HEDWIGIACEAE
Hedwigia
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HEDWIGIA P. Beauvois, Mag. Encycl. 5: 304. 1804.
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Hedwigia has narrowly lanceolate to ovate leaves, with hyaline acumina. The perichaetial leaves have ciliate margins, except H. detonsa with entire perichaetial leaves. Capsules are immersed. Leaf cells are papillose. The shape of the leaf cell papillae and the number of papillae per cell are the most important characters in distinguishing species in Hedwigia. The comparative SEM study by Biasuso (2007) provided evidence that leaf cell papillae in the abaxial surface are important character states for the distinction of species in Hedwigia. Hedwigia includes at least 12 species. Four species were recently described from Russia (Ignatova, et al. 2016, 2017). |
Hedwigia ciliata, from: Hedwig, J. 1787. Descriptio et adumbrato microscopio-analytica Muscorum Frondosorum nec non aliorum vegetantium e classe cryptogamica Linnaei novorum dubiisque vexatorum (link)
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1. Hedwigia ciliata (Hedwig) P. Beauvois, Prodr.Aethéogam., 15. 1805. Almost world wide. Widely distributed in Canada and the US. A northern European variant, Hedwigia ciliata var. leucophaea Bruch & Schimper was recognized by Hedenäs (1994). Reported as H. ciliata var ciliata from Argentina (Biasuso, 2007). Reported as H. ciliata var ciliata and as H. ciliata var leucophaea from Turkey (Erdag et al, 2003). Hedwigia ciliata is infrequent in Russia, occurring mainly in the North-Western European Russia, with few localities in its central part. It is characterized by longly recurved leaf margins, sinuose laminal cells and spores (27.5–)30–35(37.5) um. |
Hedwigia ciliata, figure from Fife 2014. Flora of New Zealand. Mosses.
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2. Hedwigia integrifolia P. Beauv., Podr. I`Aethogamie 60. 1805. = Hedwigidium integrifolium Dixon in C.E.O. Jensen, Skand. Bladmfl. 369. 1939. Eastern North America. As noticed by Allen (2010), Hooker and Taylor (1818) first synonymized Hedwigidium imberbe with Hedwigia integrifolia P. Beauv. (as Anictangium imberbe). The type of Hedwigia integrifolia was collected in North America and it was described as having ciliate perichaetial leaves. Palisot de Beauvois (1805) considered H. integrifolia very close to Hedwigia filiformis (Mich.) P. Beauv. Michaux (1803) considered Hedwigia filiformis (as Anictangium) Jones (1933) treated Hedwigia integrifolia as a synonym of Hedwigia ciliata. Not only do both taxa have ciliate perichaetial leaves, but Jones (1933), however, considered Hedwigia integrifolia a synonym of Hedwigia ciliata. Dalton et al (2012) clarified the concept of H. integrifolia. The type they examined ("Nort America, Beauvois", E) presents all characters that place the species in Hedwigia. The type material has leaves with a hyaline acumen, the leaf cells have branched papillae, and the perichaetial leaves are ciliate. This taxon in Hedwigia is different from the taxon in Hedwigidium which is correctly known as Hedwigidium imberbe (Sm.) Bruch & Schimp., Bryol. Eur. 3: 157. 1846. The type of Gymnostomum imberbe Sm. was collected in western Ireland and presents characters that define Hedwigidium: stoloniferous gametophytes, ovate leaves with concolorous short acute apices, leaf cell papillae marginal, perichaetial leaves with margins undulate, and phaneroporic stomata on the urn base (De Luna 2021). Photos of H. ciliata in Ohio actualy seem to correspond to this species, H. integrifolia.
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Hedwigia integrifolia, from the website of the Ohio Moss and Lichen Association (listed there as Hedwigia ciliata). |
3. Hedwigia nivalis (Müll. Hal.) Mitt. ---- Muller 1851, v2 p 165 29. P. nivale C. Miill.; monoicum ; P. ciliato simillmum, amocnc incano-lutcsccns, sed foliorum cellulae dolioliformes minores ad latcra maxime crenulate valde obscurae et scaberrimae, basi longiores distincte papillosae; pericli. basi multo laxius ct tenerius reticulata; thcca pachydcrmiss castanea , apophysi polytrichoideo- disciformi - coarctata pracdita. Schistitlium ciliatum Hmp. Lnn. XX. p. 70. — Anoectangium ciliatum C. Miill. Linn. XIX. p. 1999. Patria. Venezuela, prov. Merida, in montibus nivosis: Moritz. No. 174. Distinctissima species! Calyptra P. ciliati , glabra. ----- Distributed in the SW United States south to Mexico and South America. Also the Caribean, Middle Asia, Southern Africa, Western Indian Ocean; Australia, New Zealand (Allen, 2010). Reported as H. ciliata var nivalis Hampe from Argentina (Biasuso, 2007). According to Buck and Norris (1996) specimens from Latin America routinelly identified as Hedwigia ciliata most likely belong to Hedwigia nivalis. This species was recognized in Central America (Allen, 2010), where H. nivalis is best distinguished from H. ciliata by the presence of incurved upper leaf margins and non-hairy calyptrae. The leaf apex of H. ciliata can be channeled, but the upper leaf margins are plane or erect rather than incurved, and its calyptrae are hairy. Type of H nivalis at NHM, UK. 30. P. subrevolufuin C. Mulll.-, monoicum ; P. ciliato similliimum, sed folia basi tantum marglne revoluta, cellulis dolioliformis maxiec crenulatis et obscuris valdc paplllosis minorlbus, basi parictibus interruptis, punctu- latis; perich. bast laxius reticulata; theca collo angustiore longiore praedita minor, operculo conico-cupulato. Patria. Mexico, Harrisonlae sccundae associatum ad saxa pr. Malpays dc la Joya: Schiede. In locis non indicatis ibidem legit C. Ehrcnbcrg. Calyptra P. ciliati, glabra. Distinctissima species!
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Hedwigia nivalis, figure from Biasuso 2007.
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4. Hedwigia stellata Hedenäs, J. Bryol. 18: 144, figs. 1–2.1994. Almost circumboreal. Western North America (California to Oregon and Washington) Europe (Denmark, Italy, Norway, Portugal, Spain, Sweden, United Kingdom). Reported as H. stellata from Turkey (Erdag et al, 2003). Leaf of H. stellata in California, US. CalPhotos Photo Database Habit, H. stellata in Spain. FB Habits, H. stellata in Spain. https://www.biodiversidadvirtual.org/herbarium/BFI-Hedwigia-cat12600.html Habit, H. stellata in Gard, France. Mousses du Gard Habit, H. stellata in Germany. http://www.moose-deutschland.de/organismen/hedwigia-stellata
Specimens: California, US. Slopes of Mt. Knocti, Lake, Co. J. R. Shevock & K. Heise & J. Harpel & D. Toren #20598. (Berkeley herbarium) |
Hedwigia stellata, on rock; from the web site Mosses & Liverworts of the Outer Hebrides
Hedwigia stellata, figure from Flora Briofitica Iberica. |
5. Hedwigia detonsa (M. Howe) W. R. Buck & D. H. Norris, Nova Hedwigia 62: 368. 1996. Only kown from California, US. Perichaetial leaves broadly channeled to involute, margins entire. Figure from Flora of North America.
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6. Hedwigia brevipilifera Biasuso, Lindbergia 32: 16, figs. 1,2,5. 2007. Only known from Argentina. The specific epithet refers to the short hyaline acumen and hair point.
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Hedwigia brevipilifera, figure from Biasuso (2007) |
7. Hedwigia striata (Wilson ex Hook.) John Whitehead & J. Fergusson ex Hobk. & Porritt, Naturalist (Hull) 5: 29. 1879. sensu Blockeel T.L. and S. Bosanquet. 2016. Hedwigia striata in Britain sensu TROPICOS.ORG is: Hedwigia striata (Bruch & Schimp.) John Whitehead & Fergusson Hedwigia albicans var. striata (Bruch & Schimp.) Dalla Torre & Sarnth. Spain, Portugal. Hespanhol et al. (2013) and Gallego et al. (2014) have documented its occurrence in Portugal and Spain. xxxx we describe the differentiating characters of H. striata and document the known British and Irish records. In H. stellata and H. ciliata, the cilia are numerous, long and contorted, often abruptly bent or curled and sometimes bear lateral projections (Fig. 2). In H. striata the cilia are fewer, shorter and more or less straight Gallego et al. (2014) pointed out that in H. striata the marginal cells at the base of the leaf just above the insertion are often ornamented by prominent papillae and/or double teeth. The same cells are smooth in H. stellata and H. ciliata. The Naturalist n.s. 5. no L. Manchester Oryptogamic Society. — p. 29 Meeting, Aug. 11th, the president in the chair. — "The mosses exhibited by the president were Hedivigia striata, Wils, from Grasmere, which he, along with Mr. Fergusson, proposed to restore again to the rank of a distinct species; " no. LI. Vol 5. Manchester Oryptogamic Society. — Monthly meeting, Mr. John Whitehead, president, in the chair. p 46: "With regard to Hedwigia striata, exhibited at the last meeting, from near Grasmere, the president said that in looking through Mr. Ashton's collection he had found a fruiting specimen gathered in Llanberis Pass. This was a fortunate discovery, as the moss was extremely rare — three localities only being known for it. It was first found by the late Mr. W. Wilson in 1829, at Llyn Idwal, and named by him Ancectangium striatum. " The most important morphological character state of Hedwigia striata is the longitudinally plicate leaf (Gallego et al 2014). According to Jones (1933) there is Hedwigia cilata forma striata (Wils.) n. comb. in the US. Leaves yellowish green, longitudinally srtiate; margins strongly recurved; lid conical.
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Hedwigia striata, figure from Gallego et al (2014) |
8. Hedwigia nemoralis Ignatova, Fedosov & Ignatov Occurs in southern regions of Russia, from the Caucasus to the Russian Far East, with one locality in Central European Russia. It was also revealed from eastern North America. It is characterized by a small plant size, shortly acuminate, secund leaves with short, denticulate hyaline hair points and weakly ciliate perichaetial leaves.
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9. Hedwigia czernyadjevae Ignatova, Fedosov & Ignatov Wide distribution in Transbaikalia, Yakutia and high mountains of the Russian Far East, although it never occurs frequently. Previously its specimens were identified as H. stellata, but straight leaf apices, strongly incrassate and porose laminal cells and hairy calyptrae differentiate H. czernyadjevae from this species, in addition to sequence data, which place the Siberian species closer to American H. detonsa. |
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10. Hedwigia mollis Ignatova, Fedosov & Ignatov Shortly recurved leaf margins, weakly sinuose laminal cells and spores 20–25(–28) um. It was revealed in European Russia, from the southern Murmansk Province and Karelia to the Caucasus, and from South Urals and Altai Mts. |
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11. Hedwigia emodica Hampe ex Müll. Hal. Numerous collections in herbaria from the Caucasus and Asian Russia are referred to H. emodica, which turned out to be rather uniform by morphology. In addition to long hyaline leaf apices, it is characterized by straight leaves, almost plane leaf margins and dense papillae with thick, obtuse branches. Hedwigia emodica var. echinata Ignatova & Ignatov |
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12. Hedwigia kuzenevae Ignatova & Ignatov Most similar to Hedwigia czernyadjevae in straight leaves with leaf margins recurved up to hair-point and long, slighly flexuose, strongly papillose and spinulose on dorsal surface hairpoints; their main distinctive character is multipapillose vs. uniopapillose laminal cells. The distinction of H. kuzenevae from H. ciliata includes straight vs. falcatesecund |
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References: Allen B. 2010. Moss flora of Central America. Part 3: Anomodontaceae - Symphyodontaceae / B.Allen, with contributions from R.A.Purcell, R.R.Ireland. - St. Louis: Missouri Botanical Garden, 2010. - x, 731 p.: ill. - (Monographs in systematic botany from the Missouri Botanical Garden; Vol.117). - Bibliogr.: p.703-714. - Ind.: p.715-731.ISBN 978-1-930723-95-5; ISSN 0161-1542 Biasuso, B. 2007. The genus Hedwigia (Hedwigiaceae, Bryophyta) in Argentina. Lindbergia 31(1):5-17. Brotherus, V. F. 1909. Hedwigiaceae. In: A. Engler & K. Prantl, eds. Die natürlichen Pflanzenfamilien 1(3): 712–722. W. Engelman, Leipzig. Bruch, P., Schimper, W. P. & Gümbel, W. v. T. 1846. Bryologia Europaea, seu Genera Muscorum Europaeorum monographice illustrata. Vol. III. E. Schweizerbart, Stuttgart. Dalton, N., Kungu, E.M. & Long, D.G. 2012. The misapplication of Hedwigia integrifolia P. Beauv and identity of Gymnostomum imberbe Sm. (Hedwigiaceae, Bryopsida). Journal of Bryology, 34(1): 59–61. De Luna, E. 2021. Seta length variation and the refutation of Hedwigidium = Braunia (Hedwigiaceae, Bryopsida). Acta Botanica Mexicana 128: e1810. DOI: https://doi.org/10.21829/abm128.2021.1810. Erdag A, M. Kirmaci & H. Kürschner. 2003. The Hedwigia ciliata (Hedw.) Ehr. ex P.Beauv. Complex in Turkey, with a New Record, H. ciliata var. leucophaea Bruch & Schimp. (Hedwigiaceae, Bryopsida). Turk J Bot. 27: 349-356. Gallego MT, MJ Cano & J. Guerra. 2014. New chorological data on Hedwigia striata (Bruch & Schimp.) Bosw., (Hedwigiaceae, Musci) in Spain, and remarks on its morphological characters. Annales de Biología 36: 5-9. Hedenäs, L. 1994. The Hedwigia ciliata complex in Sweden, with notes on the occurrence of the taxa in Fennoscandia. Journal of Bryology 18: 139-157. Ignatova, E. A., O. I. Kuznetsova, V. E. Fedosov and M. S. Ignatov. 2016. On the genus Hedwigia (Hedwigiaceae, Bryophyta) in Russia. Arctoa 25: 241-277. Ignatova, E.A., O.I. Kuznetsova & M.S. Ignatov. 2017. Further comments on the genus Hedwigia (Hedwigiaceae, Bryophyta) in Russia. Arctoa 26(1): 132–143. Jaeger A. 1876. Genera et species muscorum systematice disposita seu Adumbratio florae muscorum totius orbis terrarum. VI, p 85-188. |
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Discussion from: Central American Mosses Hedwigia has often been considered a monospecific genus (see Hedenäs 1994 and Buck & Norris 1996), but presently it contains at least five taxa (H. ciliata (Hedw.) P. Beauv., H. ciliata var. leucophaea The genus lacks a stem central strand, paraphyllia, leaf costa, and peristome. Its primary, creeping stems are so poorly developed they are often difficult to demonstrate, and its secondary stems and branches rarely produce stolons. Hedwigia characteristically has broadly ovate, acuminate leaves that end in broad, hyaline acumina, densely papillose leaf cells, immersed, cupulate capsules, and cryptoporic stomata. Although H. detonsa has entire perichaetial leaves, all other species of Hedwigia have long-ciliate perichaetial leaves. Hedwigidium and Hedwigia have immersed capsules, and their gametophytes are very similar. Hedwigidium differs from Hedwigia in often having stoloniferous gametophytes, concolorous or narrowly |
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Records Hedwigia in France: https://inpn.mnhn.fr/espece/cd_nom/5072 H. ciliata in Italy with photos: http://www.floraitaliae.actaplantarum.org/viewtopic.php?t=87752 H. ciliata in Mecklenburg, Germany: https://www.korseby.net/outer/flora/bryophyta/hedwigiaceae/index.html H. ciliata leaves in British Columbia, Canada H. ciliata and H stellata in Czech Republic https://botanika.prf.jcu.cz/bryoweb/en/results.php?jmeno=Hedwigia
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