Indian Ocean Catalogue

History of Algal Exploration in the Indian Ocean

Three stages in Australian marine phycological history were recognized by Ducker (1981). In the first stage (late eighteenth and early nineteenth centuries), a very few seaweeds, collected along with numerous other plants by European missionaries, traders, botanical explorers, and the naturalists of expeditions, were brought back or sent to the homeland of the collector. In the second stage (late nineteenth and early twentieth centuries), the collections were made by residents of the region but were still sent to Europe for determination. In the third (and present) stage, resident phycologists study and describe their own collections. While this pattern also fits the history of marine phycology on both coasts of North America, the stages are less distinct in the Indian Ocean, where, in the twentieth century, itinerant collectors and scientific expeditions have proceeded apace along with the establishment of academic centers of phycological research.

Because the International Code of Botanical Nomenclature (ICBN) (Greuter et al., 1994) specifies that botanical nomenclature begins with the first edition of Linnaeus's Species plantarum (1753), this tremendously influential work is usually taken as the starting point for historical accounts. In this work, Linnaeus arranged into 56 species the numerous marine algae that he, his contemporaries, and his predecessors had previously described and designated by polynomials. Taking an innovative step of the greatest importance in the history of biological classification, Linnaeus applied binomials to these species (as throughout the Species plantarum). He also described a new species of marine algae (from Ascension Island in the South Atlantic Ocean). These 57 species were distributed among three genera: Fucus (cartilaginous thalli), 27 species; Ulva (membranous thalli), 9 species; and Conferva (filamentous thalli), 21 species. Only two hints of an Indian Ocean provenance are found. The first is in the protologue of Fucus natans (Sargassum natans (Linnaeus) Gaillo), in which Linnaeus cited a polynomial from his Flora zeylanica (Linnaeus, 1747). This book was based on the collections of Paul Hermann (1646--1695), a professor of botany at Leiden (Netherlands), who collected in Africa, India, and Ceylon. The second hint is in the protologue of F. tendo, in which Linnaeus cited Fucus indicus teres, setam piscatoriam referens, longissimus Plukenet (1696: 160; 1692: pl. CLXXXIV: fig. 3), but he gave the provenance as China. Fucus tendo has been shown to apply to inanimate material.

In the tenth edition of his Systema naturae (1759), Linnaeus described Fucus usneoides, without provenance, but he later (second edition of Species plantarum, 1763) cited ``Oceano Indico'' for it. At that time, Linnaeus changed the name to F. granulatus. It has been shown that the provenance was erroneous (see Cystoseira usneoides (Linnaeus) Roberts).

Hermann, whose collections formed the basis of Flora zeylanica of Linnaeus, was also one of several collectors who contributed to the Flora indica of Nicolaas Laurence Burman (1768), but that work does not mention his name in connection with any of the four species of seaweeds treated there. Burman (1734--1793) was a physician and botanist in Amsterdam and the son of the celebrated Johannes Burman (1707--1779), a professor of botany at Amsterdam and a friend and correspondent of Linnaeus. The younger Burman's book is a pioneer account of tropical botany, his concept of India including the West Indies and East Indies
as well as the Indian subcontinent (Merrill, 1921). Burman described Ulva javanica, a species of uncertain status , based on a collection by Franciscus Albertus Prayan (Pryon), who was in Java during the period 1759--1764. Burman also listed the three Linnaean species, Fucus granulatus, F. natans, and F. tendo, but did not indicate if he had original material in hand.

Burman's Flora indica was published early in 1768. Later in the same year, there appeared the first book dealing exclusively with algae in which binomial nomenclature was used, namely, Gmelin's Historia fucorum. Samuel Gottlieb Gmelin (1745--1774) was a member of a celebrated family of German naturalists with Russian connections. An uncle, Johann Georg Gmelin (1709--1755), had been encouraged by Peter the Great to move to St. Petersburg, where in 1731 he was appointed professor of chemistry and natural history at the Academy of Sciences. Soon thereafter, he undertook an exploratory journey to eastern Siberia, getting as far as Yakutsk before turning back. His nephew was born in Tübingen and obtained his medical degree at Leiden in 1763 at the age of 18. He lived for a few years by the sea in Holland, where he became intrigued by seaweeds and began making observations, collections, and drawings. In 1767 he moved to St. Petersburg, where the Academy of Sciences published his Historia fucorum. Afterward, he traveled in various parts of Russia, but died from mistreatment at the hands of a hostile tribe in the Caucasus at the age of 29.

Like Linnaeus, Gmelin referred all cartilaginous algae to the genus Fucus. Unlike Linnaeus, he had a keen interest in these plants and numerous collections at hand, many of which he perceived to represent previously undescribed species. In his remarkably scholarly treatise, Gmelin synthesized all published information on seaweeds. Of the 99 species of Fucus that he recognized, 57 were newly described while 42 were adopted from Linnaeus (1753; 1759; 1763) and Hudson (1762). For eight of the latter group, he unnecessarily changed the epithet. Although in some instances Gmelin gave fairly precise collecting information, for most species the collector must be inferred. He indicated the Indian Ocean as the provenance of four newly described species but gave no hint of the collector. Of these species, F. articulatus remains of unknown identity, F. caulescens is referable to Suhria vittata (Linnaeus) Endlicher, F. clathrus is identifiable as a well-known North Pacific kelp, Thalassiophyllum clathrum (S. Gmelin) Postels & Ruprecht , and F. muricatus is treated as a taxonomic synonym of F. denticulatus Burman (Eucheuma denticulatum (Burman) Collins & Hervey ), which was originally described from the Cape of Good Hope, South Africa.

A fifth species for which Gmelin gave the Indian Ocean as the provenance is Fucus bracteatus (Gigartina bracteata (S. Gmelin) Setchell & Gardner). For this species, he cited Rumphius (1750) and Seba (1758). Since Rumphius dealt with plants from Amboina in the Pacific Ocean part of Indonesia and many of Seba's algae came from the Cape of Good Hope in the Atlantic Ocean, we may infer that Gmelin was applying the term Indian Ocean broadly and incorrectly. It is also clear that his indications of provenance are unreliable. For three newly described species, he cited ``India orientalis'', but this term refers to the Malay Archipelago rather than to India. Fucus edulis (Gracilaria edulis (S. Gmelin) P. Silva ) is based on Alga coralloides Rumphius (1750: 181, pl. LXXIV: fig. 3, pl. LXXVI: figs. A--C) from Amboina. Fucus agarum, however, is identifiable as a well-known North Pacific kelp, Agarum clathratum Dumortier , while F. laceratus is identifiable as a well-known North Atlantic red alga, Cryptopleura ramosa (Hudson) Lily Newton. Of all the species adopted from Linnaeus and Hudson, the only one with an indication of an Indian Ocean distribution is F. sargasso, an illegitimate substitute name for F. natans, for which Gmelin followed Linnaeus in citing Flora zeylanica (Linnaeus, 1747). We must conclude that despite the important role of the Historia fucorum in the marine phycology of the North Atlantic and South Africa, Gmelin almost certainly had no Indian Ocean collections in hand.

Among numerous collections of plants of all kinds that Linnaeus received from his students were those of Johann Gerhard König (1728--1785). An ethnic German born in Kurland (now part of Latvia) and residing in Denmark, König went to the Danish settlement of Tranquebar on the southeast coast of India as surgeon and naturalist in 1768 and later served as naturalist in the Madras office of the East India Company, collecting in Malabar on the southwest coast of India, Ceylon, Malacca on the Malay Peninsula, and Siam (Rendle, 1933). He bequeathed his collections and manuscripts to Sir Joseph Banks, and these are now at the British Museum (Natural History), where they have remained unpublished. During his lifetime, however, König sent collections to various European botanists and thus we find that four such collections of algae served as the basis of new species. Fucus triqueter Linnaeus (1771), a later homonym of F. triqueter S. Gmelin (1768) from the West Indies, is treated as a taxonomic synonym of Hormophysa cuneiformis (J.F. Gmelin) P. Silva in this catalogue . Linnaeus gave the provenance of F. triqueter as ``Mari Capensi'', but König, even though he stopped briefly at the Cape of Good Hope on his way to India, must have collected this alga in India or Ceylon, since in South Africa it is restricted to Natal (warm-water coast). Linnaeus's son (Linnaeus fil., 1782) described two species from Ceylon, F. flavus, which is probably a sponge, and F. pinnatus (Caulerpa pinnata C. Agardh). Many of König's collections were described by Retzius (1791), including F. zeylanicus, of unknown identity.

Robert Wight (1796--1872), a Scottish surgeon and botanist who spent 30 years in India in various capacities, including the directorship of the Botanic Garden in Madras, was the source of many new species of algae that were described by Greville (1848--1849; 1850; 1853a,b) and J. Agardh (1848b; 1848c; 1851--1863).

William Ferguson (1820--1887), a British civil servant and amateur botanist in Ceylon from 1839 until his death in Colombo, issued an informal exsiccata, Algae ceylanicae, with the specimens determined by Albert Grunow (1826--1914), an Austrian phycologist and diatomist. The first set of his specimens, which are in the British Museum (Natural History) (BM), were included in the list of Ceylon algae compiled by G. Murray (1887).

By far the most celebrated collector in the Indian Ocean was the Irish botanist William Henry Harvey (1811--1866), who provided entertaining accounts of his adventures in letters to family and friends (Ducker, 1988). Harvey stopped in Ceylon on the way to Swan Colony (Western Australia), arriving 5 September and leaving 25 December 1853. During this period he collected sufficient specimens to be distributed as an exsiccata (Harvey, 1857a). While in Australia, he published a paper describing three spectacular reticulate Delesseriaceae from Ceylon: Claudea multifida, Martensia fragilis, and Vanvoorstia spectabilis, the latter representing a new genus named for John Van Voorst, the London publisher of some of his books (including Harvey, 1841; 1849). Most of his Ceylon collections, however, were treated by Kützing and J. Agardh, as Harvey was busy pursuing other projects. His extensive collections from Western Australia, on the other hand, received immediate attention, with the manuscript of his preliminary account (Harvey, 1855b) being posted to Dublin from Melbourne in September 1854. Many of these Western Australian species were illustrated by Harvey in his Phycologia australis (Harvey, 1858b; 1859a; 1860b; 1862; 1863).

The Mascarene Islands were particularly favored by individual collectors in the nineteenth century: Annabella Telfair (d. 1832), the wife of the supervisor of the Botanic Garden in Mauritius (Harvey, 1834a); L. Maillard, an engineer working in Réunion (Montagne & Millardet, 1862); Nicolas Pike (1815--1905), the American consul in Mauritius from 1866 to 1876 (Dickie, 1874b); Isaac Bayley Balfour (1853--1922), a Scottish botanist who collected on Rodriguez from 1874 to 1875 (Dickie, 1877b); and Fernand Jadin, a Frenchman who collected on Réunion and Mauritius in 1890 (Jadin, 1894; 1935). While Børgesen (1940 etc.) based his monumental study of the marine algae of Mauritius chiefly on contemporary collections by Theodor Mortensen (1862--1952), a Danish zoologist, and Reginald E. Vaughan (1895--1987), a Welsh botanist who lived in Mauritius from 1923, he took into account all available earlier collections.

Nineteenth-century collectors in other parts of the Indian Ocean included: Theodor Kotschy (1813--1866), an Austrian botanical explorer who collected algae on Karek [Kharg] Island in the Persian Gulf (Endlicher & Diesing, 1845a); Dr. Albrecht Roscher (1836--1860) of Hamburg, who collected seaweeds near Zanzibar in February 1859 as part of an expedition to East Africa and the Comoro Islands mounted by Baron Carl Claus von der Decken (1833--1865) (Sonder, 1879); Odoardo Beccari (1843--1920), an Italian botanical explorer who collected in Singapore and Ceylon (Zanardini, 1872); Charles Thiebaut (1837--1884), a French naval officer who collected in Madagascar (Bornet, 1885); and Johann Maria Hildebrandt (1847--1881), who collected in East Africa, Madagascar, and the Comoro Islands during the period 1872--1881 (Hauck, 1886--1889). Ludwig Preiss (1811--1883), a German plant collector, made significant contributions to our knowledge of the algae of Western Australia, where he collected from 1838 to 1842 (Sonder, 1845; 1846--1848).

The earliest collections of algae from the warm waters of South Africa were made by Carl Ludwig Philipp Zeyher (1799--1858), who traveled by himself from 1825 to 1828 and with Christian Friedrich Ecklon (1795--1868) from 1829 to 1838 (Suhr, 1834; 1836; 1840). Other important contributors in this region were Johann Franz Drège (1794--1881), who collected from 1826 to 1834 (Suhr, 1840; Drège, 1843), Ferdinand Krauss (1812--1890), who collected from 1838 to 1840 (Krauss, 1846; Areschoug, 1851), and Wilhelm Gueinzius (1814--1874), whose collections were sent to Eduard Friedrich Pöppig (1798--1868), a professor in Leipzig (Endlicher & Diesing, 1845b). All of these collectors in South Africa were German except for Ecklon, who was Danish.

The itineraries of the earliest voyages of exploration favored Australia over non-Australian portions of the Indian Ocean. After rounding the Cape of Good Hope, the ships would head for Mauritius (called Île de France until captured by the British in 1810), thence eastward to Western Australia. Ducker (1979b) has provided an insightful and detailed account of these expeditions. Only two resulted in collections of algae from the west coast of Western Australia. The expedition commanded by Thomas Nicolas Baudin, with the ships Geographe and Naturaliste, spent a significant part of 1801 exploring the coast south of Perth (where we now have Geographe Bay and Cape Naturaliste) and Shark Bay. The celebrated naturalist Jean Baptiste Geneviève Marcellin Bory de Saint-Vincent (1778--1846) was a member of the expedition, but discord among the personnel caused him to depart at Mauritius. He published a separate account of his adventures (1804a; 1804b), including the description of the pantropical species Conferva antennina (Chaetomorpha antennina (Bory de Saint-Vincent) Kützing ) from Réunion. The expedition commanded by Louis de Saulces de Freycinet, with the ships Uranie and Physicienne, explored Shark Bay in 1817. The resulting collections of algae were treated by C. Agardh (1822--1823; 1824; in Gaudichaud, 1826) and Lamouroux (in Quoy & Gaimard, 1824). Perhaps the best-known species is Acetabularia calyculus Lamouroux.

The numerous expeditions that entered the Indian Ocean after the middle of the nineteenth century focused on shores other than those of Australia. The voyage of the Austrian frigate Novara (1857--1859) resulted in collections of algae by Anton Jelinek, the naturalist of the expedition, at St. Paul and Amsterdam islands, Ceylon, India, and the Nicobar Islands (Grunow, 1868). The Prussian expedition to East Asia on board the Thetis (1860--1862) brought back algae collected by Eduard von Martens (1831--1904), a zoologist, at Singapore and on the Indian Ocean shores of present-day Indonesia. These were treated by the collector's father, G. von Martens (1868). The voyage of the German corvette Gazelle (1874--1876) resulted in collections of algae from St. Paul Island, northwestern Australia (including Dirk Hartog Island and the Monte Bello Islands), and Timor (Askenasy, 1888). During the round-the-world cruise of the Italian corvette Vettor Pisani (1882--1885), Cesare Marcacci collected in Singapore and Ceylon (Piccone, 1886; 1889a). Andreas Franz Wilhelm Schimper (1856--1901), the botanist of the German deep-sea expedition on board the Valdivia (1898--1899), collected seaweeds at St. Paul and New Amsterdam islands, Sumatra, the Nicobar Islands, Chagos Archipelago, Seychelles, and Tanzania. These were treated by Reinbold (1907).

The scientific expedition of greatest importance to marine phycology in the Indian Ocean in the nineteenth century was the voyage of the Siboga through the Dutch East Indies (1899--1900), organized by Max Weber (1852--1937), a Dutch zoologist. His wife, Anna Weber-van Bosse (1852--1942), made extensive algal collections which were treated in a large number of papers, most importantly Weber-van Bosse (1901; 1904a,b,c; 1913a; 1921; 1923; 1928), Barton (1901), Foslie (1904a), and A. Gepp & E. Gepp (1911). Cambridge University sent a student, John Stanley Gardiner (1872--1946), to make a faunistic and geographical survey of the Maldive and Laccadive archipelagos (1899--1900). His algal collections were studied by Barton (1903a) and Foslie (1903b).

In the twentieth century, Emil Werth (1869--1958), the botanist of the German antarctic expedition on board the Gauss (1901--1903), collected at New Amsterdam Island. These algae were treated by Reinbold (1908). The Percy Sladen Trust Expedition on board the Sealark (1905) targeted the various islands in the west central Indian Ocean. Gardiner, the leader of the expedition, collected algae, which were treated by Foslie (1907c,d), A. Gepp & E. Gepp (1908; 1909), and Weber-van Bosse (1913b; 1914). The John Murray Expedition (1933--1934), organized by the British Museum (Natural History), operated from the Mabahiss in the northwestern part of the Indian Ocean and collected algae in the Gulf of Aden, along the southeastern coast of Arabia, Zanzibar, and the Maldives (Newton, 1953). The University of Cape Town Ecological Survey (1937--1940) was a program organized by T.A. Stephenson and Anne Stephenson, his wife, to study intertidal zonation and biogeography on South African shores. The algae from this survey were studied by Kylin (1938), Levring (1938), and Papenfuss and his students (numerous papers). The overall results were published by T. Stephenson (1939; 1944; 1947).

Despite the activities of these expeditions, and of many others not mentioned because they produced no phycological results, the Indian Ocean was by far the least known of the three major oceans at the time of the International Geophysical Year (1957--1958). Attempts to create a similar plan in support of oceanographic investigations in the Indian Ocean were successful, and the International Indian Ocean Expedition (IIOE) emerged under the aegis of the Special (later Scientific) Committee on Oceanic Research (SCOR), itself under the International Council of Scientific Unions, in turn under the United Nations Educational, Scientific and Cultural Organization (UNESCO) (Behrman, 1981). Many independent expeditions were mounted under this umbrella organization during the period 1959--1965. Three such efforts were of particular significance to marine phycology. Foremost was the land-based expedition carried out by G.F. Papenfuss (University of California, Berkeley) and R.F. Scagel (University of British Columbia), funded by the National Science Foundation (Washington, D.C.). Papenfuss and Scagel collected seaweeds along the East African coast from Djibouti to the Cape Peninsula during the period 15 September--16 December 1962. Before joining Papenfuss at Djibouti, Scagel collected in Western Australia, Mauritius, and Madagascar. Much of the resulting material has been studied by specialists (Abbott, 1970; Ducker, 1967; Johansen, 1977; R. Norris, 1987a,d, 1988b,c,e, 1989a, 1990b, 1991a; Papenfuss & Edelstein, 1974; Papenfuss & Jensen, 1967; Papenfuss, Mshigeni, & Chiang, 1982; Scagel & Chihara, 1968; W.R. Taylor, 1966b, 1967; Wollaston, 1984), but many large and taxonomically difficult genera remain unstudied at the University of California. An expedition to the Maldives was organized by Cambridge University under the direction of David R. Stoddart with algae collected by David C. Sigee during the period 1 July--10 September 1964 (Hollenberg, 1968a; Sigee, 1966; Tsuda & Newhouse, 1966). Although ships from several nations participated in the Indian Ocean Expedition, only one has contributed significantly to marine phycology, namely, the Te Vega, a two-masted schooner operated by Stanford University with funds provided by the National Science Foundation to train students and carry out biological investigations in shallow waters. The most important publications that resulted from the various cruises of this ship are the accounts of benthic marine algae from the Maldives (Wynne, 1993) and from the Seychelles (Wynne, 1995). Other papers based on Te Vega collections include Aregood & Hackett (1971), Hackett (1969; 1977), Hollenberg (1968c), and Wynne (1970a; 1970b).

In the thirty years since the end of the International Indian Ocean Expedition, marine phycology throughout the world has changed markedly. An ever-increasing proportion of phycological publications deals with biochemistry, physiology, ecology, and commercial utilization. Consequently, the need for accurate identification of the algae used in these studies is greater than ever. The classical diagnostic tools of comparative morphology, biochemistry, reproductive biology, and ecology are now routinely supplemented by cladistic and genomic analysis. In the Indian Ocean region, most of the resources of taxonomists have been applied to the study of collections resulting from the International Indian Ocean Expedition or collections made by resident phycologists in Australia, South Africa, and India. The Indian Ocean shares with other oceans the problems of such troublesome genera as Gracilaria, Hypnea, Ceramium, Laurencia, Polysiphonia, Dictyota, Sargassum, Enteromorpha, Ulva, Chaetomorpha, Cladophora, and Caulerpa, all of which are badly in need of monographic study. Especially in these genera, determinations of Indian Ocean seaweeds have often been made on the basis of illustrations of seaweeds in the Atlantic and Pacific oceans. Undoubtedly many of these determinations would prove incorrect if subjected to critical study. There is still the need to locate and examine type specimens of names of uncertain application, although the number of such names has been greatly reduced through the efforts of G.F. Papenfuss, H.B.S. Womersley, S.C. Ducker, and their academic offspring. Because taxonomy is a synthesis of all previously published data from all fields of investigation that provide distinguishing characters, the older literature may not be ignored. The present catalogue was designed to facilitate the task of integrating the new with the old.