, dark-green, mycorrhizal prothalli that can live for several years. An old prothallus may be several centimetres long and may closely resemble a large thalloid liverwort (Sporne 1962).
The young parts of Angiopteris, Ptisana, Eupodium and Marattia are covered with short simple hairs, while those of Christensenia and Danaea bear peltate scales. Microscopic characters, such as hairs, scales and indusia, which are of great taxonomic importance in other fern groups, are absent or vary little within marattioid genera.
Vegetative reproduction appears to be an important way of dispersal. Reproduction by means of stipule cuttings is also the easiest way of propagation in cultivation for Marattia and Angiopteris (Uffelen 1994, Chiou et al. 2006), but Christensenia and Danaea are difficult to cultivate because their requirements are unknown (Schneider 1893). Some Danaea species reproduce vegetatively, by means of apical buds on the leaves, where the terminal pinna is replaced or aborted; the bud can grow into a new plant when the leaf arches and the tip reaches the substrate. In this way, large uniform populations are sometimes formed. (After Christenhusz 2007)
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| Rights holder/Author | Maarten Christenhusz, Tree of Life web project |
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The first attempt to investigate evolutionary trends in Marattiales was published by Stidd (1974), who compared stelar structures among various fossil and extant genera. Hill and Camus (1986) examined generic relationships using a cladistic analysis of morphological characters, and proposed a new classification for Marattiales. In that study, they hypothesized that Christensenia was sister to a clade comprising Danaea, Marattia, and Angiopteris. Later phylogenetic studies based on DNA sequence data, which included some representatives from the Marattiaceae, consistently showed Danaea as sister to a clade uniting Marattia and Angiopteris (Hasebe et al. 1995, Pryer et al. 2004).
More recently molecular studies focussing on Danaea (Christenhusz et al. 2008) and Ptisana (Murdock 2008) have given greater insights into the indivisual relationships of this family.
The tree presented here is a consensus of these previous studies, which is inevitable because no good outgroup is available; all genera are too distant from the most closely related extant group, the Equisetaceae. Moreover not all species could be included in the sampling of the previous studies, and they are therefore here included based on their closest morphological resemblance.
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| Rights holder/Author | Maarten Christenhusz, Tree of Life web project |
| Source | http://tolweb.org/Marattiaceae/134249 |
Barcode of Life Data Systems (BOLD) Stats
Specimen Records:128
Specimens with Sequences:125
Specimens with Barcodes:125
Species:71
Species With Barcodes:67
Public Records:122
Public Species:67
Public BINs:0
Class Marattiopsida is a group of ferns containing a single order, Marattiales, and family, Marattiaceae.[1] Class Marattiopsida diverged from other ferns very early in their evolutionary history and are quite different from many plants familiar to people in temperate zones. Many of them have massive, fleshy rootstocks and the largest known fronds of any fern. The Marattiaceae is one of two groups of ferns traditionally known as eusporangiate fern,[2] meaning that the sporangium is formed from a group of cells vs the leptosporangium in which there is a single initial cell. There have long been four traditional extant genera (Angiopteris, Christensenia, Danaea and Marattia), but recent genetic/cladistic analysis [3] has determined the genus Marattia to be paraphyletic, and the genus has been split into three genera, the two new ones being Eupodium and Ptisana. This fern group has a long fossil history with many extinct taxa (Psaronius, Asterotheca, Scolecopteris, Eoangiopteris, Qasimia, Marantoidea, Danaeites, Marattiopsis, etc.).
In this group, such fronds are found in the genus Angiopteris, native to Australasia, Madagascar and Oceania. These fronds may be up to 9 meters long in the species Angiopteris teysmanniana of Java. In Jamaica the species Angiopteris evecta is widely naturalized and is registered as an invasive species. The plant was introduced by Captain Bligh from Tahiti as a staple food for slaves and cultivated in the Castleton Botanical Garden in 1860. From there it was able to distribute itself throughout the eastern half of the island.
The East-Asian genus Christensenia is named in honor of the Danish pteridologist Carl Christensen is an uncommon fern with distinctive fronds resembling a horse chestnut leaf, hence the species Christensenia aesculifolia, meaning horse-chestnut-leaved Christensenia.
The most widespread genus in Marattiaceae is the pantropical Marattia, usually occurring at higher elevations. These are also large ferns with globular rhizomes, but fronds can be up to 4 times pinnate. The sporangia are fused into bivalvate structures called a synangium.
The fourth genus Danaea is endemic to the Neotropics. They have bipinnate leaves with opposite pinnae, which are dimorphic, the fertile leaves much contracted, and covered below with sunken synangia.
The genus Eupodium is also neotropical, with two species. It has fronds that are 2-5 times pinnate, stalked synangia, and awns on the adaxial side of distal blade segments. Blade division decreases towards the apex of the frond. Plants of Eupodium usually only have one frond per plant per year (sometimes two).
Ptisana is a paleotropical genus. These plants are 2-4 times pinnate. Terminal segments usually have a prominent suture where they attach. The sporangia lack the labiate apertures of Marattia and Eupodium, and synangia are deeply cut. The King Fern, Ptisana salicina, from New Zealand and the South Pacific and known in Māori as para now has been placed in this genus. Sometimes called the Potato Fern, this is a large fern with an edible fleshy rhizome that is used as a food source by some indigenous peoples.
Several other genera have been named in the Marattiaceae, namely: Archangiopteris, Macroglossum, Protangiopteris, and Protomarattia. These are all synonyms of Angiopteris.
According to recent molecular studies it appears that these eusporangiate ferns may be a sister group to the horsetails (Equisetaceae). Both groups are certainly of ancient lineage.
References[edit]
- ^ Smith, Alan R.; Kathleen M. Pryer; Eric Schuettpelz; Petra Korall; Harald Schneider; Paul G. Wolf (2006). "A classification for extant ferns" (PDF). Taxon 55 (3): 705–731. doi:10.2307/25065646. JSTOR 25065646.
- ^ C.Michael Hogan. 2010. Fern. Encyclopedia of Earth. National council for Science and the Environment. Washington, DC
- ^ Murdock, Andrew G. (2008). "A taxonomic revision of the eusporangiate fern family Marattiaceae, with description of a new genus Ptisana". Taxon 57 (3): 737–755.
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Marattiidae is a subclass of Equisetopsida in the sense used by Mark W. Chase and James L. Reveal in their 2009 article "A phylogenetic classification of the land plants to accompany APG III."[1] This subclass comprises a single fern order, Marattiales, and family, Marattiaceae. It is equivalent to the class Marattiopsida in previous treatments, including Smith et al., 2006.[2]
Phylogeny[edit]
The following diagram shows a likely phylogenic relationship between subclass Marattiidae and the other Equisetopsida subclasses.[3]
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References[edit]
- ^ Mark W. Chase & James L. Reveal (2009). "A phylogenetic classification of the land plants to accompany APG III". Botanical Journal of the Linnean Society 161: 122–127. doi:10.1111/j.1095-8339.2009.01002.x.
- ^ Alan R. Smith, Kathleen M. Pryer, Eric Schuettpelz, Petra Korall, Harald Schneider & Paul G. Wolf (2006). "A classification for extant ferns". Taxon 55 (3): 705–731. doi:10.2307/25065646.
- ^ Maarten J. M. Christenhusz, Xian-Chun Zhang & Harald Schneider (2011). "A linear sequence of extant families and genera of lycophytes and ferns". Phytotaxa 19: 7–54.
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