|Subfamilies and genera|
Ophioglossaceae, the adder's-tongue family, is a small family of ferns. In the Pteridophyte Phylogeny Group classification of 2016 (PPG I), it is the only family in the order Ophioglossales, which together with the Psilotales is placed in the subclass Ophioglossidae. The Ophioglossidae are one of the groups traditionally known as eusporangiate ferns. Members of the family differ from other ferns in a number of ways. Many have only a single fleshy leaf at a time. Their gametophytes are subterranean and rely on fungi for energy.
Members of Ophioglossaceae are usually terrestrial (excepting a few epiphytic species of Ophioglossum) and occur in both temperate and tropical areas. They differ from the other ferns in several respects:
- Many species only send up one frond or leaf-blade per year, producing only a single leaf at a time. The leaves are usually fleshy, and in temperate areas will often turn brownish or reddish during colder months.
- Instead of the leptosporangia typical of most ferns they produce eusporangia, which are larger, contain more spores, and have thicker walls.
- Their sporophylls (spore-bearing leaves) are divided into two distinct parts, the sporophore, which produces sporangia and has a greatly reduced and modified blade, and the rest of the leaf, the trophophore.
- Their gametophytes are subterranean and rely on fungi for their energy (in other words, they are myco-heterotrophic), unlike the terrestrial, photosynthetic gametophytes found in most ferns.
In addition to having mycoheterotrophic gametophytes, there are a few members of Botrychium that are unique among ferns in having the sporophytes also mycoheterotrophic, producing only small, ephemeral sporophylls that do not photosynthesize.
The plants have short-lived spores formed in sporangia lacking an annulus, and borne on a stalk that splits from the leaf blade; and fleshy roots. A few species send up fertile spikes only, without any conventional leaf-blade. The spores will not germinate if exposed to sunlight, and the gametophyte can live some two decades without forming a sporophyte.
The genus Ophioglossum has the highest chromosome counts of any known plant. The record holder is Ophioglossum reticulatum, with about 630 pairs of chromosomes (1260 chromosomes per cell). Other measurements have indicated a chromosome number up to 1440 (n = 720). For comparison, humans have 46 chromosomes, consisting of n = 23 pairs.
History of classification
The ferns in this group were originally treated as a family and later as the separate order Ophioglossales. In some classifications, they were placed in a separate division, Ophioglossophyta, but molecular phylogenetic studies have shown the Ophioglossales to be closely related to the Psilotales, and both are placed in the class Ophioglossidae.
In the molecular phylogenetic classification of Smith et al. in 2006, Ophioglossales, in its present circumscription, was placed in the class Psilotopsida, along with the order Psilotales. The linear sequence of Christenhusz et al. (2011), intended for compatibility with the classification of Chase and Reveal (2009), which placed all land plants in Equisetopsida, made it a member of subclass Ophioglossidae, equivalent to Smith's Psilotopsida. This approach has subsequently been followed in the classifications of Christenhusz and Chase (2014) and PPG I (2016).
Older treatments recognized segregate families within the Ophioglossales, such as Botrychiaceae for the moonworts and grape ferns and Helminthostachyaceae for Helminthostachys, but modern treatments combine all members of the order into the single family Ophioglossaceae.
Subfamilies and genera
|Phylogeny of Ophioglossaceae|
|Phylogeny of Ophioglossaceae|
The number of genera into which the family is divided has varied. The Smith system of 2006 used four genera, treating Botrychium and Ophioglossum broadly. Cheiroglossa has been segregated from Ophioglossum, or included within it. The PPG I system of 2016 divides the family into four subfamilies:
- Mankyuoideae J.R.Grant & B.Dauphin
- Mankyua B.Y.Sun, M.H.Kim & C.H.Kim
- Ophioglossoideae C.Presl (equivalent to Ophioglossum s.l. in other systems)
- Helminthostachyoideae C.Presl
- Helminthostachys Kaulf.
- Botrychioideae C.Presl (equivalent to Botrychium s.l. in other systems)
- The Pteridophyte Phylogeny Group (November 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.
- Raven, Peter H.; Evert, Ray F. & Eichhorn, Susan E. (2005). Biology of Plants (7th ed.). New York: W.H. Freeman. ISBN 978-0-7167-1007-3.
- Khandalwal, Sharda (1990). "Chromosome evolution in the genus Ophioglossum L.". Botanical Journal of the Linnean Society. 102 (3): 205–217. doi:10.1111/j.1095-8339.1990.tb01876.x.
- Patel, Mitesh & Reddy, Mandadi (2018). "Discovery of the World's Smallest Terrestrial Pteridophyte". Scientific Reports. 8 (1): 5911. Bibcode:2018NatSR...8.5911P. doi:10.1038/s41598-018-24135-2. PMC 5897345. PMID 29651115.
- 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.
- Christenhusz, Maarten J. M.; Zhang, Xian-Chun & Schneider, Harald (2011). "A linear sequence of extant families and genera of lycophytes and ferns" (PDF). Phytotaxa. 19: 7–54. doi:10.11646/phytotaxa.19.1.2.
- Chase, Mark W.; Reveal, James L. (October 2009). "A phylogenetic classification of the land plants to accompany APG III". Botanical Journal of the Linnean Society. 161 (2): 122–127. doi:10.1111/j.1095-8339.2009.01002.x.
- Christenhusz, Maarten J. M. & Chase, Mark W. (2014). "Trends and concepts in fern classification". Annals of Botany. 113 (4): 571–594. doi:10.1093/aob/mct299. PMC 3936591. PMID 24532607.
- Zhang, Liang; Fan, Xue-Ping; Petchsri, Sahanat; Zhou, Lin; Pollawatn, Rossarin; Zhang, Xin; Zhou, Xin-Mao; Lu, Ngan Thi; Knapp, Ralf; Chantanaorrapint, Sahut; Limpanasittichai, Ponpipat; Sun, Hang; Gao, Xin-Fen; Zhang, Li-Bing (17 January 2020). "Evolutionary relationships of the ancient fern lineage the adder's tongues (Ophioglossaceae) with description of Sahashia gen. nov". Cladistics. 36 (4): 380–393. doi:10.1111/cla.12408. PMID 34618972. S2CID 212939923. Retrieved 25 March 2021.
- Nitta, Joel H.; Schuettpelz, Eric; Ramírez-Barahona, Santiago; Iwasaki, Wataru; et al. (2022). "An Open and Continuously Updated Fern Tree of Life". Frontiers in Plant Science. 13. doi:10.3389/fpls.2022.909768. PMID 36092417.
- "Tree viewer: interactive visualization of FTOL". FTOL v1.5.0 [GenBank release 256]. 2023. Retrieved 17 August 2023.
- C.Michael Hogan. 2010. Fern. Encyclopedia of Earth. National council for Science and the Environment. Washington, DC
- Ophioglossaceae in Flora of North America (incl. Botrychiaceae)