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Pteridophyte taxa known or reported from Sulawesi
Pteris aquilina
EOL Text
Bracken (Pteridium aquilinum) is a conspicuous fern that forms large clonal colonies in a variety of habitats. The large, more or less triangular leaves develop from fiddleheads that develop widely spaced along the branches of an extensive subterranean rhizome that may reach nearly 400 m in length. The taxonomy of the genus remains controversial, but most botanists currently favor a classification involving five or more species. In this sense, Pteridium aquilinum is distributed widely in mostly the northern hemisphere, in both the New and Old Worlds.
Bracken produces a pharmacopeia of toxic compounds, including: thiaminase (which breaks down the amino acid thiamine and results in vitamin B deficiency), ecdysomes (hormones that stimulate uncontrolled early molting in insects), tannins (which bind to proteins and other compounds), and hydrogen cyanide, and also produces carcinogenic compounds. The combination of chemicals renders the plants toxic to most animals, both invertebrates and vertebrates, although some insect specialists ingest bracken tissue to become poisonous to their predators. Humans have long eaten the fiddleheads (emerging young leaves) of bracken, but over-ingestion of fresh or dried fronds has been linked to stomach and esophageal cancers.
Bracken is an aggressive colonizer of disturbed and other successional habitats and has been considered an invader of pastureland. It has been shown to be allelopathic (to produce compounds that inhibit the growth of other plant species) and can for dense monocultures. It is difficult to eradicate or control and, because it is toxic, renders such pasturage unfit for grazing. Pteridium aquilinum is considered a noxious weed, especially in portions of Great Britain and mainland Europe.
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Rights holder/Author | Missouri Botanical Garden, George Yatskievych |
Source | No source database. |
The species Pteridium aquilinum is considered globally Secure. Conversely, it is considered a noxious weed (officially or unofficially) in portions or North America, Europe, New Zealand, and Australia.
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Rights holder/Author | George Yatskievych |
Source | No source database. |
Pteridium aquilinum is a terrestrial species that occurs in a wide variety of habitats, ranging from sand dunes to peatlands and open meadows to forests. It is especially aggressive in disturbed or successional habitats, including those prone to periodic disturbance by natural processes such as fires. Human-mediated perturbations such as grazing and timber removal tend to stimulate its growth and spread. It occurs from sea level to ca. 3300 m in soils derived from a wide variety of substrates and with various pH levels.
License | http://creativecommons.org/licenses/by/3.0/ |
Rights holder/Author | George Yatskievych |
Source | No source database. |
License | http://creativecommons.org/licenses/by/3.0/ |
Rights holder/Author | George Yatskievych |
Source | No source database. |
- BioImages (extensive images from Britain plus table of associated invertebrates)
- USDA Plants Profile: Pteridium aquilinum (extensive summary information and several images from the U.S.A.)
- International Bracken Group
- Native American ethnobotany
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Rights holder/Author | George Yatskievych |
Source | No source database. |
In the New World, Pteridium aquilinum in the strict sense occurs nearly throughout the United States and southern Canada, with extensions northward to about 55° N Latitude. The species is distributed southward through the mountains of Mexico into Central America as far as Honduras. It is also present in the Caribbean Islands and one subspecies is endemic to Hawaii. In the Old World, P. aquilinum occurs nearly throughout Europe (north to near the Arctic Circle) and Africa (excluding the Saharan portion), is present on a number of islands, including Madagascar, Réunion, the Mascarenes, Mauritius, and Macaronesia. Its Asian distribution extends eastward across Russia and south through China and Indochina into Malaysia. Farther south in both the Old and New Worlds, other species of Pteridium replace P. aquilinum, but there is distributional overlap between P. aquilinum and the other taxa, mainly in Latin America and portions of Asia.
License | http://creativecommons.org/licenses/by/3.0/ |
Rights holder/Author | George Yatskievych |
Source | No source database. |
Alonso-Amelot, M. E., U. F. Castillo, M. Avedaño, B. L. Smith and D. R. Lauren. 2000. Milk as a vehicle for the transfer of ptaquiloside, a bracken carcinogen. Chapter 12, pp. 86. J. A. Taylor and R. T. Smith, eds. Bracken Fern: Toxicity, Biology and Control. International Bracken Group Special Publication, vol. 4, Aberystwyth, Wales.
Bridges, K. M., C. J. Ashcroft, and E. Sheffield. 1998. Population analysis of the type localities of some recently recognised taxa of British Pteridium (Dennstaedtiaceae: Pteridophyta). Fern Gaz. 15: 205-213.
Brown, R. W. 1995. Bracken and the ecology of Lyme disease. Chapter 24, pp. 116-119 in R. T. Smith and J. A. Taylor, eds. Bracken: An Environmental Issue. International Bracken Group Special Publication, vol. 2, Aberystwyth, Wales.
Burrows, G. E., and R. J. Tyrl. 2001. Toxic Plants of North America. Iowa State University Press, Ames, Iowa.
Döpp, W. 1950. Eine der Antheridienbildung bei Farnen fördernde Substanz in den Prothallien von Pteridium aquilinum (L.) Kuhn. Ber. Deutsch. Bot. Ges. 63: 139-147.
Evans, I. A. 1976. Relationship between bracken and cancer. Bot. J. Linn. Soc. 73: 105-112.
Evans, I. A. 1986. The carcinogenic, mutagenic, and teratogenic toxicity of bracken. Pp. 139-146 in R. T. Smith and J. A. Taylor, eds. Bracken: Ecology, Land Use, and Control Technology. Parthenon Publishing, Carnforth, England.
Evans, W. C. 1976. Bracken thiaminase-mediated neurotoxic syndromes. Bot. J. Linn. Soc. 73: 113-131.
Evans, W. C. 1986. The acute diseases caused by bracken in animals. Pp. 121-132 in R. T. Smith and J. A. Taylor, eds. Bracken: Ecology, Land Use, and Control Technology. Parthenon Publishing, Carnforth, England.
Farlow, W. G. 1889. Apospory in Pteris aquilina. Ann Bot. (Oxford) 21: 360-368.
Ferguson, D. E. and R. J. Boyd. 1988. Bracken fern inhibition of conifer regeneration in northern Idaho. Res. Pap. I.N.T. U.S. Forest Serv. INT-388: i, 1-11.
Fowler, S. V. 2002. The potential for control of bracken in the UK using introduced herbivorous insects. Pesticide Sci. 37: 393-397.
Gaskin, R. E., N. A. Davenhill and J. A. Zabkiewicz. 1986. Preliminary screening of asulam and glyphosate-surfactant formulations for control of bracken (Pteridium esculentum). Pp. 413-420 in R. T. Smith and J. A. Taylor, eds. Bracken: Ecology, Land Use, and Control Technology. Parthenon Publishing, Carnforth, England.
Gliessman, S. R. 1976. Allelopathy in a broad spectrum of environments as illustrated by bracken. Bot. J. Linn. Soc. 73: 95-104.
Hannam, D. A. R. 1986. Bracken poisoning in farm animals with special reference to the North York moors. Pp. 133-138 in R. T. Smith and J. A. Taylor, eds. Bracken: Ecology, Land Use, and Control Technology. Parthenon Publishing, Carnforth, England.
Hirono, I. 1990. Carcinogenicity of bracken fern and its causal principle. Pp. 233-240 in J. A. Thomson and R. T. Smith, eds. Bracken Biology and Management. Australian Institute of Agricultural Science Occasional Papers 40, Sydney.
Hodge, W. H. 1973. Fern foods of Japan and the problem of toxicity. Amer. Fern J. 63: 77-80.
Hopkins, A. 1995. Factors influencing cattle bracken-poisoning in Great Britain. Chapter 25, pp. 120-123 in R. T. Smith and J. A. Taylor, eds. Bracken: An Environmental Issue. International Bracken Group Special Publication, vol. 2, Aberystwyth, Wales.
Hudson, P. J. 1986. Bracken and ticks on grouse moors in the north of England. Pp. 161-170 in R. T. Smith and J. A. Taylor, eds. Bracken: Ecology, Land Use, and Control Technology. Parthenon Publishing, Carnforth, England.
Jacobs, C. A. and J. H. Peck. 1993. Pteridium Gleditsch ex Scopoli. Pp. 201-204 in Flora of North America Editorial Committee, eds. 1993. Flora of North America North of Mexico. Volume 2. Pteridophytes and Gymnosperms. Oxford University Press, New York.
Jarrett, F. M., I. Manton and S. K. Roy. 1868. Cytological and taxonomic notes on a small collection of living ferns from the Galapagos. Kew Bull. 22: 475-480.
Jubrael, J. M. S., E. Sheffield and D. Moore. 1986. Polymorphisms in DNA of Pteridium aquilinum. Pp. 309-313 in R. T. Smith and J. A. Taylor, eds. Bracken: Ecology, Land Use, and Control Technology. Parthenon Publishing, Carnforth, England.
Jubrael, J. M. S., E. Sheffield and D. Moore. 1990. Detection of genetically useful variants in Pteridium. Pp. 79-86 in J. A. Thomson and R. T. Smith, eds. Bracken Biology and Management. Australian Institute of Agricultural Science Occasional Papers 40, Sydney.
Klekowski, E. J., Jr. 1972. Evidence against genetic self-incompatibility in the homosporous fern Pteridium aquilinum. Evolution 26: 66-73.
Kramer, K. U. 1990. Dennstaedtiaceae. Pp. 81-94 in K. Kubitzki, ed. The Families and Genera of Vascular Plants. Volume 1. Pteridophytes and Gymnosperms. Volume eds. K. U. Kramer and P. S. Green. Springer-Verlag, Berlin.
Kuhn, M. 1879. III. Cryptogamae vasculares. Vol. 3, fasc. 3, Pp. 7-71 in C. C. von der Decken. 1869-1879. Baron Carl Claus von der Decken’s Reisen in Ost-Afrika in den Jahren 1859 bis 1861, 4 vols in 6 pts. C. F. Winter’she Verlagshandlung, Leibzig.
Linnaeus, C. 1753. Species Plantarum : Exhibentes Plantas Rite Cognitas, Ad Genera Relatas, cum Differentiis Specificis, Nominibus Trivialibus, Synonymis Selectis, Locis Natalibus, Secundum Systema Sexuale Digestas, vol. 2. Impensis Laurentii Salvii, Stockholm.
Löve, Á. and Kjellqvist. 1972. Cytotaxonomy of Spanish plants. I. Introduction, Pteridophyta and Gymnospermae. Lagascalia 2: 23-35.
Löve, Á., D. Löve, and R. E. G. Pichi Sermolli. 1977. Cytotaxonomical Atlas of the Pteridophyta. J. Cramer, Vaduz, Lichtenstein.
Mickel, J. T. and A. R. Smith. 2004. The pteridophytes of Mexico. Mem. New York Bot. Gard. 88: i-xiii, 1-1055.
Moerman, D. E. 1998. Native American Ethnobotany. Timber Press, Portland, Oregon.
Moran, R. C. 2004. A Natural History of Ferns. Timber Press, Portland, Oregon.
Oliver, W. R. B. 1928. The Flora of the Waipaoa Series (Later Pliocene) of New Zealand. Trans. & Proc, Roy. Soc. New Zealand 59: 287-303.
den Ouden, J. 1995. Allelopathy in bracken in The Netherlands. Chapter 8, pp. 43-48 in R. T. Smith and J. A. Taylor, eds. Bracken: An Environmental Issue. International Bracken Group Special Publication, vol. 2, Aberystwyth, Wales.
Petrov, P. and R. H. Marrs. 2000. Follow-up methods for bracken control following an initial glyphosate application: the use of weed wiping, cutting and reseeding. P. 185 in J. A. Taylor and R. T. Smith, eds. Bracken Fern: Toxicity, Biology and Control. International Bracken Group Special Publication, vol. 4, Aberystwyth, Wales.
Potter, D. M. and R. M. Pitman. 1995. The extraction and characterization of carcinogens from bracken and the effect of composting. Chapter 23, pp. 110-115 in R. T. Smith and J. A. Taylor, eds. Bracken: An Environmental Issue. International Bracken Group Special Publication, vol. 2, Aberystwyth, Wales.
Raghavan, V. 1969. Photocontrol of growth pattern in a tissue isolated from the gametophytes of bracken fern. Pl. Cell Physiol. 10: 481-484.
Raghavan, V. 1970. Germination of bracken spores. Exp. Cell Res. 63: 341-352.
Robinson, R. C. 2000. The means to large-scale, effective bracken clearance in the UK. Chapter 25, pp. 156-162 in J. A. Taylor and R. T. Smith, eds. Bracken Fern: Toxicity, Biology and Control. International Bracken Group Special Publication, vol. 4, Aberystwyth, Wales.
Robinson, R. A. and C. N. Page. 2000. Protection of non-target ferns during extensive spraying of bracken. Chapter 26, pp. 163-174 in J. A. Taylor and R. T. Smith, eds. Bracken Fern: Toxicity, Biology and Control. International Bracken Group Special Publication, vol. 4, Aberystwyth, Wales.
Rumsey, F. J., E. Sheffield and C. H. Haufler. 1991. A re-assessment of Pteridium aquilinum (L.) Kuhn in Britain. Watsonia 18: 297-301.
Rymer, L. 1976. The history and ethnobotany of bracken. Bot. J. Linn. Soc. 73: 1651-176.
Scopoli, J. A. 1760. Flora Carniolica Exhibens Plautas Carniolae Indigenas et Distributas in Classes Naturales cum Differentiis Specificis, Synonymis Recentiorum, Locis Natalibus, Nomimbus Incolarum. Obscrvationibus Selectis, Viribus Medicis, ed. 1. Joannis Thomae Trattner, Vienna.
Sharma, S. D. and R. C. Kirkwood. 1995. The role of non-ionic surfactants as adjuvants for use with systemic herbicides on bracken. Chapter 27, pp. 130-136 in R. T. Smith and J. A. Taylor, eds. Bracken: An Environmental Issue. International Bracken Group Special Publication, vol. 2, Aberystwyth, Wales.
Sheffield, E., P. G. Wolf, C. H. Haufler and J. M. S. Jubrael. 1986. Pteridium herediae: a dangerous diploid bracken? Pp. 301-307 in R. T. Smith and J. A. Taylor, eds. Bracken: Ecology, Land Use, and Control Technology. Parthenon Publishing, Carnforth, England.
Sheffield, E., P. G. Wolf, C. H. Haufler, T. A. Ranker and A. C. Jermy. 1989. A re-evaluation of plants referred to as Pteridium herediae (Colmeiro) Löve and Kjellqvist. Bot. J. Linn. Soc. 99: 377-386.
Sheffield, E., P. G. Wolf, and C. H. Haufler. 1989. How big is a bracken plant? Weed Res. 29: 455-460.
Sheffield, E., P. G. Wolf, F. J. Rumsey, D. J. Robson, T. A. Ranker and S. M. Challinor. 1993. Spatial distribution and reproductive behaviour of a triploid bracken (Pteridium aquilinum) clone in Britain. Ann Bot. (Oxford) 72: 231-237.
Sheffield, E., P. G. Wolf and T. A. Ranker. 1995. Genetic analysis of bracken in the Hawaiian Islands. Chapter 5, pp. 29-32 in R. T. Smith and J. A. Taylor, eds. Bracken: An Environmental Issue. International Bracken Group Special Publication, vol. 2, Aberystwyth, Wales.
Simán, S. E., A. C. Povey, P. J. O’Connor, T. H. Ward, G. P. Margison and E. Sheffield. 2000. The genotoxicity of fern spores. Chapter 15, pp. 99-105 in J. A. Taylor and R. T. Smith, eds. Bracken Fern: Toxicity, Biology and Control. International Bracken Group Special Publication, vol. 4, Aberystwyth, Wales.
Smith, B. L., D. R. Lauren and A. S. Prakash. 2000. Bracken fern (Pteridium) toxicity in animal and human health. Chapter 11, pp. 76-85 in J. A. Taylor and R. T. Smith, eds. Bracken Fern: Toxicity, Biology and Control. International Bracken Group Special Publication, vol. 4, Aberystwyth, Wales.
Sobota, A. E. and C. R. Partanen. 1967. The growth and division of cells in relation to morphogenesis in fern gametophytes. II. The effect of biochemical agents on the growth and development of Pteridium aquilinum. Canad. J. Bot. 45: 595-603.
Speer, W. 2000. A systematic assessment of British and North American Pteridium using cpDNA gene sequences. Chapter 5, pp. 37-42 in J. A. Taylor and R. T. Smith, eds. Bracken Fern: Toxicity, Biology and Control. International Bracken Group Special Publication, vol. 4, Aberystwyth, Wales.
Speer, W. D., C. R. Werth and K. W. Hilu. 1999. Relationships between two infraspecific taxa of Pteridium aquilinum (Dennstaedtiaceae). II. Isozyme evidence. Syst. Bot. 23: 313-325.
Tan, M. K. and J. A. Thomson. 1990. Variation of genome size in Pteridium. Pp. 87-93 in J. A. Thomson and R. T. Smith, eds. Bracken Biology and Management. Australian Institute of Agricultural Science Occasional Papers 40, Sydney.
Tan, M. K. and J. A. Thomson. 1990. Evolutionary studies of the chloroplast genome in Pteridium. Pp. 95-103 in J. A. Thomson and R. T. Smith, eds. Bracken Biology and Management. Australian Institute of Agricultural Science Occasional Papers 40, Sydney.
Thomson, J. A. 2000. New perspectives on taxonomic relationships in Pteridium. Chapter 3, pp. 15-34 in J. A. Taylor and R. T. Smith, eds. Bracken Fern: Toxicity, Biology and Control. International Bracken Group Special Publication, vol. 4, Aberystwyth, Wales.
Thomson, J. A. 2000. Morphological and genomic diversity in the genus Pteridium (Dennstaedtiaceae). Ann. Bot. (Oxford) 85 (suppl. B): 77-99.
Thomson, J. A. 2004. Towards a taxonomic revision of Pteridium (Dennstaedtiaceae). Telopea 10: 793-803.
Thomson, J. A. and M. E. Alonso-Amelot. 2002. Clarification of the taxonomic status and relationships of Pteridium caudatum (Dennstaedtiaceae) in Central and South America. Bot. J. Linn. Soc. 140: 237-248.
Thomson, J. A., P. H. Weston and M. K. Tan. 1995. A molecular approach to tracing major lineages in Pteridium. Chapter 4, pp. 21-28 in R. T. Smith and J. A. Taylor, eds. Bracken: An Environmental Issue. International Bracken Group Special Publication, vol. 2, Aberystwyth, Wales.
Thomson, J. A., P. H. Weston and M. K. Tan. 2000. A molecular approach to tracing major lineages in Pteridium: update and amendment. Chapter 4, pp. 35-36 in J. A. Taylor and R. T. Smith, eds. Bracken Fern: Toxicity, Biology and Control. International Bracken Group Special Publication, vol. 4, Aberystwyth, Wales.
Thomson, J. A., A. C. Chikuni and C. S. McMaster. 2005. The taxonomic status and relationships of bracken ferns (Pteridium: Dennstaedtiaceae) from sub-Saharan Africa. Bot. J. Linn. Soc. 148: 311-321.
Tryon, A. F. and B. Lugardon. 1991. Spores of the Pteridophyta. Springer-Verlag, New York.
Tryon, R. M., Jr. 1941. A revision of the genus Pteridium. Rhodora 43: 1-31, 37-67.
Tryon, R. M. and A. F. Tryon. 1982. Ferns and Allied Plants, with Special Reference to Tropical America. Springer-Verlag, New York.
Villalobos-Salazar, J., A. Meneses and J. Salas. 1990. Carcinogenic effects in mice of milk from cows fed on bracken fern Pteridium aquilinum. Pp. 247-251 in J. A. Thomson and R. T. Smith, eds. Bracken Biology and Management. Australian Institute of Agricultural Science Occasional Papers 40, Sydney.
Villalobos-Salazar, J., J. Mora, A. Meneses, and B. Pashov. 1995. The carcinogenic effects of bracken spores. Chapter 21, pp. 102-103 in R. T. Smith and J. A. Taylor, eds. Bracken: An Environmental Issue. International Bracken Group Special Publication, vol. 2, Aberystwyth, Wales.
Whittier, D. P. 1966. Natural apospory in Pteridium. Amer. Fern J. 56: 61-64.
Whittier, D. P. 1966. Induced apogamy in diploid gametophytes of Pteridium. Canad. J. Bot. 44: 1717-1721.
Whittier, D. P. 1966. The influence of growth substances on the induction of apogamy in Pteridium gametophytes. Amer. J. Bot. 53: 882-886.
Wolf, P. G., C. H. Haufler and E. Sheffield. 1988. Electrophoretic variation and mating system of the clonal weed Pteridium aquilinum (L. Kuhn) (bracken). Evolution 42: 1350-1355.
Wolf, P. G., E. Sheffield and C. H. Haufler. 1991. Estimates of gene flow, genetic substructure and population heterogeneity in bracken (Pteridium aquilinum). Biol. J. Linn. Soc. 42: 407-423.
Wolf, P. G., E. Sheffield, J. A. Thomson and R. B. Sinclair. 1995. Bracken taxa in Britain: a molecular analysis. Chapter 3, pp. 16-20 in R. T. Smith and J. A. Taylor, eds. Bracken: An Environmental Issue. International Bracken Group Special Publication, vol. 2, Aberystwyth, Wales.
Womack, J. G., G. M. Eccleston and M. N. Burge. 1995. Progress in the development of a mycoherbicidal formulation for bracken control. Chapter 28, pp. 137-140 in R. T. Smith and J. A. Taylor, eds. Bracken: An Environmental Issue. International Bracken Group Special Publication, vol. 2, Aberystwyth, Wales.
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Rights holder/Author | George Yatskievych |
Source | No source database. |
The mostly northern hemisphere distribution of Pteridium aquilinum sensu stricto vs. the mostly southern hemisphere distribution of the remaining diploid species in the genus is suggestive of a Laurasian/Gondwanan split, however, there is no fossil evidence to directly confirm the development this biogeographic pattern. Currently, there is a large area of geographic overlap between the two groups in Latin America and portions of southern Asia. Rymer (1973) reviewed the fossil history of Pteridium. The oldest macrofossils attributed to the genus date to the Oligocene of Hungary, and unequivocal leaf compressions of Pteridium have been recorded from the late Miocene of England and late Pliocene of New Zealand (Oliver, 1928). Rymer reviewed the Quaternary history of Pteridium in Great Britain in somewhat more detail, noting that its distribution waxed and waned during successive glacial and interglacial stages, and that generally the current distribution of P. aquilinum is a relatively recent phenomenon affected greatly by prehistoric and historic human-mediated perturbations such as clearing of forests and introduction of livestock.
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Source | No source database. |
Medium- to large-sized ferns with a usually extensive system of long-creeping, branched, subterranean, hairy rhizomes, often forming large clonal colonies. Leaves mostly 0.5–1.5 m long, rarely to 2.5 m long, glabrous or variously hairy, long-petiolate, the petioles grooved along the upper side. Leaf blades 2 to 3 times pinnately compound and lobed, broadly deltate to deltate-triangular, ovate-triangular, or nearly pentagonal, the ultimate lobes or segments rounded to pointed at the tip, with strongly recurved, often pale margins, these irregular and usually sparsely to densely fringed along the edge, acting as a pseudoindusium to cover the developing sporangia (a true, outward-oriented indusium absent or vestigial). Sporangia in a continuous band along the leaf margins, hidden by the pseudoindusium until maturity. Spores 64 per sporangium, brown, nearly spherical (slightly tetrahedral), with a 3-branched tetrad scar (trilete), 23–40 μm in diameter, the outer surface irregularly granular. Source documents: Tryon (1941), Tryon and Tryon (1982), Kramer (1990), Tryon and Lugardon (1991), Jacobs and Peck (1993), Mickel and Smith (2004).
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Rights holder/Author | George Yatskievych |
Source | No source database. |
Although the taxonomy of the genus Pteridium is still controversial, four or five species are often accepted in modern accounts, including: P. aquilinum (L.) Kuhn sensu stricto, P. arachnoideum (Kaulf.) Maxon, P. caudatum (L.) Maxon, P. esculentum (G. Forst.) Cockayne, and P. semihastatum (Wall. ex J. Agardh) S.B. Andrews (P. yarrabense (Domin) N.A Wakef.) (Thomson, 2004).
The list of epithets that have been applied as infraspecific taxa within P. aquilinum when the genus was still considered monospecific includes nearly 20 names. A number of these are now applied to segregate species within the genus and the taxonomic status of most others still has not been fully resolved.
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Rights holder/Author | George Yatskievych |
Source | No source database. |