Fungi of Puerto Rico and the United States Virgin Islands a History Of
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FUNGI OF PUERTO RICO AND T H E UNITED STATES VIRGIN ISLANDS A History of Previous Surveys, Current Status, and the Future D. JEAN LODGE Center for Forest Mycology Research Forest Products Laboratory United States Department of Agriculture Forest Service PO. Box 1377 Luquillo, Puerto Rico 00773 H E NEW YORK ACADEMY OF SCIENCES published two issues containing fungi in the Scientific Survey of Porto Rico and the Virgin T Islands in 19261 and 19322,3 (Volume VIII) under the Phyllum Thalo- phyta. Algae, bacteria, actinomycetes and lichens were also included in Phyllum Thalophyta by the survey, but are not discussed in this chapter. Today, fungi are classified in a separate kingdom, with the true fungi in a separate subkingdom (Eumycota) from the slime molds (Myxomycota). With the pub- lication of the Supplement to Mycology in 1932,1 the New York Academy of Sciences (NYAS) Scientific Survey covered approximately 1610 fungi in- cluding the slime molds. According to Stevenson,4 the first fungi that were recorded from Puerto Rico were apparently collected by an Italian physician, Carlos Bertero, in 1818 and described by Montagne and Leveille. Seaver and Chardon I summarized the history of mycology in Puerto Rico, beginning with a few collections dating from 1847-1850 by Carl Schwanecke, a German botanist, that were identified and published by J. F. Klotzsch in 1852.5 A second, larger collec- tion was made by P. Sintenis from 1884-1887, and these were identified by 6 J. Bresadola, P. Hennings and P. Magnus. Attention to mycology increased after Puerto Rico became a part of the United States of America at the end of the Spanish-American war. Various collections were made from 1900-1910 by pathologists with the Insular Department of Agriculture and visiting bot- anists. Among the visitors were A. A. Heller and Mrs. Heller in 1900, G. P. Clinton in 1904, and E. W. D. Holway in 1910. Dr. and Mrs. N. L. Britton and others from the New York Botanical Garden made numerous collections of fungi during collecting trips from 1906 to 1932. In 1911, J. R. Johnston joined the Insular Department of Agriculture in Rio Piedras, Puerto Rico as the Pathologist, and several agaric fungi that he collected were named by others 123 124 ANNALS New York Academy of Sciences in honor of him. During 1915, B. Fink visited Puerto Rico to study lichens but also collected many fungi, and H. H. Whetzel and E. W. Olive came to study rust fungi (their collections were reported by J.C. Arthur). In 1918,J. A. Steven- sen of the Insular Department of Agriculture published a checklist of fungi 7 1-3 for Puerto Rico, which served as the foundation for the NYAS survey.- In addition to the fungi on the 1918 checklist,7 the latter survey1-3 included an extensive collection of more than a thousand fungi that was made in Puerto Rico and the U.S. Virgin Islands during 1923 by Fred A. Seaver of the New York Botanical Garden, Bronx, NY, and Carlos E. Chardon of the Insular Experi- ment Station in Río Piedras, PR, in the company of Dr. and Mrs. N. L. Britton. The largest fungal collections in Puerto Rico prior to Seaver and Chardon were by Professor F. L. Stevens when he was the Dean of the College of Agri- culture at Mayagüez. Stevens's collections of rusts were studied and reported by J. C. Arthur, while F. L. Stevens and his collaborators at the University of Illinois published papers on various other groups of fungi. A later collection of rust fungi was made in 1924 by H. H. Whetzel of Cornell University, and F . D. Kern of the Pennsylvania State College, often accompanied by Rafael A. Toro of the Agricultural Experiment Station at Rio Piedras, PR. These and other records of Puerto Rican fungi were reported in 1932 by Seaver et al.2 in the Supplement to Mycology of the NYAS Scientific Survey. The New York Academy of Sciences' survey of fungi in the U.S. Virgin Islands was preceded by a Danish Survey, with collections dating from at least 1715 to 1906. Raunkiaer, F. Børgesen, and Ø. Paulsen collected in what were then the Danish West Indies, and some of the collections were described or 8 reported by B0rgensen and Paulsen, Ellis and Kelsey,9 Ferdinandsen and 10, 12 Winge, 11 and Millspaugh. Raunkiaer13 also published a survey of myxomy- cetes in the Danish West Indies, and Ferdenandsen and Winge14 later de- scribed a pathogenic water mold collected by Ostenfeld on St. Croix. Rust and smut fungi (18 spp .) and species in the Xylariaceae (17 spp.) were also well rep- resented in the Danish works,10,11 and these records were incorporated in the later survey by the NYAS.1-3 Apparently, no American mycologists had col- lected in the U.S. Virgin Islands prior to the expedition by Seaver and Chardon in 1924. In the taxonomic survey of fungi in Puerto Rico and the U.S. Virgin Islands by the NYAS- 1 3 rust fungi and other plant pathogens, discomycetes, polypore fungi and slime molds were especially well represented. However, relatively few ephemeral species of basidiomycetes (55 spp.) were reported, and many of these were not from forests. These patterns can be best understood in a his- torical context . During the time covered by the NYAS survey, almost all of the island of Puerto Rico was in agricultural use,15 and travel across the island was quite difficult. 16 By 1912, less than 1% of the forest in Puerto Rico was still Virgin.,16 17 Destruction of native forest was also severe in the Virgin Islands .13 It was probably for these reasons, in addition to the economic im- portance of fungi as plant pathogens, that coverage of the fungi was primarily from agricultural rather than natural forested areas. As a result of work by Stevens, Arthur, Whetzel and Kern, the rust flora of Puerto Rico was largely known and well covered by the NYAS survey.1,2 Other plant pathogenic fungi Fungi of Puerto Rico and the Virgin Islands LODGE 125 were also quite extensively covered. 1,2 Collections of forest fungi were largely those of Seaver, Chardo n, Johnston, Toro, and Brittain, but numerous species of wood decomposing fungi in the Xylariaceae (Ascomycetes) and Polyporaceae (Basidiomycetes, determined by Overholts) were collected by various botanists and mycologists and are reported in the survey. 1 Most of the species in these groups are rather tough and retain their characteristics when dried, and are hence easy to collect and preserve in an identifiable state. Another group that was especially well covered by the survey' was the Pezi- zales (57 spp.), owing to the special interests of Seaver. Although the number of slime molds listed by the survey2 probably only represents a part of the true diversity in this group, numerous species of Myxomycetes were reported owing to the special interests of Raunkiaer 13 and Hagelstein.3 The absence of agaricologists among the visiting mycologists undoubtedly contributed to the underrepresentation of this group. In addition, these fungi are more difficult to collect, annotate, and preserve in good condition, and their fruiting is often sparse and sporadic. 13,18 RECENT SURVEYS OF FUNGI Almost all reports of fungi from Puerto Rico and the American Virgin Islands were summarized in an annotated bibliography by J. A. Stevenson4 in 1975. Many citations of more recent reports on plant pathogenic fungi were added to those published in the NYAS survey,1-3 and the taxonomy and sys- tematics was updated . Stevenson4 also included reports of slime molds iden- tified by Dr. C. Alexopoulos from the El Verde Research Area in the Caribbean National Forest, Luquillo Mountains that were reported in an ecological re- search volume .19 A few species of macro fungi identified by Cowley,20 and a survey of micro fungi on living leaf surfaces by Hutton and Rassmussen21 at El Verde in the same volume were not included by Stevenson.4 What was probably the first survey in Puerto Rico of micro fungi growing on forest leaf litter and soil by Holler22 was also not included in Stevenson.4 There are several studies of fungi in Puerto Rico that were published or sub- mitted after 1970 and consequently were not included in the Fungi of Puerto Rico published by Stevenson in 1975.4Among these are several surveys of Caribbean discomycetes by D. Pfister23-26 while he was a professor of the Uni- versity of Puerto Rico at Mayagüez, a survey of aquatic hyphomycetous fungi on leaf litter in streams at El Verde by Padgett,27 and miscellaneous reports and descriptions of ascomycetes by G. Samuels and collaborators,28-31 J. D. Rogers and collaborators,32-34 and Laessøe and Lodge.35 The U.S. Forest Ser- vice surveys of forest pests in Puerto Rico were conducted in 1973, 1976, 1981, and 1984,36-39 and included fungi that infect trees, but emphasis was placed on introduced tree species and plantation forestry. From 1983-1992, D. J. Lodge was employed by the Division of Terrestrial Ecology, University of Puerto Rico in San Juan, to study the role of microorganisms in rainforest nutrient cycles and food webs at the El Verde Research Area in the Luquillo Mountains. As a result of this work, new records of fungi were reported in the context of fungal ecology and nutrient cycling.40,41 New reports and descrip- tions of new species of Hygrocybe (Hygrophoraceae),42Mycena (Tricholo- 126 ANNAL s New York Academy of Sciences mataceae),43 and Neopaxillus (Paxillaceae) 44 were also published in the taxo- nomic literature .