Abstract
Isoetes jarmaniae sp. nov. is described as a new lycophyte endemic to Tasmania, Australia, where it is confined to peat-bound karstic wetlands in several river valleys in the south-west wilderness. While seemingly morphologically closest to I. drummondii, this quillwort has features that are globally uncommon in Isoetes and unknown in other Australasian taxa. Most notable are its markedly flattened, strongly recurved leaves and disproportionately large sporangium ligules that are more suggestive of South American than Australian taxa. As well, the exceptionally thin and wide (alate) megaspore equatorial ridge is swollen at suture intersections, presenting a slightly triangular shape suggestive of the Indian taxon I. udupiensis. The microspores of I. jarmaniae exhibit exceptionally, perhaps uniquely, fine-papillate ornamentation. An original key placing I. jarmaniae in context with the other Tasmanian Isoetes species is provided. This diminutive, apparently diploid species is evidently maintaining a self-sustaining population within a regionally unique habitat and small geographic range.
References
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<p>Bentham, G. (1878) <em>Flora Australiensis: a description of the plants of the Australian territory Volume 7: Roxburghiaceae to Filices</em>. Reeves & Company, London. https://doi.org/10.1017/CBO9781139096072.003</p>
<p>Braun, A. (1853) Plantae Muellerianae, Beitrag zur Flora Sudaustraliens, Isoetes Rich. <em>Linnaea; Ein Journal für die Botanik in ihrem ganzen Umfange</em> 25 (6): 722. [https://www.biodiversitylibrary.org/item/109556#page/728/mode/1up]</p>
<p>Braun, A. (1868) Uber die australischen Arten der Gattung Isoetes. <em>Monatsberichte der Königlich preussischen Akademie der Wissenschaften zu Berlin</em> 13: 523–546. [https://www.biodiversitylibrary.org/page/35989553#page/572/mode/1up]</p>
<p>Brown, M.J., Crowden, R.K. & Jarman, S.J. (1982) Vegetation of an alkaline pan − acidic peat mosaic in the Hardwood River Valley, <em>Tasmania. Australian Journal of Ecology</em> 7: 3−12. https://doi.org/10.1111/j.1442-9993.1982.tb01295.x</p>
<p>Brunton, D.F. & Britton, D.M. (2006) <em>Isoetes</em> ×<em>novae-angliae</em>, a new quillwort (Isoetaceae) from northeastern North America. <em>Rhodora</em> 108: 228-241. https://doi.org/10.3119/0035-4902(2006)108[228:INIAAH]2.0.CO;2</p>
<p>Brunton, D.F. & Troia, A. (2018) Global review of recent taxonomic research into <em>Isoetes</em> (Isoetaceae), with implications for biogeography and conservation. <em>Fern Gazette</em> (UK) 20 (Part 8): 309–333.</p>
<p>Butler, G.D. (1878) The Species of Isoetes of the Indian Territory. <em>Botanical Gazette</em> 3: 1–2. https://doi.org/10.1086/325109</p>
<p>Chinnock, R.J. (1998) <em>Isoetaceae</em>, in <em>Flora of Australia</em>, Volume 48. Australian Biological Resources Study, Department of the Environment and Energy, Canberra.</p>
<p>Curtis, W.M. (1985) New species of Tasmanian Monocotyledones in the families Juncaceae, Centrolepidaceae and Cyperaceae. <em>Burnonia</em> 7 (2): 297–304. https://doi.org/10.1071/BRU9840297</p>
<p>Field, A.R. (2020) Classification and typification of Australian lycophytes and ferns based on Pteridophyte Phylogeny Group classification PPG I. <em>Australian Systematic Botany</em> 33: 1–102. https://doi.org/10.1071/SB18011</p>
<p>Fuchs, H.P. (1960) Una nueva especie de Isoetes de Columbia y algunas notas sabre nuevas colecciones de Isoetes killpii <em>Caldasia</em> 8 (38): 314. [https://revistas.unal.edu.co/index.php/cal/article/view/33030]</p>
<p>Garrett, M. (1996) <em>The ferns of Tasmania; their ecology and distribution</em>. Tasmanian Forest Research Council Inc., Hobart.</p>
<p>Garrett, M. & Kantvilas, G. (1992) Morphology, ecology and distribution of <em>Isoetes</em> L. in Tasmania. <em>Papers and Proceedings of the Royal Society of Tasmania</em> 126: 115–122. https://doi.org/10.26749/rstpp.126.115</p>
<p>Gomez, L.D. (1981) Vegetative reproduction in a Central American <em>Isoëtes</em> (Isoetaceae): its morphological, systematic and taxonomical significance. <em>Brenesia</em> 18 (4): 1–13.</p>
<p>Harris, S. & Kitchener, A. (2005) <em>From Forest to Fjaeldmark. Descriptions of Tasmania’s Vegetation</em>. Department of Primary Industries, Water and Environment, Printing Authority of Tasmania, Hobart.</p>
<p>Hickey, R.J. (1986) <em>Isoetes</em> megaspore surface morphology: nomenclature, variation and systematic importance. <em>American Fern Journal</em> 76 (1): 1–16. https://doi.org/10.2307/1547394</p>
<p>Hickey, R.J. (1994) Isoetaceae, in R.M Tryon & R.G. Stolze. <em>Pteridophytes of Peru</em>, Part VI. <em>Fieldiana</em> (New Series) 34: 88–97.</p>
<p>Hooker, J.D. (1847) Flora Tasmaniae Spicilegium, or, Contributions towards a flora of Van Diemen’s Land. <em>London Journal of Botany</em> 6: 461–479. [https://www.biodiversitylibrary.org/page/2925973#page/520/mode/1up]</p>
<p>Hooker, J.D. (1858) <em>Flora Tasmaniae, Part 2.</em> Reeves Brothers, London.</p>
<p>IUCN Standards and Petitions Committee (2019) Guidelines for Using the IUCN Red List Categories and Criteria. Version 14. Prepared by the Standards and Petitions Committee. Available from: http://www.iucnredlist.org/documents/RedListGuidelines.pdf (accessed 4 October 2021)</p>
<p>Jarman, S.J. & Crowden, R.K. (1978) <em>A survey of vegetation. Lower Gordon Scientific Survey</em>. Hydro-Electric Commission, Hobart.</p>
<p>Jarman, S.J. & Crowden, R.K. (1979) An examination of vegetation from the Lower Gordon River and associated catchments against a background of regional botany.<em> In: </em>Hale, G.E.A. (ed.) <em>Lower Gordon Region. Land Use, Resources and Special Features</em>. Hydro-Electric Commission, Hobart.</p>
<p>Jarman, S.J., Kantvilas, G. & Brown, M.J. (1988) <em>Buttongrass Moorland in Tasmania</em>. Research Report 2. Tasmanian Forest Research Council Inc., Hobart.</p>
<p>Johnson, E.R.L. (1984) Taxonomic revision of <em>Isoetes</em> L. in Western Australia. <em>Journal of the Royal Society of Western Australia</em> 67 (1): 23–43.</p>
<p>Koyama, T. (1956) Taxonomic study of Cyperaceae V. <em>Botanical Magazine</em> (Tokyo) 68: 59–67. https://doi.org/10.15281/jplantres1887.69.59</p>
<p>Limpricht, K.G. (1899) <em>Die Laubmoose Deutschland, Volume 3</em>. Leipzig.</p>
<p>Marsden, C.R.<strong> (</strong>1979) <em>Morphology and Taxonomy of Isoetes in Australia, India, North-East and South-East Asia, China and Japan</em>. Department of Botany, University of Adelaide [Phd Thesis].</p>
<p>Pfeiffer, N.E. (1922) Monograph on the Isoetaceae. <em>Annals of the Missouri Botanical Garden</em> 9: 79–233. https://doi.org/10.2307/2990000</p>
<p>Pant, D.D., Srivastava, G.P. & Srivastava, P.C. (2000) Ligule in Lycopods – A Review. <em>Proceedings of the Indian Natural Sciences Academy</em> Part B 66:195–222. https://doi.org/10.2307/2990000</p>
<p>Paris, E.G. (1904) <em>Index bryologicus sive enumeratio muscorum ad diem ultimam anni 1900 cognitorum adjunctis synonymia distributioneque geographica locupletissimis.</em> Librairie Scientifique A. Hermann, Paris.</p>
<p>Proemse, B.C., Eberhard, R.S., Sharples, C., Bowman, J.P., Richards, K., Comfort, M. & Barmuta, L.A. (2017) Stromatolites on the rise in peat-bound karstic wetlands. <em>Scientific Reports</em> 7: 1−8. https://doi.org/10.1038/s41598-017-15507-1</p>
<p>Rodway, L. (1898) Notes on a new Trithuria (Plate). <em>Papers and Proceedings of the Royal Society of Tasmania for 1897 (1898)</em> 27: 47–48.</p>
<p>Shukla, P.K., Srivastava, G.K., Shukla, S.K. & Rajagopal, P.K. (2005) Two new species of the genus <em>Isoetes</em> L. (Isoetaceae–Lycopsida) from India. <em>Taxon</em> 54 (1): 109–116. https://doi.org/10.2307/25065306</p>
<p>Singh, S.K., Shukla, P.K., Brunton, D.F., Dubney, N.K. & Shukla, S.K. (2021) The taxonomy and conservation status of <em>Isoetes</em> (Isoetaceae; Lycopodiopsida) in India. <em>Botany Letters</em> 168 (2): 200–226. https://doi.org/10.1080/23818107.2020.1848621</p>
<p>Small, R.L. & Hickey, R.J. (2001) Systematics of the Northern Andean <em>Isoetes karstenii</em> Complex. <em>American Fern Journal</em> 91: 41–69. https://doi.org/10.1640/0002-8444(2001)091[0041:SOTNAI]2.0.CO;2</p>
<p>Tasmanian State Government (2021) <em>Tasmanian Species Protection Act 1995-83: Schedule 5-Taxa of native flora and fauna which are rare</em>. Available from: https://www.legislation.tas.gov.au/view/html/inforce/current/act-1995-083#JS5@HS2@HT3@EN (accessed 13 June 2021)</p>
<p>Thiers, B. (2021) [continuously updated] <em>Index Herbariorum: A global directory of public herbaria and associated staff.</em> New York Botanical Garden’s Virtual Herbarium. Available from: http://sweetgum.nybg.org/science/ih (accessed 4 October 2021)</p>
<p>Troia, A. (2001) The genus Isöetes L. (Lycophyta, Isöetaceae): synthesis of kayological data. <em>Webbia</em> 56: 201–218. https://doi.org/10.1080/00837792.2001.10670712</p>
<p>Troia, A. & Azzella, M.M. (2013) <em>Isoetes sabatina</em> (Isoetaceae, Lycopodiophyte), a new aquatic species from central Italy. <em>Plant Biosystems</em> 147: 1052–1058. https://doi.org/10.1080/11263504.2013.782902</p>
<p>Troia, A., Pereira, A.B, Kim, C. & Taylor, W.C. (2016) The genus <em>Isoetes </em>(Isoetaceae): a provisional checklist of the accepted and unresolved taxa. <em>Phytotaxa</em> 277: 101–145. https://doi.org/10.11646/phytotaxa.277.2.1</p>
<p>Wood, D., Besnard, G., Beerling, D.J., Osborne, C.P. & Christin, P.-A. (2020) Phylogenomics indicates the “living fossil” <em>Isoetes</em> diversified in the Cenozoic<em>. PLoS ONE</em> 15 (6): e0227525. https://doi.org/10.1371/journal.pone.0227525</p>
<p>Bentham, G. (1878) <em>Flora Australiensis: a description of the plants of the Australian territory Volume 7: Roxburghiaceae to Filices</em>. Reeves & Company, London. https://doi.org/10.1017/CBO9781139096072.003</p>
<p>Braun, A. (1853) Plantae Muellerianae, Beitrag zur Flora Sudaustraliens, Isoetes Rich. <em>Linnaea; Ein Journal für die Botanik in ihrem ganzen Umfange</em> 25 (6): 722. [https://www.biodiversitylibrary.org/item/109556#page/728/mode/1up]</p>
<p>Braun, A. (1868) Uber die australischen Arten der Gattung Isoetes. <em>Monatsberichte der Königlich preussischen Akademie der Wissenschaften zu Berlin</em> 13: 523–546. [https://www.biodiversitylibrary.org/page/35989553#page/572/mode/1up]</p>
<p>Brown, M.J., Crowden, R.K. & Jarman, S.J. (1982) Vegetation of an alkaline pan − acidic peat mosaic in the Hardwood River Valley, <em>Tasmania. Australian Journal of Ecology</em> 7: 3−12. https://doi.org/10.1111/j.1442-9993.1982.tb01295.x</p>
<p>Brunton, D.F. & Britton, D.M. (2006) <em>Isoetes</em> ×<em>novae-angliae</em>, a new quillwort (Isoetaceae) from northeastern North America. <em>Rhodora</em> 108: 228-241. https://doi.org/10.3119/0035-4902(2006)108[228:INIAAH]2.0.CO;2</p>
<p>Brunton, D.F. & Troia, A. (2018) Global review of recent taxonomic research into <em>Isoetes</em> (Isoetaceae), with implications for biogeography and conservation. <em>Fern Gazette</em> (UK) 20 (Part 8): 309–333.</p>
<p>Butler, G.D. (1878) The Species of Isoetes of the Indian Territory. <em>Botanical Gazette</em> 3: 1–2. https://doi.org/10.1086/325109</p>
<p>Chinnock, R.J. (1998) <em>Isoetaceae</em>, in <em>Flora of Australia</em>, Volume 48. Australian Biological Resources Study, Department of the Environment and Energy, Canberra.</p>
<p>Curtis, W.M. (1985) New species of Tasmanian Monocotyledones in the families Juncaceae, Centrolepidaceae and Cyperaceae. <em>Burnonia</em> 7 (2): 297–304. https://doi.org/10.1071/BRU9840297</p>
<p>Field, A.R. (2020) Classification and typification of Australian lycophytes and ferns based on Pteridophyte Phylogeny Group classification PPG I. <em>Australian Systematic Botany</em> 33: 1–102. https://doi.org/10.1071/SB18011</p>
<p>Fuchs, H.P. (1960) Una nueva especie de Isoetes de Columbia y algunas notas sabre nuevas colecciones de Isoetes killpii <em>Caldasia</em> 8 (38): 314. [https://revistas.unal.edu.co/index.php/cal/article/view/33030]</p>
<p>Garrett, M. (1996) <em>The ferns of Tasmania; their ecology and distribution</em>. Tasmanian Forest Research Council Inc., Hobart.</p>
<p>Garrett, M. & Kantvilas, G. (1992) Morphology, ecology and distribution of <em>Isoetes</em> L. in Tasmania. <em>Papers and Proceedings of the Royal Society of Tasmania</em> 126: 115–122. https://doi.org/10.26749/rstpp.126.115</p>
<p>Gomez, L.D. (1981) Vegetative reproduction in a Central American <em>Isoëtes</em> (Isoetaceae): its morphological, systematic and taxonomical significance. <em>Brenesia</em> 18 (4): 1–13.</p>
<p>Harris, S. & Kitchener, A. (2005) <em>From Forest to Fjaeldmark. Descriptions of Tasmania’s Vegetation</em>. Department of Primary Industries, Water and Environment, Printing Authority of Tasmania, Hobart.</p>
<p>Hickey, R.J. (1986) <em>Isoetes</em> megaspore surface morphology: nomenclature, variation and systematic importance. <em>American Fern Journal</em> 76 (1): 1–16. https://doi.org/10.2307/1547394</p>
<p>Hickey, R.J. (1994) Isoetaceae, in R.M Tryon & R.G. Stolze. <em>Pteridophytes of Peru</em>, Part VI. <em>Fieldiana</em> (New Series) 34: 88–97.</p>
<p>Hooker, J.D. (1847) Flora Tasmaniae Spicilegium, or, Contributions towards a flora of Van Diemen’s Land. <em>London Journal of Botany</em> 6: 461–479. [https://www.biodiversitylibrary.org/page/2925973#page/520/mode/1up]</p>
<p>Hooker, J.D. (1858) <em>Flora Tasmaniae, Part 2.</em> Reeves Brothers, London.</p>
<p>IUCN Standards and Petitions Committee (2019) Guidelines for Using the IUCN Red List Categories and Criteria. Version 14. Prepared by the Standards and Petitions Committee. Available from: http://www.iucnredlist.org/documents/RedListGuidelines.pdf (accessed 4 October 2021)</p>
<p>Jarman, S.J. & Crowden, R.K. (1978) <em>A survey of vegetation. Lower Gordon Scientific Survey</em>. Hydro-Electric Commission, Hobart.</p>
<p>Jarman, S.J. & Crowden, R.K. (1979) An examination of vegetation from the Lower Gordon River and associated catchments against a background of regional botany.<em> In: </em>Hale, G.E.A. (ed.) <em>Lower Gordon Region. Land Use, Resources and Special Features</em>. Hydro-Electric Commission, Hobart.</p>
<p>Jarman, S.J., Kantvilas, G. & Brown, M.J. (1988) <em>Buttongrass Moorland in Tasmania</em>. Research Report 2. Tasmanian Forest Research Council Inc., Hobart.</p>
<p>Johnson, E.R.L. (1984) Taxonomic revision of <em>Isoetes</em> L. in Western Australia. <em>Journal of the Royal Society of Western Australia</em> 67 (1): 23–43.</p>
<p>Koyama, T. (1956) Taxonomic study of Cyperaceae V. <em>Botanical Magazine</em> (Tokyo) 68: 59–67. https://doi.org/10.15281/jplantres1887.69.59</p>
<p>Limpricht, K.G. (1899) <em>Die Laubmoose Deutschland, Volume 3</em>. Leipzig.</p>
<p>Marsden, C.R.<strong> (</strong>1979) <em>Morphology and Taxonomy of Isoetes in Australia, India, North-East and South-East Asia, China and Japan</em>. Department of Botany, University of Adelaide [Phd Thesis].</p>
<p>Pfeiffer, N.E. (1922) Monograph on the Isoetaceae. <em>Annals of the Missouri Botanical Garden</em> 9: 79–233. https://doi.org/10.2307/2990000</p>
<p>Pant, D.D., Srivastava, G.P. & Srivastava, P.C. (2000) Ligule in Lycopods – A Review. <em>Proceedings of the Indian Natural Sciences Academy</em> Part B 66:195–222. https://doi.org/10.2307/2990000</p>
<p>Paris, E.G. (1904) <em>Index bryologicus sive enumeratio muscorum ad diem ultimam anni 1900 cognitorum adjunctis synonymia distributioneque geographica locupletissimis.</em> Librairie Scientifique A. Hermann, Paris.</p>
<p>Proemse, B.C., Eberhard, R.S., Sharples, C., Bowman, J.P., Richards, K., Comfort, M. & Barmuta, L.A. (2017) Stromatolites on the rise in peat-bound karstic wetlands. <em>Scientific Reports</em> 7: 1−8. https://doi.org/10.1038/s41598-017-15507-1</p>
<p>Rodway, L. (1898) Notes on a new Trithuria (Plate). <em>Papers and Proceedings of the Royal Society of Tasmania for 1897 (1898)</em> 27: 47–48.</p>
<p>Shukla, P.K., Srivastava, G.K., Shukla, S.K. & Rajagopal, P.K. (2005) Two new species of the genus <em>Isoetes</em> L. (Isoetaceae–Lycopsida) from India. <em>Taxon</em> 54 (1): 109–116. https://doi.org/10.2307/25065306</p>
<p>Singh, S.K., Shukla, P.K., Brunton, D.F., Dubney, N.K. & Shukla, S.K. (2021) The taxonomy and conservation status of <em>Isoetes</em> (Isoetaceae; Lycopodiopsida) in India. <em>Botany Letters</em> 168 (2): 200–226. https://doi.org/10.1080/23818107.2020.1848621</p>
<p>Small, R.L. & Hickey, R.J. (2001) Systematics of the Northern Andean <em>Isoetes karstenii</em> Complex. <em>American Fern Journal</em> 91: 41–69. https://doi.org/10.1640/0002-8444(2001)091[0041:SOTNAI]2.0.CO;2</p>
<p>Tasmanian State Government (2021) <em>Tasmanian Species Protection Act 1995-83: Schedule 5-Taxa of native flora and fauna which are rare</em>. Available from: https://www.legislation.tas.gov.au/view/html/inforce/current/act-1995-083#JS5@HS2@HT3@EN (accessed 13 June 2021)</p>
<p>Thiers, B. (2021) [continuously updated] <em>Index Herbariorum: A global directory of public herbaria and associated staff.</em> New York Botanical Garden’s Virtual Herbarium. Available from: http://sweetgum.nybg.org/science/ih (accessed 4 October 2021)</p>
<p>Troia, A. (2001) The genus Isöetes L. (Lycophyta, Isöetaceae): synthesis of kayological data. <em>Webbia</em> 56: 201–218. https://doi.org/10.1080/00837792.2001.10670712</p>
<p>Troia, A. & Azzella, M.M. (2013) <em>Isoetes sabatina</em> (Isoetaceae, Lycopodiophyte), a new aquatic species from central Italy. <em>Plant Biosystems</em> 147: 1052–1058. https://doi.org/10.1080/11263504.2013.782902</p>
<p>Troia, A., Pereira, A.B, Kim, C. & Taylor, W.C. (2016) The genus <em>Isoetes </em>(Isoetaceae): a provisional checklist of the accepted and unresolved taxa. <em>Phytotaxa</em> 277: 101–145. https://doi.org/10.11646/phytotaxa.277.2.1</p>
<p>Wood, D., Besnard, G., Beerling, D.J., Osborne, C.P. & Christin, P.-A. (2020) Phylogenomics indicates the “living fossil” <em>Isoetes</em> diversified in the Cenozoic<em>. PLoS ONE</em> 15 (6): e0227525. https://doi.org/10.1371/journal.pone.0227525</p>