Aspleniaceae

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Aspleniaceae
Illustration Asplenium trichomanes0.jpg
Scientific classification OOjs UI icon edit-ltr.svg
Kingdom: Plantae
Clade: Tracheophytes
Division: Polypodiophyta
Class: Polypodiopsida
Order: Polypodiales
Suborder: Aspleniineae
Family: Aspleniaceae
Newm.
Type genus
Asplenium
L.
Genera

The Aspleniaceae (spleenworts) are a family of ferns, included in the order Polypodiales. [1] The composition and classification of the family have been subject to considerable changes. In particular, there is a narrow circumscription, Aspleniaceae s.s. (adopted here), in which the family contains only two genera, and a very broad one, Aspleniaceae s.l., in which the family includes 10 other families kept separate in the narrow circumscription, with the Aspleniaceae s.s. being reduced to the subfamily Asplenioideae. The family has a worldwide distribution, with many species in both temperate and tropical areas. Elongated unpaired sori are an important characteristic of most members of the family.

Contents

Description

Sori of Asplenium trichomanes, showing linear arrangement with a thin membranous indusium along one edge Asplenium trichomanes sori (40523421534).jpg
Sori of Asplenium trichomanes , showing linear arrangement with a thin membranous indusium along one edge
Asplenium nidus in habitat: an epiphyte with undivided leaves Birds nest ferns in tropical montane forest on Mt Manucoco, Atauro, 30 Dec 2003.jpg
Asplenium nidus in habitat: an epiphyte with undivided leaves

Members of the family grow from rhizomes, that are either creeping or somewhat erect, and are usually but not always unbranched, and have scales that usually have a lattice-like (clathrate) structure. In some species, for example Asplenium nidus , the rhizomes form a kind of basket which collects detritus. The leaves may be undivided or be divided, with up to four-fold pinnation. The sori are characteristic of the family. They are elongated, and normally located on one side of a vein. More rarely, they may be in pairs on a single vein, but then they never curve over the vein. A flap-like indusium arises along one edge of a sorus. The leaf stalks (petioles) have two vascular bundles, uniting to form an X-shape in cross-section towards the tip of the leaf. The stalks of the sporangia are one cell wide in the middle. [2]

Taxonomy

The family Aspleniaceae was first described by Edward Newman in 1840. [3] Newman included three genera: Athyrium , Asplenium and Scolopendrium. [4] Athyrium is now placed in a different family, Athyriaceae, not considered very strongly related to the Aspleniaceae, and Scolopendrium is regarded as synonym of Asplenium. [1]

The narrow circumscription of the family adopted by the Pteridophyte Phylogeny Group classification of 2016 (PPG I) recognizes only two genera, Asplenium and Hymenasplenium . Asplenium has previously been split into a dozen or so genera, including Diella, found only in Hawaii. The consensus of molecular phylogenetic studies is that all are nested within Asplenium. [5] [6] PPG I places Aspleniaceae in the suborder Aspleniineae of the order Polypodiales.

Earlier, Christenhusz and Chase had proposed a much broader circumscription of Aspleniaceae, in which it consisted of all the separate families that PPG I places in the suborder Aspleniineae (eight at the time), with the families reduced to subfamilies. Thus the Aspleniaceae of PPG I became the subfamily Asplenioideae. [7] As of July 2019, the broader circumscription of the Aspleniaceae is used by Plants of the World Online, which lists 24 genera. [8]

Phylogenic relationships

Aspleniaceae is placed in a clade known as eupolypods II, or more formally as suborder Aspleniineae. The following cladogram, based on Lehtonen (2011) [9] and Rothfels & al. (2012), [2] shows a likely phylogenic relationship between the Aspleniaceae and the other families in the clade.

Aspleniineae  (eupolypods II)

Genera

In the PPG I system, Aspleniaceae s.s. contains two genera: [1]

Distribution and habitat

The Aspleniaceae have a worldwide distribution, with the large genus Asplenium being native to almost all parts of the world except Antarctica and some high Arctic areas. [10] The family is unusual in having high diversity in both temperate and tropical areas, and more-or-less equal numbers of terrestrial and epiphytic species. Plants are terrestrial, growing in the ground, lithophytic, growing on rocks, or epiphytic, growing on other plants; less often they are aquatic, growing in moving water. [2]

Related Research Articles

<span class="mw-page-title-main">Thelypteridaceae</span> Family of ferns

Thelypteridaceae is a family of about 900 species of ferns in the order Polypodiales. In the Pteridophyte Phylogeny Group classification of 2016, it is placed in the suborder Aspleniineae. Alternatively, the family may be submerged in a very broadly defined family Aspleniaceae as the subfamily Thelypteridoideae.

<span class="mw-page-title-main">Dryopteridaceae</span> Family of ferns

The Dryopteridaceae are a family of leptosporangiate ferns in the order Polypodiales. They are known colloquially as the wood ferns. In the Pteridophyte Phylogeny Group classification of 2016 (PPG I), the family is placed in the suborder Polypodiineae. Alternatively, it may be treated as the subfamily Dryopteridoideae of a very broadly defined family Polypodiaceae sensu lato.

<span class="mw-page-title-main">Polypodiales</span> Order of ferns

The order Polypodiales encompasses the major lineages of polypod ferns, which comprise more than 80% of today's fern species. They are found in many parts of the world including tropical, semitropical and temperate areas.

<span class="mw-page-title-main">Blechnaceae</span> Family of ferns

Blechnaceae is a family of ferns in the order Polypodiales, with a cosmopolitan distribution. Its status as a family and the number of genera included have both varied considerably. In the Pteridophyte Phylogeny Group classification of 2016, the family has 24 genera, and excludes genera placed in the separate family Onocleaceae. The family is divided into three subfamilies, including Blechnoideae s.s. Alternatively, the entire family may be treated as the subfamily Blechnoideae s.l. of a very broadly defined family Aspleniaceae, and include genera others place in Onocleaceae.

<span class="mw-page-title-main">Pteridaceae</span> Family of ferns

Pteridaceae is a family of ferns in the order Polypodiales, including some 1150 known species in ca 45 genera, divided over five subfamilies. The family includes four groups of genera that are sometimes recognized as separate families: the adiantoid, cheilanthoid, pteridoid, and hemionitidoid ferns. Relationships among these groups remain unclear, and although some recent genetic analyses of the Pteridales suggest that neither the family Pteridaceae nor the major groups within it are all monophyletic, as yet these analyses are insufficiently comprehensive and robust to provide good support for a revision of the order at the family level.

<span class="mw-page-title-main">Athyriaceae</span> Family of ferns

The Athyriaceae are a family of terrestrial ferns in the order Polypodiales. In the Pteridophyte Phylogeny Group classification of 2016 (PPG I), the family is placed in the suborder Aspleniineae, and includes two genera. Alternatively, it may be treated as the subfamily Athyrioideae of a very broadly defined family Aspleniaceae. The family has with a cosmopolitan distribution.

<span class="mw-page-title-main">Onocleaceae</span> Family of ferns

Onocleaceae is a small family of terrestrial ferns in the order Polypodiales. It is placed in the suborder Aspleniineae in the Pteridophyte Phylogeny Group classification of 2016. Alternatively, the family, along with Blechnaceae, may be placed in a very broadly defined family Aspleniaceae as the subfamily Blechnoideae. The family may contain from one to four genera, consisting of five species largely in north temperate climes. The four genera, Matteuccia, Onoclea, Onocleopsis and Pentarhizidium, may be included under the single genus Onoclea.

<span class="mw-page-title-main">Woodsiaceae</span> Family of ferns

Woodsiaceae is a family of ferns. In the Pteridophyte Phylogeny Group classification of 2016 (PPG I), it is placed in the suborder Aspleniineae. The family can also be treated as the subfamily Woodsioideae of a very broadly defined family Aspleniaceae sensu lato. In PPG I, the family contained only one genus, Woodsia. In 2020, Physematium was split off from Woodsia on the basis of molecular phylogenetic evidence. As of June 2023, Plants of the World Online continued to treat Physematium as a synonym of Woodsia.

<span class="mw-page-title-main">Aspleniineae</span> Suborder of ferns

Aspleniineae is a suborder of ferns in the order Polypodiales. It is equivalent to the clade eupolypods II in earlier systems; it is also treated as a single very broadly defined family Aspleniaceae. The suborder generally corresponds with the order Blechnales as described by J. L. Reveal in 1993. Aspleniineae includes some important ferns, including Onoclea sensibilis, the sensitive fern, which grows as a virtual weed throughout much of its temperate North American range, and ferns of the genus Thelypteris, a genus that has shown remarkable speciation. It also includes one of the more common horticultural ferns, Matteuccia struthiopteris, the ostrich fern.

<i>Homalosorus</i> Genus of ferns

Homalosorus is a genus of fern with only one species, Homalosorus pycnocarpos. It may also be referred to by its older synonyms Athyrium pycnocarpon and Diplazium pycnocarpon. Commonly referred to as the narrow-leaved glade fern, narrow-leaved-spleenwort, or glade fern, it is endemic to eastern North America and typically grows in moist woodlands. Once classified in the family Athyriaceae due to its linear, often doubled sori, in the Pteridophyte Phylogeny Group classification of 2016, it is placed in the small family Diplaziopsidaceae, whose other three species are native to east Asia. Other sources place the genus in the subfamily Diplaziopsidoideae of a very broadly defined family Aspleniaceae, equivalent to the suborder Aspleniineae in PPG I.

<span class="mw-page-title-main">Cystopteridaceae</span> Family of ferns

Cystopteridaceae is a family of ferns in the order Polypodiales. In the Pteridophyte Phylogeny Group classification of 2016 (PPG I), the family is placed in the suborder Aspleniineae of the order Polypodiales, and includes three genera. Alternatively, it may be treated as the subfamily Cystopteridoideae of a very broadly defined family Aspleniaceae.

<span class="mw-page-title-main">Hypodematiaceae</span> Family of ferns

Hypodematiaceae is a family of ferns in the order Polypodiales. In the Pteridophyte Phylogeny Group classification of 2016 (PPG I), the family is placed in the suborder Polypodiineae. Alternatively, it may be treated as the subfamily Hypodematioideae of a very broadly defined family Polypodiaceae sensu lato. The family consists of two, or in some versions three, small genera.

<span class="mw-page-title-main">Tectariaceae</span> Family of ferns

Tectariaceae is a family of leptosporangiate ferns in the order Polypodiales. In the Pteridophyte Phylogeny Group classification of 2016 (PPG I), the family is placed in the suborder Polypodiineae. Alternatively, it may be treated as the subfamily Tectarioideae of a very broadly defined family Polypodiaceae sensu lato. The family comprises seven genera, of which Tectaria is by far the largest.

<span class="mw-page-title-main">Polypodiineae</span> Suborder of ferns

Polypodiineae is a suborder of ferns in the order Polypodiales. It is equivalent to the clade eupolypods I in earlier systems, and to the very broadly defined family Polypodiaceae in the classification of Christenhusz & Chase (2014). It probably diverged from the suborder Aspleniineae during the mid-Cretaceous. The divergence is supported by both molecular data and an often overlooked morphological characteristic which lies in the vasculature of the petiole. Most species that make up the suborder have three vascular bundles. The only exceptions are the grammitid ferns which have one, and the genus Hypodematium which has two. This differs from eupolypods II which mostly have two vascular bundles.

Hemidictyum is a genus of ferns with a single species, Hemidictyum marginatum, commonly known as the marginated half net fern. In the Pteridophyte Phylogeny Group classification of 2016, it is the only genus in the family Hemidictyaceae. Alternatively, the family, along with Aspleniaceae sensu stricto, may be placed in a much more broadly defined family Aspleniaceae as the subfamily Asplenioideae.

<span class="mw-page-title-main">Diplaziopsidaceae</span> Family of ferns

Diplaziopsidaceae is a family of ferns in the order Polypodiales. In the Pteridophyte Phylogeny Group classification of 2016 (PPG I), the family is placed in the suborder Aspleniineae, and includes two genera. Alternatively, it may be treated as the subfamily Diplaziopsidoideae of a very broadly defined family Aspleniaceae.

Rhachidosorus is a genus of ferns in the order Polypodiales. It is the only genus in the family Rhachidosoraceae in the Pteridophyte Phylogeny Group classification of 2016. Alternatively, the genus may be placed in the subfamily Rhachidosoroideae of a more broadly defined family Aspleniaceae, the family placement used in Plants of the World Online as of November 2019.

<i>Hymenasplenium</i> Genus of ferns

Hymenasplenium is one of three genera of ferns in the Aspleniaceae, in the eupolypods II clade of the order Polypodiales. The others are Hemidictyum and Asplenium. Hymenasplenium was segregated because it is a natural grouping with differing rhizome morphology – dorsiventral v. radial for the rest of Asplenium, differing chromosome count – x=39 v. x=36 for the rest of Asplenium, and a clear monophyletic grouping based on genetic analysis. It was confirmed as a sister group to Asplenium in a 2015 molecular study of the genera.

The Pteridophyte Phylogeny Group (PPG) is an informal international group of systematic botanists who collaborate to establish on the classification of pteridophytes that reflects knowledge about plant relationships discovered through phylogenetic studies. In 2016, the group published a classification for extant pteridophytes, termed "PPG I". The paper had 94 authors . The classification was presented as a consensus classification supported by the community of fern taxonomists, but it has been partially exclusive and is highly contested. Alternative classifications of ferns exist and are preferred by more general taxonomists.

References

  1. 1 2 3 PPG I (2016). "A community-derived classification for extant lycophytes and ferns". Journal of Systematics and Evolution. 54 (6): 563–603. doi: 10.1111/jse.12229 . S2CID   39980610.
  2. 1 2 3 Carl J. Rothfels; Anders Larsson; Li-Yaung Kuo; Petra Korall; Wen- Liang Chiou; Kathleen M. Pryer (2012). "Overcoming Deep Roots, Fast Rates, and Short Internodes to Resolve the Ancient Rapid Radiation of Eupolypod II Ferns". Systematic Biology. 61 (1): 490–509. doi: 10.1093/sysbio/sys001 . PMID   22223449.
  3. ↑ "Aspleniaceae Newman". The International Plant Names Index . Retrieved 2019-07-30.
  4. ↑ Newman, Edward (1840). "Aspleniaceae". A history of British ferns. London: J. Van Voorst. pp. 6–7. Retrieved 2019-07-30.
  5. ↑ Smith, Alan R.; Pryer, Kathleen M.; Schuettpelz, Eric; Korall, Petra; Schneider, Harald & Wolf, Paul G. (2006). "A classification for extant ferns" (PDF). Taxon. 55 (3): 705–731. doi:10.2307/25065646. JSTOR   25065646. Archived from the original (PDF) on 2008-02-26. Retrieved 2008-02-26.
  6. ↑ Schneider, Harald; Ranker, Tom A.; Russell, Stephen J.; Cranfill, Raymond; Geiger, Jennifer M. O.; Aguraiuja, Ruth; Wood, Ken R.; Grundmann, Michael; Kloberdanz, Keelie & Vogel, Johannes C. (2005). "Origin of the endemic fern genus Diellia coincides with the renewal of Hawaiian terrestrial life in the Miocene". Proceedings of the Royal Society B. 272 (1561): 455–460. doi:10.1098/rspb.2004.2965. PMC   1634989 . PMID   15734701.
  7. ↑ Christenhusz, Maarten J.M. & Chase, Mark W. (2014). "Trends and concepts in fern classification". Annals of Botany . 113 (9): 571–594. doi:10.1093/aob/mct299. PMC   3936591 . PMID   24532607.
  8. ↑ "Aspleniaceae Newman". Plants of the World Online. Royal Botanic Gardens, Kew. Retrieved 2019-07-31.
  9. ↑ Samuli Lehtonen (2011). "Towards Resolving the Complete Fern Tree of Life". PLOS ONE. 6 (10): e24851. Bibcode:2011PLoSO...624851L. doi: 10.1371/journal.pone.0024851 . PMC   3192703 . PMID   22022365.
  10. 1 2 "Asplenium L." Plants of the World Online. Royal Botanic Gardens, Kew. Retrieved 2019-07-31.
  11. ↑ "Hymenasplenium Hayata". Plants of the World Online. Royal Botanic Gardens, Kew. Retrieved 2019-07-31.

Bibliography

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