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Peter H. Raven

From Wikipedia, the free encyclopedia
American botanist

Peter Raven
Peter Raven, after receiving theAddison Emery Verrill Medal
Born
Peter Hamilton Raven

June 13, 1936 (1936-06-13) (age 88)
Shanghai, China
NationalityAmerican
Alma materUniversity of California, Berkeley,
University of California, Los Angeles
Spouses
Children4
AwardsSee Awards and honors
Scientific career
FieldsBotany,Evolutionary biology,Biodiversity
InstitutionsStanford University,
Missouri Botanical Garden
Washington University in St. Louis
Doctoral studentsDennis E. Breedlove
Warren L. Wagner
Author abbrev. (botany)P.H.Raven

Peter Hamilton Raven (born June 13, 1936) is an Americanbotanist andenvironmentalist, notable as the longtime director, now President Emeritus, of theMissouri Botanical Garden.[1][2]

Early life

[edit]

On June 13, 1936, Raven was born inShanghai, China, to American parents, Walter Francis Raven and Isabelle Marion Breen. His father's uncle Frank Jay Raven was, for a time, one of the wealthiest Americans in China but was later jailed in a banking scandal.[3][4] That incident and Japanese aggression in China led the Raven family to return toSan Francisco, California, in the late 1930s.

After becoming a member of theCalifornia Academy of Sciences while still a youth, Raven went on to graduate with a BSc in biology from theUniversity of California, Berkeley, in 1957 and a Ph.D. in botany from theUniversity of California, Los Angeles, in 1960.

Career

[edit]

After teaching atStanford University, Raven went on to become Director of theMissouri Botanical Garden in 1971. In 2006, his position was renamed President and Director. Raven announced his plans to retire in 2011, to coincide with his 75th birthday and his 40th year at the garden.Peter Wyse Jackson was appointed as Raven's successor at the Missouri Botanical Garden in September 2010.

Raven is possibly best known for his work "Butterflies and Plants: A Study in Coevolution", published in the journalEvolution in 1964, which he coauthored withPaul R. Ehrlich. Since then he has authored numerous scientific and popular papers, many on the evening primrose family,Onagraceae. Raven is also an author of the widely used textbookBiology of Plants, now in its eighth edition, coauthored with Ray F. Evert and Susan E. Eichhorn (both ofUniversity of Wisconsin, Madison).

He is a frequent speaker on the need for biodiversity and species conservation.

In 2000, theAmerican Society of Plant Taxonomists established thePeter Raven Award in his honor to be conferred to authors with outstanding contributions to planttaxonomy and "for exceptional efforts at outreach to non-scientists".

He serves on the advisory council ofCRDF Global. He served on the board of trustees for Science Service, now known asSociety for Science & the Public, from 1993 to 1996.

Research

[edit]

Raven has published more than 700 articles, books, and monographs covering topics in Evolution, Taxonomy and Systematics, Biogeography, Coevolution, Plant Conservation, Ethnobotany, and Public Policy, including several text books.[5]

During his early years he was associated with and led Sierra Club outings for several weeks at a time, after which he published "Base Camp Reports." Published from 1950 to 1956, these reports covered a wide range of subjects, including plant lists, insects, and ecology. His first such report, at the age of 14, summarized 506 plant collections representing 337 species collected in theSierra Nevada Mountains inInyo andFresno Counties.G. Ledyard Stebbins was a counselor on this particular trip, identified by Raven as Prof. G. L. "Led" Stebbins.[6]

During this time he also published on new weed species and other plants found in and around San Francisco as well as the Sierra Nevada Mountains.[7][8][9][10][11]

In 1950 Raven, at the age of 14, had collected a plant calledC. rubicunda.[12] In the early 1950s, in the course of revising the genusClarkiaHarlan Lewis and his wife Margaret Lewis discovered the herbarium specimen collected by Raven.[12] They visited him in 1952 when he was 16, and wanted to know where the collection was made.[12] Lewis eventually located the new species, and in 1958 Lewis and Raven[13] published a botanical description of this plant, calledC. franciscana, which was morphologically very closely related toC. rubicunda andC. amoena.

Evolution

[edit]

While a graduate student at the University of California, Los Angeles, Raven andHarlan Lewis published a major paper in 1958 on the evolution ofC. franciscana, and generalized to what was by then a general a pattern of speciation inClarkia.[14] They concluded thatC. franciscana had evolved fromClarkia rubicunda; and they asserted thatC. franciscana's origin mirrored a recurring theme inClarkia of a derived species showing a close morphological similarity to a parental species, the derived species being geographically proximal, but differing from the parent by chromosomal differences and showing interspecific sterility. Further, they hypothesized that such speciation inClarkia was rapid, and perhaps occurred within the last 12,000 years.

Additionally, they hypothesized that this rapid mode of speciation seen inClarkia was analogous to a mode of speciation known asquantum evolution.[citation needed]

Following his early publication in 1958 on evolution ofC. franciscana, Raven went on to publish many papers on evolutionary topics. While at Stanford University, withPaul R. Ehrlich, he coined the termcoevolution after a 1964 review of butterflies and their food plants.[15]

In a 1969 paper Ehrlich and Raven were also critical of the idea that the definition ofspecies as advocated byErnst Mayr,Theodosius Dobzhansky, andG. Ledyard Stebbins had very little meaning for plants.[16]

In 1978 Sussman and Raven[17] advanced the idea that nonflying mammals, such as primates and marsupials, could have been significant pollinators but were outcompeted by nectar-feeding birds and bats. Any coevolved relationships between flowering plant species and non-flying mammal pollinators that persist at the present would appear to be "living fossils, which have a great deal to tell us about the evolution of both the mammals, including some of our antecedents, and of the flowering plants."

Raven wrote a review of the plant population data as of 1979, and identified several themes that he felt had potential for future research, including the above theme of the species problem.[18] He went on to assert that developmental biology would be more important in the future He advocated another theme, that being that funding should be provided for study on a few species rather than spread amongst many in order to solve population biology problems:[18]

In 1980 Raven continued discussing problems associated with defining species in plants.[19] He discussed the widespread ability of plant species to hybridize, especially in perennial plants, and the historical observations of such back to 1717. He used as examples of perennial plants in thegeneraEpilobium,Scaevola,Bidens, andCeanothus as examples of plants that appeared to use hybridization as a means to adapting to new environments. He stated "If the hybrids are particularly favored in specific ecological situations, asexual reproduction, polyploidy, or simply autogamy may favor the perpetuation of specific genotypes through a narrowing of the spectrum of genetic recombination characteristic of the population. No general conclusions about the most appropriate way to treat these populations taxonomically appear to be possible." In annual plants, using examples fromClarkia, he asserted that several species ofClarkia often occur sympatrically, yet hybrids are very rare in the wild," and that much of the sterility is due to chromosomal repatterning between species."

In 1980 Raven and coauthors reviewed the literature concerning fungal symbiosis in vascular plants.[20] They reviewed two kinds of fungal-plant associations:ectomycorrhizal andendomycorrhizal. They reported that endomycorrhizal fungi, which penetrate plant cells, are found in 80% of all vascular plants, including ferns, gymnosperms, and angiosperms, and are found in forests of high species richness. On the other hand, ectomycorrhizal fungi, which do not penetrate plant cells, occur in forests of low species richness, are usually in temperate forests, or infertile soils of the tropics. Further, they hypothesized that ectomycorrhizal forests have expanded through the MiddleCretaceous at the expense of endomycorrhizal forests.

Raven's Ph.D. thesis was on a genus within the Onagraceae, and his interest on the evolution of plants within this family as well as theMyrtales runs through his entire career.[12][5] In 1988 he published a review of the Onagraceae, covering its taxonomy, evolution,cytogenetics, anatomy, breeding systems, and geographic distribution.[21] He asserted that the family was the best known plant family of its size, and proposed that further studies of the family would be useful in understanding of "variation and evolution of plants in the future."

Biogeography

[edit]

Raven showed an early interest in plantdisjunctions prior to the wide acceptance of plate tectonic theory of the late 1960s,[22] and was an early adopter of plate tectonics in explaining plant disjunctions by the early 1970s.[23]

TheNothofagus plant genus illustrates Gondwanan distribution, having descended from the supercontinent and existing in present-day Australia, New Zealand, New Caledonia, and theSouthern Cone. Fossils have also recently been found in Antarctica.[24]

In 1963 Raven published a review ofamphitropical distributions of plant species in North and South America.[22] He divided species into three groups: biopolar or high-latitude species, temperate species, and desert species.

In 1974, withDaniel I. Axelrod, Raven published an extensive article on plant and animal biogeography in the context ofplate tectonics.[25] They stated that the new plate tectonic theory "did not require any new modifications of previously established major principles of evolution...however there were new principles of biogeography..."[citation needed]

In 1978, again with Axelrod, they published on the origin and complexity of Californian flora.[26] They reviewed that the flora of California consisted of "northern, temperate elements and xeric, southern elements, and is characterized by a high degree ofendemism." They proposed that the reasons for the large number of species in the state as well as the endemics is due to the favorable climate that has prevailed in California for most of theTertiary, as well as the recent elevation of the Sierra Nevada and other ranges, together with: "The concomitant development of a cold off-shore current which ultimately resulted in the development of a mediterranean, summer-dry climate during the past million years...The endemics of California are a mixture of relicts and newly produced species...and it is the latter that have contributed most to the size of the flora and to the high proportion of endemism in it."

Raven and Axelrod wrote a paper in 1985 on the origin of theCordilleran flora, a region bounded by the east slope of theSierra Nevada andTransverse Ranges andPeninsular Ranges of California to the eastern front of theRocky Mountains, north to theSnake River Plain-western Wyoming, and south to central Arizona-New Mexico.[27]

In 1996 Raven, Axelrod, and Al-Shehbaz wrote a paper on the history of the modern flora of China, Europe, and the continental United States.[28] They said that the three regions have approximately the same geographic area, yet China has two times the number of species as the United States, and three times as many as Europe. They asserted that all three regions had essentially the same flora as of 15 million years ago, but China came to possess the most species because of three reasons. First, China has a tropical rain forest. Second, there is an unbroken gradient of vegetation from the tropical rain forest to "boreal coniferous forests that has persisted and afforded habitats characterized by equable climates during the last 15 million years, when massive extinctions were taking place elsewhere in the Northern Hemisphere...such continuity is interrupted in North America by the Gulf of Mexico and in Europe by the Alps, the Mediterranean, and the Sahara Desert." The third reason was due to the impact of the Indian subcontinent with Asia starting 50 million years ago, making a "highly dissected, elevated geography."

Taxonomy/Systematics/Floras

[edit]

Raven has produced a wide variety of works in the area of plant systematics. Most of them are related to the plant family Onagraceae. In 1969 he published a 235-page report onCamissonia;[29] In 1976 he, along with his wife at the time Tamara Engelhorn, published a 321-page monograph onEpilobium in Australasia;[30] an embryological analysis of species in the Myrtales;[31] In 1992 a 209-page monograph on the systematics ofEpilobium in China;[32] in 1997 a 234-page monograph on the systematics ofOenothera;[33] and in 2007 a 240-page monograph on a reclassification of the Onagraceae.[34] And in 1981 he published a 1,049 page monograph on the systematics of Legumes.[35]

He has also published a number of books (Floras) devoted to the systematics of plants found in particular regions. These include a 1966 book on the native shrubs of Southern California;[36] a 1966 flora on the Santa Monica Mountains in California;[37] and, with various editors and authors, an ongoing 33 volume set (to date) on the flora of China, organized by plant family.[38][39][40][41]

Ethnobotany

[edit]

Raven (along with Dennis E. Breedlove) was a collaborator on a team led byBrent Berlin that published a seminal work on the classification of plants by theTzeltal Mayan-speaking people of HighlandChiapas.[42][43][44][45] They concluded that plant, as well as animal, descriptions could be grouped into five different hierarchical "taxonomic ethnobiological categories"; these included 1.) "unique beginner," such as plants and animals, 2.) "life form", such as tree, vine, bird, grass, mammal, etc.; 3.) the largest category, consisting usually of ~500 taxa, is "generic", and consists of names such as oak, pine, catfish, perch, and robin. Further, some "generics" were not included with in the classification of "life forms", and were called "aberrant". These included names such as cactus, bamboo, pineapple, platypus, etc., and were often of economic value—Agave, bean, and corn as examples. Two other taxa were called "specific" and "varietal," and were generally less numerous. Examples of "specific" include blue spruce, white fir, and post oak, and examples of "varietal" included baby beans, button beans, etc.

Berlin, Breedlove and Raven later extended their analysis of plants and animals to other indigenous peoples[44] including theHanunoo speaking people of the island ofMindoro,Philippines; the Karam ofPapua New Guinea; the Cantonese speakingboat people of Castle Peak Bay, Hong Kong; theNavajo of theSouthwestern United States; theFore people ofPapua New Guinea; theGuaraní people ofSouth America; and theNahuatl speaking people ofMexico, and concluded that their five to six taxonomic ethnobiological categories were generalizable.

Personal life

[edit]

From 1996 to 2000, Raven was married to Kathryn Fish, who served as the director of public policy forMonsanto. The couple met while both were serving on an environmental advisory committee for Monsanto.[46]

Awards and honors

[edit]
The standardauthor abbreviationP.H.Raven is used to indicate this person as the author whenciting abotanical name.[59]

Works

[edit]
  • Paul R. Ehrlich and Peter H. Raven (1964), "Butterflies and Plants: A Study in Coevolution",Evolution, 18: 586–608.
  • Peter H. Raven and Helena Curtis (1970),Biology of Plants, New York: Worth Publishing. [Early presentation of five-kingdom system.]

References

[edit]
  1. ^"President Emeritus".www.missouribotanicalgarden.org. Retrieved10 August 2019.
  2. ^"Royal Patrons and Honorary Fellows".The Linnean Society of London. Archived fromthe original on 14 July 2014. Retrieved25 July 2014.
  3. ^Correspondence, Special; TIMES, THE NEW YORK (2 February 1936)."CONVICTED BANKER WAS MYTH IN CHINA; F.J. Raven Rose Spectacularly in Shanghai to Become a Financial Power. BEGAN WORK AS SURVEYOR He Branched Off Into Realty, Banking and Trust Business -- Then Came Collapse".The New York Times.ISSN 0362-4331. Retrieved2 October 2021.
  4. ^"The Frank Jay Raven Story, an Ebook by Shanghai.Manholes".Smashwords. Retrieved2 October 2021.
  5. ^abRaven, Peter H."Publications of Peter H. Raven"(PDF).Discoverlife.org.
  6. ^Raven, Peter H. (1950). "Base Camp Botany".Sierra Club:1–19.
  7. ^Raven, Peter H. (1952). "Parsley for Marin County".Leafl. West. Bot.6: 204.
  8. ^Raven, Peter H. (1952). "Plant notes from San Francisco, California".Leafl. West. Bot.6:208–211.
  9. ^Raven, Peter H. (1954). "New weeds for the Sierra Nevada, California".Leafl. West. Bot.7: 151.
  10. ^Raven, Peter H. (1955). "A range extension for Allocarya cusickii in California".Leafl. West. Bot.7: 255.
  11. ^Raven, Peter H. (1956). "The grasses of San Francisco, California".Leafl. West. Bot.8:198–200.
  12. ^abcdNewbold, Heather, ed. (2000).Life Stories: World-renowned scientists reflect on their lives and the future of life on earth. Berkeley and Los Angeles, California: University of California Press. pp. 30–31.
  13. ^Lewis, Harlan; Raven, Peter (1958). "Clarkia franciscana, a new species from central California".Brittonia.10 (1):7–13.Bibcode:1958Britt..10....7L.doi:10.2307/2804688.JSTOR 2804688.S2CID 19700807.
  14. ^Lewis, Harlan; Raven, Peter H. (1958)."Rapid Evolution in Clarkia".Evolution.12 (3):319–336.doi:10.1111/j.1558-5646.1958.tb02962.x.
  15. ^Ehrlich, Paul R.; Raven, Peter (1964). "Butterflies and Plants: A study in Coevolution".Evolution.18 (4):586–608.doi:10.2307/2406212.JSTOR 2406212.
  16. ^Ehrlich, Paul R.; Raven, Peter H. (1969). "Differentiation of Populations".Science.165 (3899):1228–1232.Bibcode:1969Sci...165.1228E.doi:10.1126/science.165.3899.1228.PMID 5803535.S2CID 206567035.
  17. ^Sussman, Robert W.; Raven, Peter H. (1978). "Pollinatin by Lemurs and Marsupials: An Archaic Coevolutionary System".Science.200 (4343):731–736.Bibcode:1978Sci...200..731S.doi:10.1126/science.200.4343.731.PMID 17743224.S2CID 28858423.
  18. ^abRaven, Peter H. (1979).Future Directions in Plant Population Biology. In: Topics in Plant Population Biology. O. T. Solbrig, S. Jain, G. B. Johnson, and P. H. Raven, (eds). New York: Columbia University Press. pp. 461–481.
  19. ^Raven, Peter H. (1980). "Hybridization and the Nature of Species in Higher Plants".Canadian Botanical Association Bulletin.13 (1):3–10.
  20. ^Malloch, D. W.; Pirozynski, K. A.; Raven, P. H. (1980)."Ecological and Evolutionary Significance of mycorrhizal Symbioses in Vascular Plants (A Review)".Proc. Natl. Acad. Sci.77 (4):2113–2118.Bibcode:1980PNAS...77.2113M.doi:10.1073/pnas.77.4.2113.PMC 348662.PMID 16592806.
  21. ^Raven, Peter H. (1988).Onagraceae as a Model of Plant Evolution. In: Plant Evolutionary Biology. Leslie Gottlieb and Subodh K. Jain, eds. London, New York: Chapman and Hall. pp. 85–107.
  22. ^abRaven, Peter H. (1963). "Amphitropical Relationships in the Floras of North and South America".Quarterly Review of Biology.38 (2):151–177.doi:10.1086/403797.S2CID 85310607.
  23. ^Raven, Peter H. (1972)."Plant Species Disjunctions: A Summary".Annals of the Missouri Botanical Garden.59 (2):234–246.doi:10.2307/2394756.JSTOR 2394756.
  24. ^HaoMin, Li; ZheKun, Zhou (1 September 2007)."Fossil Nothofagaceous Leaves from the Eocene of Western Antarctica and their Bearing on the Origin, Dispersal and Systematics of Nothofagus"(PDF).Science in China Series D: Earth Sciences.50 (10):1525–1535.Bibcode:2007ScChD..50.1525H.doi:10.1007/s11430-007-0102-0.S2CID 130395392. Retrieved10 September 2017.
  25. ^Raven, Peter H.; Axelrod, Daniel I. (1974). "Angiosperm Biogeography and Past Continental Movements".Annals of the Missouri Botanical Garden.61 (3):539–673.doi:10.2307/2395021.JSTOR 2395021.
  26. ^Raven, Peter H.; Axelrod, Daniel I. (1978). "Origin and Relationships of the Californian Flora".Univ. Of California. Publ. Bot.72:1–134.
  27. ^Axelrod, Daniel I.; Raven, Peter H. (1985). "Origins of the Cordilleran Flora".Journal of Biogeography.12 (1):21–47.Bibcode:1985JBiog..12...21A.doi:10.2307/2845027.JSTOR 2845027.
  28. ^Axelrod, Daniel I.; Al-Shehbaz, Ihsan; Raven, Peter H. (1996). "History of the Modern Flora of China".Proceedings of the IFCD:43–55.
  29. ^Raven, P. H. (1969). "A revision of the genus Camissonia (Onagraceae)".Contrib. U.S. Natl. Herb.37:161–396.
  30. ^Raven, P. H.; Raven, T. E. (1976). "The Genus Epilobium (Onagraceae) in Australasia: A Systematic and Evolutionary Study".New Zealand Department of Scientific and Industrial Research Bulletin.216: 321.
  31. ^Tobe, H.; Raven, P. H. (1983)."An embryological analysis of Myrtales: its definition and characteristics"(PDF).Ann. Missouri Bot. Gard.70 (1):71–94.doi:10.2307/2399008.JSTOR 2399008.
  32. ^Chen, C. J.; Hoch, P. C.; Raven, P. H. (1992). "Systematics of Epilobium (Onagraceae) in China".Syst. Bot. Monogr.34:1–209.doi:10.2307/25027806.JSTOR 25027806.S2CID 89989636.
  33. ^Dietrich, W.; Wagner, W. L.; Raven, P. H. (1997). "Systematics of Oenothera Section Oenothera Subsection Oenothera (Onagraceae)".Syst. Bot. Monogr.50:1–234.doi:10.2307/25027870.JSTOR 25027870.
  34. ^Wagner, Warren L.; Hoch, Peter C.; Raven, Peter H. (2007). "Revised Classification of the Onagraceae".Syst. Bot. Monogr.83:1–240.
  35. ^Polhill, R. M.; Raven, P. H. (1981). "Advances in Legume Systematics".Royal Botanic Gardens, Kew.1, 2:1–1049.
  36. ^Raven, P. H. (1966).Native Shrubs of Southern California. Berkeley and Los Angeles: University of California Press. pp. 132.
  37. ^Thompson, H. H.; Prigge, B. A.; Raven, P. H. (1986).Flora of the Santa Monica Mountains, California. Los Angeles: University of California. pp. 185.
  38. ^Zheng-yi, Wu; Raven, P. H. (1994).Flora of China. Verbenaceae through Solanaceae, Vol. 17. Science Press (Beijing) and Missouri Botanical Garden (St. Louis). p. 378.
  39. ^Zheng-yi, Wu; Raven, P. H. (1994).Flora of China. Gentianaceae through Boraginaceae, Vol. 16. Science Press (Beijing) and Missouri Botanical Garden (St. Louis). p. 479.
  40. ^Zheng-yi, Wu; Raven, P. H. (1996).Flora of China. Myrsinaceae through Loganiaceae, Vol. 15. Science Press (Beijing) and Missouri Botanical Garden (St. Louis). p. 387.
  41. ^Zheng-yi, Wu; Raven, P. H. (1998).Flora of China. Scrophulariaceae through Gesneriaceae, Vol. 18. Science Press (Beijing) and Missouri Botanical Garden (St. Louis). p. 449.
  42. ^Berlin, Brent; Breedlove, Dennis E.; Raven, Peter H. (1968)."Covert Categories and Folk Taxonomies".American Anthropologist.70 (2):290–299.doi:10.1525/aa.1968.70.2.02a00050.
  43. ^Berlin, Brent; Breedlove, Dennis E.; Rave, Peter H. (1974).Principles of Tzeltal Plant Classification: An Introduction to the Bothanical Ethnography of a Mayan-Speaking People of Highland Chiapas. New York, London: Academic Press. pp. 660.ISBN 0-12-785047-3.
  44. ^abBerlin, Brent; Breedlove, Dennis E.; Raven, Peter H. (1973)."General principles of classification and nomenclature in folk biology".American Anthropologist.75:214–242.doi:10.1525/aa.1973.75.1.02a00140.
  45. ^Berlin, Brent; Breedlove, Dennis E.; Raven, Peter H. (1973)."General Principles of Classification and Nomenclature in Folk Biology".American Anthropologist.75:214–242.doi:10.1525/aa.1973.75.1.02a00140.
  46. ^Batz, Jeanne (3 November 1999)."PETER AND THE WOLF. Why Missouri Botanical Garden's Peter Raven, world-renowned environmentalist, courts Monsanto's favor, boosts its biotech and takes its money".Riverfront Times. Retrieved2 October 2015.
  47. ^Staff writer (2014)."The 2010 William L. Brown Award".William L. Brown Center for Plant Genetic Resources. Missouri Botanical Garden. Retrieved31 March 2014.
  48. ^abPeter H. Raven -- Curriculum Vita, retrieved8 September 2010
  49. ^"2014 RHS Awards for Exceptional Contributions to Horticulture Announced". Archived fromthe original on 17 March 2017. Retrieved24 December 2016.
  50. ^"Golden Plate Awardees of the American Academy of Achievement".www.achievement.org.American Academy of Achievement.
  51. ^St. Louis Walk of Fame."St. Louis Walk of Fame Inductees". stlouiswalkoffame.org. Archived fromthe original on 31 October 2012. Retrieved25 April 2013.
  52. ^"APS Member History".search.amphilsoc.org. Retrieved28 April 2022.
  53. ^"Peter H. Raven".www.nasonline.org. Retrieved28 April 2022.
  54. ^"Peter Hamilton Raven".American Academy of Arts & Sciences. Retrieved28 April 2022.
  55. ^"Royal Patrons and Honorary Fellows". Retrieved10 January 2016.[permanent dead link]
  56. ^Cockburn, Andrew (September 2015)."Weed Whackers".Harper's Magazine. Retrieved3 February 2025.
  57. ^"Megacorax S.González & W.L.Wagner | Plants of the World Online | Kew Science".Plants of the World Online. Retrieved27 May 2021.
  58. ^Weerakoon, Gothamie; Lücking, Robert; Lumbsch, H. Thorsten (2014). "Thirteen new species of Graphidaceae (lichenized Ascomycota: Ostropales) from Sri Lanka".Phytotaxa.189 (1):331–347.doi:10.11646/phytotaxa.189.1.24.
  59. ^International Plant Names Index. P.H.Raven.

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