History of phycology

The history of phycology is the history of the scientific study of algae. Human interest in plants as food goes back into the origins of the species, and knowledge of algae can be traced back more than two thousand years. However, only in the last three hundred years has that knowledge evolved into a rapidly developing science.

Early days

The study of botany goes back into pre-history as plants were the food of people from the beginning of the human race. The first attempts at plant cultivation are believed to have been made shortly before 10,000 BC in Western Asia (Morton, 1981)Morton, A.G. 1981 ''History of Botanical Science.'' Academic Press Inc. (London) Ltd. and the first references to algae are to be found in early Chinese literature. Records as far back as 3000 BC indicate that algae were used by the emperor of China as food (Huisman, 2000 p. 13).Huisman, J.M. 2000. ''Marine Plants of Australia.'' University of Western Australia Press, Australia. The use of ''Porphyra'' in China dates back to at least A.D. 533–44 (Mumfard and Miura, 1988),Mumford, T.F. and Miura, A. 1988. 4. p.87–117. ''Porphyra'' as food: cultivation and economics. in Lembi, C.A. and Waaland, J.R. 1988. ''Algae and Human Affairs.'' Cambridge University Press, Cambridge there are also references in Roman and Greek literature. The Greek word for algae was "Phycos" whilst in Roman times the name became '' Fucus''. There are early references to the use of algae for manure. The first coralline algae to be recognized as living organisms were probably ''Corallina'', by Pliny the Elder in the 1st century AD (Irvine and Chamberlain, 1994 p. 11).

The classification of plants suffered many changes since Theophrastus (372–287 B.C.) and Aristotle (384–322 B.C.) grouped them as "trees", "shrubs" and "herbs" (Smith, 1955 p. 1).Smith, G.M. 1955. ''Cryptogamic Botany. Volume 1. Algae and Fungi.'' McGraw-Hill Book Company, New York.,

Little is known of botany during the Middle Ages — it was the Dark Ages of botany.

The development of the study of phycology runs in a pattern comparable with, and parallel to, other biological fields but at a different rate. After the invention of the printing-press in the 15th century (with the publication of the first printed book: Gutenberg's ''Bible'' of 1488) Hawksworth, D.L and Seaward, M.R.D. 1977. ''Lichenology in the British Isles 1568 - 1975'' The Richmond Publishing Co. Ltd. education enabled people to read and knowledge to spread.

Exploration of the world and the advance of knowledge

Written accounts of the algae of South Africa were made by the Portuguese explorers of the 15th and 16th centuries, however it is not clear to which species reference was being made (Huisman, 2000 p. 7).

17th Century

In the 17th Century there was a great awakening of scientific interest all over Europe, and after the invention of the printing-press books on botany were published. Among them was the work of John Ray

who wrote in 1660: ''Catalogus Plantarum circa Cantabrigiam.'', this initiated a new era in the study of Botany (Smith, 1975 p. 4).Smith, A.L. 1975. ''Lichens''. The Richmond Publishing Co. Ltd. England Ray "influenced both the theory and the practice of botany more decisively than any other single person in the latter half of the seventeenth century" (Morton, 1981).

However no real progress was made in the scientific study of algae until the invention of the microscope — in about 1600. It was Anton van Leeuwenhoek (1632–1723) who discovered bacteria and saw the cell structure of plants. His unsystematic glimpses of plant structure, reported to the Royal Society between 1678 and his death in 1723, produced no significant advances (Morton, 1981 p. 180).

As adventurers explored the world more species of all animals and plants were discovered, this demanded efforts to bring order out of this quickly accumulating knowledge.

The first Australian marine plant recorded in print was collected from Shark Bay on the Western Australian coast by William Dampier who described many new species of Australian wildlife in the 17th century (Huisman, 2000 p. 7).

18th century

Before Carl von Linné (1707–1778) animals and plants had names, but it took him to arrange the names and group the plants of this Earth in some sort of order. Carl Linnaeus (Carl von Linné

was a Swedish botanist, the son of a pastor of the Lutheran church, a physician and zoologist. He laid the foundations of modern biological systematics and nomenclature in his ''Species Plantarum'' (1753).Linnaeus, C. 1753 ''Species plantarum...,'' 2 vols. Salvius, Stackholm. He adopted and popularized a binomial (or binary) system of designation (Morton, 1981) using one name as the genus and a second name as the species name both in Latin or ''Latinised''. This specific name he
referred to as a trivial name ''nomen triviale'' consisting of a single word, normally a Latin adjective, but any single word would suffice, to identify a particular species, but not intended to describe it. He developed a coherent system for naming organisms and divided the plant kingdom into 25 classes (according to Smith p. 1 and p. 24 according to Dixon, 1973) (Smith, 1955 p. 1),Dixon, P.S. 1973. ''Biology of the Rhodophyta.'' Oliver and Boyd, Edinburgh. one of which, the ''Cryptogamia'', included all plants with ''concealed'' reproductive organs. He divided the Cryptogamia into four orders: Filices (ferns), Musci (mosses), Algae (including lichens), and fungi (Smith, 1955 p. 1).

Examination for the reproductive structures had already started. In 1711, R.A.F de Réaumur gave an account of '' Fucus'' in which noted the two types of external openings in the thallus: the non-sexual cryptostromata (sterile surface cavities) and the conceptacles (fertile cavities, immersed but with a surface opening) containing the sexual organs, which he thought were female flowers. With a lens he was able to see the oogonoa (the female sex organs) and the antheridia (the male sex organs) within the conceptacles, but he interpreted these as seeds (Morton, 1981 p. 245). Johann Hedwig (1730–1799) provided further evidence of the sexual process in algae, and figured conjugation in ''Spirogyra'' Hedwig in 1797. He also illustrated '' Chara'' ('' Charales'') and identified the antheridia and oogonia as male and female sexual organs (Morton, 1981 p. 323 & 357).

Harvey commented on ''...motion, apparently spontaneous, among the seeds at the period of germination.'' Some found it ''difficult...to account for these anomalous motions.'' ''...that the seeds becomes (how is not said) a perfect animalcule, which after enjoying an animal existence for a time ceases to live animally, and, reverting to its original nature, gives birth to a vegetable. Thus, this seed was first vegetable, then animal, and then again vegetable,...'' .Harvey, W.H. 1841. ''A Manual of the British Algae:'' London: John van Voorst During the 18th Century there was a stormy controversy as to whether coralline algae were plants or animals. Up to the mid-18th century coralline algae (and coral animals) were generally treated as plants. By 1768 many, but by no means all authorities, considered them animal. Five years later, Harvey concluded that they were certainly of vegetable material he noted: "The question of the vegetable nature of Corallines, among which the ''Melobesia'' take rank, may now be considered as finally set at rest, by the researches of Kützing, Phillipi and Decaisne." (Harvey, 1847, pl. 73).Harvey, W.H. 1847. ''Phycologia Britannica.'' Vol. 1, Fasc.13 (plates 73–78) Reeve & Benham, London, London.Woelkerling, Wm. J. 1988 ''The Coralline Red Algae:...'' British Museum (Natural History), Oxford University Press.

The first scientific species description of a South African seaweed accepted for most nomenclatural purposes is that of ''Ecklonia maxima'', published in 1757 as ''Fucus maximus'' (Stegenga ''et al.'', 1997).Stegenga, H., Bolton, J.J. and Anderson, R.J. 1997. ''Seaweeds of the South African West Coast.'' Bolus Herbarium, University of Cape Town.

Knowledge of North American Pacific algae begins with the 1791–95 expedition of Captain George Vancouver (Papenfuss, 1976 p. 21).Papenfuss, G.F. pp.21–46 Landmarks in Pacific North American Marine Phycology in Abbott, I.A. and Hollenberg, G.J. 1976. ''Marine Algae of California.'' Stanford University Press, California.

Archibald Menzies (1754–1842) was the appointed botanist on the expedition led by Captain George Vancouver in the ships ''Discovery'' and ''Chatham'' of 1791–1795 to the Pacific coast of North America and south-western Australia. The algae collected by Menzies were passed to Dawson Turner (1775–1858) who described and illustrated them in a four-volumed work published in 1808–1819. However Turner only referred to the taxa referable to '' Fucus''; either Menzies collected very few or he gave only a few to Turner. Three of these species described by Turner later became the types of new genera (Papenfuss, 1976) and (Huisman, 2000) Turner also received plants from Robert Brown (1773–1858) the botanist who accompanied Captain Matthew Flinders on the ''Investigator'' (1801–1805). This collection also included many plants from Australia (Huisman, 2000).

The real awakening of interest in American algae resulted from a visit by William Henry Harvey in 1849–1850 when he visited areas from Florida to Nova Scotia and produced three volumes of ''Nereis Boreali-Americana.'' These gave an incentive to others to study algae (Taylor, 1972 p. 21).Taylor,W.R. 1972 ''Marine Algae of the Northeastern Coast of North America.'' Ann Arbor, Mich

The first collector of marine algae in Greenland waters seems to have been J.M.Vahl who lived in Greenland from 1828 to 1836. Vahl's East Greenland species were not recorded until 1893 when Rosenvinge included them in his work of 1893 together with the species collected by Sylow (Lund, 1959).Lund,S. 1959. The Marine Algae of East Greenland I. Taxonomic part. ''Meddelelser om Grønland'' 156: 1–248 F.R.Kjellman records only 12 species from East Greenland 4 of which are doubtful, these records are based on Zeller's list (Lund, 1959).

Early 19th Century

Carl Adolph Agardh was one of the most prominent algologists of all time, he was born in Sweden on 23 January 1785 and died on 28 January 1859. He was Professor of Botany at the University of Lund and later Bishop of Karlstad Diocese (Papenfuss, 1976). Many species still show his name as the authority of the scientific name. He traveled widely in Europe visiting Germany, Poland, Denmark, the Netherlands, Belgium, France and Italy and was the first to emphasize the importance of the reproductive characters of algae and use them to distinguish the different genera and families. His son, Jacob Georg Agardh (1813–1901), who became Professor of Botany at Lund in 1839, made a study of the life-histories of algae, described many new genera and species. It was to him that many workers sent specimens for determination and as donations. Because of this the herbarium at Lund is the most important algal herbarium in the world (Papenfuss, 1976).

The first records of algae from the Faroe Islands were made by Jørgen Landt in his book of 1800 where he mentions about 30 species. Following this, Hans Christian Lyngbye visited the Faroe Islands in 1817 and published his work in 1819. In this, he described several new genera and species, some 100 new species were listed. Emil Rostrup who visited the Faroe Islands in 1867 listed ten new species and a total not far from 100. In 1895, ''Herman G. Simmons'' mentioned 125 species. In that year F. Børgesen (1866–1956) started work and in 1902 published his work (Børgesen, 1902).'' Børgesen, F.'' 1902. Marine algæ (of the Færöes). Botany of the Færöes (ed. E. Warming), Vol. II, pp. 339-532.'' Part II. Copenhagen

Jean Vincent Félix Lamouroux (1779–1825) was the first, in 1813, to separate the algae into groups on the basis of colour (Dixon and Irvine, 1977 p. 59).Dixon, P.S. and Irvine, L.M. 1977. ''Seaweeds of the British Isles. Vol.1. Rhodophyta. Part 1.'' British Museum atural History London. At this time all coralline algae were considered animals, it was R. Philippi who in 1837 published his paper in which he finally recognized that coralline algae were not animals and he proposed the generic names '' Lithophyllum'' and '' Lithothamnion'' (Irvine and Chamberlain, 1994 p. 11).Irvine, L.M. and Chamberlain, Y.M. 1994. ''Seaweeds of the British Isles Vol.1. Part 2B.'' Natural History Museum, London.

Freshwater algae are commonly treated separately from marine algae and may be considered not correctly placed in phycology. Lewis Weston Dillwyn (1778–1855) "British Confervae" (1809) was one of the earliest attempts to bring together all that was then known on the British Freshwater algae .West, G.S. and Fritsch, F.E. 1927. ''A Treastise on the British Freshwater Algae.'' Cambridge University Press, Cambridge

Specimens of Anne E. Ball (1808–1872) have been found in both the Herbarium of the Irish National Botanic Gardens, Dubli

and the Ulster Museum (BEL). A.E.Ball was an Irish algologist who corresponded with W.H. Harvey and whose records appear in his ''Phycologia Britannica.'' The specimens in Dublin do not contain any unusual or rare items. However, they are well documented.Parkes, H.M. and Scannell, M.J.P. 1970. Anne E. Ball, two volumes of algae in Herbarium. National Botanic Gardens, Dublin. ''Irish Naturalists' Journal'' 16: 349

W.H. Harvey

Willian Henry Harvey (1811–1866), Keeper of the Herbarium and professor in botany at Trinity College, Dublin, was one of the most distinguished algologists of his time (Papenfuss, 1976 p. 26). Apart from Ireland he visited South Africa, the Atlantic seaboard of America as far south as the Florida Keys on the east coast of North America and Australia (1854–1856). Between 1853 and 1856 he visited Ceylon, Australia and New Zealand and various parts of the South Pacific (Huisman, 2000 & Papenfuss, 1976). His collection in Australia resulted in one of the most extensive collections of marine plants and it inspired others (Huisman, 2000). He published: ''Nereis Australis Or Algae of the Southern Ocean'' in 1847–1849 and in 1846–51 his ''Phycologia Britannica'' appeared. His ''Nereis Boreali-Americana'' was published in three parts (1852–1858) this was the first, and still is (1976) is the only marine algal flora of North America as it includes taxa from the Pacific coast (Papenfuss, 1976 p. 27). His five-volume '' Phycologia Australica'' was published in 1858 to 1863. These volumes remain to this day a most important reference to Australian algae (Huisman, 2000). His primary herbarium is in Trinity College, Dublin (TCD). However large collections of Harvey material are to be found in the Ulster Museum (BEL) (Morton, 1977; Morton, 1981);Morton, O. 1977. A note on W.H.Harvey's algae in the Ulster Museum. ''Irish Naturalists' Journal.'' 19:26Morton, O. 1981. American algae collected by W.H.Harvey and others, in the Ulster Museum Herbarium. ''Taxon'': 30:867–868 University of St Andrews (STA) and National Herbarium of Victoria (MEL), Melbourne, Australia (May, 1977).May, V. 1977 Harvey's Australian Algae at the National Herbarium of New South Wales (NSW), Sydney, Australia. ''Taxon'': 26: 496 Many of the collectors of this period sent, and exchanged, specimens freely one to another, as a result Harvey's books show a remarkable knowledge of the distribution of algae elsewhere in the world. His ''Phycologia Britannica'' lists species recorded and collected from various parts of the British Isles. For example, he notes William Thompson (1805–1852), William McCalla (c.1814–1849), John Templeton (1766–1825) and D. Landsborough (1779–1854) who collected, as he did, from distinct sites in Ireland. The collections of these botanists, and many others, are represented separately by collections in the Ulster Museum (BEL).

Sir William Jackson Hooker (1785–1865) was a lifelong friend of Harvey (Papenfuss, 1976 p. 26), he was appointed Professor of Botany at Glasgow University in 1820 and became director in Kew 1841–1865. Hooker recognized the talent in Harvey and lent him books, encouraged and invited him to write the section on algae in his ''British Flora.'' as well as the section on algae for ''The Botany of Captain Beechey's Voyage'' (Papenfuss, 1976). Margaret Gatty (1809–1873) (née Margaret Scott) (author of ''British Seaweeds'', 1863), and others, corresponded with William Henry Harvey (Desmond, 1977 and Evans, 2003).Desmond,R. 1977. ''Dictionary of British and Irish Botanists and Horticulturists.'' Taylor and Francis Ltd., London Evans, F. 2003. Mrs Alfred Gatty (1809–1873), author of ''British Seaweeds'' ''The Phycologist'' No.65: 14–17

Late 19th century

Much work was done in this period by many workers and the many specimens became very valuable. Harvey's specimens, are to be found in at least several herbaria as well as those of other phycologists whose names are to be found in historic publications.
In the same period Friedrich Traugott Kützing (1807–1893) in Germany described more new genera than anyone either before or after (Chapman, 1968 p. 13).Chapman, V.J. 1968. ''The Algae.'' Mackmillan, New York His publications span the period 1841 to 1869 and added materially to knowledge of algae of cold waters of the Arctic seas. Some of his specimens are stored in the Ulster Museum Herbarium (BEL) catalogued: F1171; F10281–F10318. In 1883 Frans Reinhold Kjellman, professor of botany at Uppsala University, published ''The Algae of the Arctic Sea''. He divided the "Arctic Sea" into different regions which surround the North Pole (Kjellman, 1883).Kjellman, F.R. Reprint 1971. The Algae of the Arctic Sea.''K. Svenska VetenskAkad. Handl.'' 20(5): 1–351 Further research work on the marine algae of the world included: Charles Lewis Anderson (1827–1910) who collaborated with William Gilson Farlow and with Professor Daniel Cady Eaton to produce on the first exsiccatae of North American Algae (Papenfuss, 1976). Edward Morell Holmes (1843–1930), was an expert on seaweeds, mosses, liverworts and lichens, as well as from Norway, Sweden, Florida, Tasmania, France, Cape of Good Hope, Ceylon and Australia. He also exchanged specimens (Furley, 1989).Furley, D.D. 1989 Notes on the correspondence of W.M.Holmes (1843–1930).''The Linnean'' 5: 23–30 and some are in the herbarium of the Ulster Museum (BEL).
George Clifton (1823–1913) an Australian phycologist is mentioned in Harvey's Memoirs, as the superintendent of the Water Police in Perth, West Australia sent algal specimens to Harvey (Blackler, H.1977).Blackler, H. 1977. Harvey's Australian Algae in the Herbarium of Mrs Margaret Gatty in the Department of Botany of the University of St. Andrew's (STA), Scotland. ''Taxon'': 26: 495–496 In these years there were many workers in this field: W.G. Farlow, mentioned above, who was appointed in 1879 Professor of Cryptogamic Botany at University of Harvard (U.S.A.) in 1879 and published, among other works, the ''Marine algae of New England and Adjacent Coasts.''; in 1876 John Erhard Areschoug, a Swedish Professor of Botany at Upsalla University, reported on some brown algae collected in California by Gustavus A. Eisen (Papenfuss. 1976). George W.Traill (1836–1897) was a clerk in the Standard Life Company in Edinburgh where he worked long hours, yet he was one of the greatest authorities on Scottish algae. Despite bad health he was an indefatigable collector. In 1892 he gave his collection to the Herbarium of the Edinburgh Botanic Gardens (Furley, 1989).

Mikael Heggelund Foslie ( M.Foslie) (1855–1905) published 69 papers between 1887 and 1909. During this time he increased the number of species and forms (of corallines) from 175 to 650 (Irvine and Chamberlain, 1994). After his death his collection of specimens was purchased by the Museum of the Royal Norwegian Society for Sciences and Letters (Thor ''et al.'', 2005) Thor, E., Johansen, S and Nielsen, L.S. 2005. The collection of botanical letters to Michael H.Foslie in the Gunnerus Library: a Catalogue ''Gurreria'' 78: 7–22 and there is a small collection of his in the Ulster Museum Herbarium: (Collection No. 42) entitled: ''Algae Norvegicae'' (Ulster Museum Herbarium catalogue (BEL): F10319–F10334). F.Heydrich also described 84 taxa and was a bitter foe of Foslie. This left a legacy of complicated and still unresolved problems.

It was in the 19th Century that the true nature of lichens

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