.V; “W' 1‘ “ :r -- o s' .n. Q i! ::':I.‘ .'vv. L.‘ I--.‘ '0 Do 0. O.¢’,‘ 3216: gm N ”‘43:?“ . ‘ a: "v0... 0 I...- .n ABSTRACT THE LICHENS OF LONG ISLAND, NEW YORK: A VEGETATIONAL AND FLORISTIC ANALYSIS by Irwin ' M. Brodo The lichen vegetation of Long Island is discussed in broad perspective, and yet with considerable detail, in an attempt to present a relatively complete picture of an important segment of the North American east coast lichen flora. A floristic list based on complete collections made throughout Long Island and some adjacent islands is supplemented by a number of investigations of local problems in lichen ecology. The ecological studies consist of transect analyses along the island's north shore, transplant experiments concerning the vertical distribution of corticolous species as well as the city effect, analyses of the present distributions of various species by vegetation type, and ob- servations on succession and related phenomena in terricolous, saxicolous, and corticolous commities. A habitat classification is used to group assemblages of lichens into "cominities." Some discussion is presented on the relative merits of such a loose classification as compared with a more formal lichen ”union" or "association" system used by many European workers. A consideration of some of the environmental factors influencing lichen microdistributions is presented along with some supporting mgurements and correlations, but no extensive work along these lines is pursued. The effect of New York City on Long Island lichen distributions is discussed in some detail. Empirical data and theoretical considerations ~‘ 1- ML?! [it 1:73.; at? ?;a:'.‘._' 3'13th -2 1': cute." 3?!!!” Z: are used in concluding that the lichen distributions are influenced ‘by air pollution as well as city-induced drought, with the former acting over longer distances than the latter. Placing the Long Island lichen flora into phytogeographic perspective involved setting up a scheme of "elements" and "subelements" for eastern North America into which the lichens could be fit. The presence of Long Island species in Asia and Europe was noted and consideration given to problems of migration and vicariism. The lichen flora consists of 260 species. Keys to the identifi- cation of these species, including keys to sterile material, precedes an extensive annotated list. Included under each species in this list is reference to material seen, notes on habitat ecology, a state- ment on North American and world-wide distribution, and where necessary, notes on nomenclature, morphological and chemical variation, and closely related and/or confusing species. Three species are described as new: Polyblastiopsis uercicola, Pertusaria sub ertusa, and Lepraria gggggg, In adddition, three new combinations are made: Micarea prasina var. sordidescens (Nyl.) Brodo, Parmelia‘michauxiana var. laciniata (Hale) Brodo, and Buellia curtisii (Tuck.) Imsh. in Brodo. THE LICHENS OF LONG ISLAND, NEW YORK: A VEGETATIONAL AND FLORISTIC ANALYSIS by Irwin M. Brodo A THESIS Submitted to Michigan State University in partial fulfillment of the requirements for the degree of DOCTOR OF PHILOSOPHY Department of Botany and Plant Pathology 1965 311;: is f...“ :e'.’ 3:13.114 31.1.32! I 13: 1113.41: . I a! u.... ‘0... _ ACKNOWLEDGMENTS This work had its genesis due to the generosity of the New York State Means: and Science Service through their Sumner Honararium program. Financial help from the Science Service, in combination with the encouragement, help, and guidance of Dr. Eugene Ogden made the first two summer's work possible. In 1961 Dr. George Woodwell at Brookhaven National Laboratory provided a room, and laboratory facilities, and for this I am indeed grateful. I sincerely thank the National Science Foundation for awarding me a Graduate Fellowship for the years 1961-62 and 1962-63, and giving me the opportunity for uninterrupted research during that time. Without the skilled and dedicated help of Dr. Henry Imshaug, this paper would have suffered greatly. His guidance, advice, and good humor 'were truly an inspiration. Dr. John Cantlon's many critical comments and valuable suggestions are greatly appreciated. Drs. Ervin Barnes, Edward Cantino, Roland Fischer, and the late Dr. Philip Clark, all contributed suggestions and comments on the manuscript, and for these I am.grateful. I would like to especially thank‘Mr. Roy Latham, not only for lending me his entire lichen collection which was of such fundamental importance to this work, but for his amiable and informative letters concerning the Long Island of past years, and for his companionship on several exciting and fruitful field trips. Of the many Long Island residents and naturalists who led me to rich collecting areas and provided trans- portation to some relatively inaccessible areas, I would like to extend special thanks to Mr. Leroy Wilcox, Mr. Gilbert Raynor, and Miss Linda Quinby. Mr. Fred McKeaver's hospitality and guidance on Nantucket Island ii “r- y--.r__-_‘ --.- Tl? 0. Os "1: u .. are greatly appreciated. my sincerest gratitude goes to my very patient and tireless wife, Fenja, for her many and varied assistances. Thanks are due to the following lichenologists for identifying or verifying Long Island material in their special fields of interest: T. Ahti (Cladina), W. L. Culberson (Cetraria), F. Erbisch (Chaenotheca), A. W. Evans (Cladonia), M. E. Hale (Parmelia), W. Harris (Polyblastiopsis, Leptorhaphis), A. Henssen (Placygthium), A. W. C. T. Herre (Eggga), G. Howard (Ochrolechia), I. M. Lamb (Stereocaulon), A. H. Magnusson (Ramalina), E. D. Rudolph o laca), H. Sierk (Leptogium), D. Swinscow (Porina), 'W. weber (Agarospora), C. H; wetmore (Nephroma). Mr. W. D. Margadant of the Hunt Library kindly helped me with the latin diagnoses. iii g1 ‘. .- Os ‘ I 2 4.. , 1 c to . TABLE OEJCONTENTS' Sections 1. Introduction A. General 8. History II. Description of Long Island III. A. Geography C. D. Geology Climate Vegetation types Habitat Ecology A. C. D. F. General methods 1. Collection data 2. Statistical studies 3. Transplant experiments Substrate 1. Texture 2. Mbisture holding capacities 3. Stability 4. Chemical composition Climate l. Illumination and temperature 2. Mbisture Vertical distribution Succession Species composition within habitats iv Page 18 18 18 18 21 22 24 25 29 31 34 35 36 38 43 51 Co. 2" . nu . . 'Q’ . o 0- .1! o E. ,3- . .' 4 O C '00.... A. '... a . ‘ ‘ A O. -.“ 'tuz . 0 " .13: f . .. :’..: ‘ .. ‘ IV. City Effect A. Introduction B. Methods and observations 1. Transect studies 2. Distributions 3. Transplants C. Results D. Discussion E. Conclusion V. Floristic Elements A. Introduction B. Classification of elements C. Summary of significant features D. Discussion E. Summary VI. The Lichen Flora A. Collections B. Additional specimens seen C. Taxonomy 1. Species concept 2. Ecological forms 3. Infraspecific taxa 4. Keys and annotated list ‘VII. General Discussion A. Distribution of lichens on Long Island 1. Substrate 2. Climate 75 75 77 77 80 80 83 84 94 95 95 96 100 101 108 110 110 114 115 115 118 120 120 359 359 359 361 U“ .‘. e -4 B. 3. Vegetation type Floristic changes 'VIII. Summary and Conclusions D. Habitat ecology and lichen communities Lichen distributions The city effect The lichen flora Appendices (see page xi) Literature Cited vi 362 367 372 372 373 374 375 376 415 e0. o.. .01-. 3. 4. 6. 10. 11. 12. l3. 14. LIST OF TABLES Degree of similarity of the lichen vegetation growing on various species of oaks in red oak forests. Coefficients of association of lichen vegetation on different tree species at base and breast height quadrats. Bark characters of some common Long Island phorophytes. Sand and soil pH. Vertical distribution of some corticolous lichens in red oak and pine-oak forests Results of the east-west corticolous lichen transplant experiments using Parmelia caperata. Lichen species diversity in various habitat types in ten localities closest to New York City. Growth forms and city tolerance along a north shore red oak transect. Atmospheric pollutants in New York City and a suburb. Phytogeographic categories represented in the Long Island lichen flora. Phytogeographic affinities of Long Island lichens. European-American vicarious sub-generic taxa in the Long Island lichen flora. Distribution of some common lichens in various vegetation types on Long Island. Average annual growth rates of some corticolous lichens growing on oak. vii 440 441 442 443 444 445 446 447 448 449 455 456 457 462 10. ll. 12. 13. 14. 15. 16. 17. 18. 19. 20. LIST OF FIGURES Soil types. Original vegetation. . Precipitation. a. Growing season. b. Annual. Temperature. a. January. b. July. The fog belt. . Humidity levels. a. July. b. January. Sand dune vegetation. Sand dunes and sand plains. A sand plain community. North shore bluffs. Pine-oak forest. Black oak forest. Gravel pit bog. Verrucaria microspora habitat. Collection localities. Non-directional fluctuations in a corticolous community. Succession in a corticolous lichen community. East-west transect and transplant localities. Distribution of the common trees along the north shore transect. Tree size along the north shore transect. viii 463 463 463 464 464 464 465 465 465 466 466 466 466 467 467 467 467 468 469 470 471 472 473 .. .. . "- .I . Q. .‘ .- 5"‘.‘ ‘e e O ' C D. 4!. ‘ \ h. “ :9. 5‘. . . n A. .1..- ...‘ e 's .. . C' ‘ ‘1‘ s ‘e 9. .. : o. a“ -‘. - 5 I o. . ‘ e.\ . U s ..1 .. a .D I. I'. .I. D a"... 21. Distribution of some common lichens along the north a. Arithmetic plot. b. Logarithmic plot. 22. Bark-borer. 23. Transplanted bark disks. 24. A possible mechanism for the city-effect. 25. Arctic-Boreal element. 26. North Temperate subelement. 27. Appalachian unit. 28. Appalachian-Ozark unit. 29. Appalachian-Great Lakes unit. 30. Appalachian-Great Lakes-Rocky Mountain unit. 31. Coastal Plain subelement. 32. East Temperate subelement. 32A. Oceanic subelement. 33. Phytogeographic affinities of Long Island lichens. shore transect. 34. Historic relationships between floristic elements, subelements, and units. 35. Localities of oceanic species. 36. Bog and swamp localities. 37-85. Distributions of some lichens on Long Island showing the influence of vegetation type and climatic factors. 85-100. Distributions of some lichens on Long Island showing the city effect. 101. Pertusaria subpertusa (holotype). 102. Polyblastiopsis guercicola (holotype). 103. Lepraria‘zonata (holotype). 104. Polyblastiopsis guercicola (spores, etc.). ix 490- 474 475 476 477 478 479 480 481 482 483 484 485 486 487 488 489 490 490 493 493 494 494 494 495 105. Ascocarps. 106. Thallus types. 107. Lichen phycobionts. 108. Some ascospore types. 496 497 497 497 -;:c‘ Appendix A - Appendix B - Appendix C - I- II - Appendix D - APPENDICES Studies of Lichen Growth Rates on Long Island A. - Methods B - Results C - Discussion D - Summary Long Island Collectors Glossary Mbrphological and Ecological Terms Chemical Terms Checklist of the Lichens of Long Island Xi 376 376 379 380 382 384 386 386 401 409 I. INTRODUCTION A. General. Eastern North America has probably received more lichenological study than .eny'other part of the continent. Such famous and productive workers as Edward Tuckerman, Henry Willey, Lincoln W. Riddle, R. Heber Howe, George K. Herrill, Charles A. Robbins, Alexander W. Evans, and Guy G. Nearing devoted much of their lives to the study of northeastern lichens. Yet with this exceptionally fine.background of basic taxonomic knowledge, no recent workers studied this area using modern methods of floristic analysis and taxonomy until Gunnar Degelius visited the United States in 1939 and published two excellent papers, one dealing with the lichens of Maine (Degelius, 1940) and the other with the lichens of the Smoky Mbuntains of Tennessee (Degelius, 1941). In 1950, Hale wrote an account of the lichens of Aton Forest in northeastern Connecticut, and inl954, I. Hackenzie Lamb published a study of the lichens of Cape Breton Island, Nova Scotia, both papers significantly adding to our.knowledge of the northeastern coast lichen vegetation. Culberson (1958) reported on some lichens of North Carolina but dealt only with the pine-inhabiting vegetation. This paper, then, is mainly designed to contribute to our knowledge of the eastern coastal plain vegetation, and, by so doing, to provide a link between the studies of the northern coastal regions and the Appalachians. The principles which guided the research summarised here were that a vegetation cannot be adequately written without a thorough knowledge of the flora, and that a flora cannot be understood without a study of the ecological and phytogeographic factors which fashioned it. In a study of this scope, it is impossible to answer all or even most of the questions asked concerning relationships and factors involved in the vegetational picture. It is my earnest hope that this study will point to the many taxonomic, ecologic, I: ~ ’.V._'r ‘11 31.11! 5“: '°' .' a “- ..Q ”:4 '. «fl 3. 2 .and.phytogeographic problems still in need of clarification and solution, and will provide a stimulus for other workers to add to our knowledge in these and related fields. B. History. Long Island lichenology surprisingly had its beginnings quite early in the history of American botany. Halsey (1823) published a list of lichens collected ”in the vicinity of New York," but he did not state explicitely that he collected east of the East River and there is some doubt as to whether he listed any Long Island specimens. Specimens which were collected in Brooklyn and Queens by George B. Brainerd and by George D. Hulst during the 1860's may very well be the earliest from Long Island. Their collections, which are deposited in the Brooklyn Botanic Garden Herbarium, provide a good basis for reconstructing the probable state of the lichen vegetation of eastern New York City prior to urbanization (see page 368). Other collectors of Long Island lichens during the late 19th century include Charles H. Peck who collected all forms of plant life throughout New York State during his tenure as New York State Botanist. His collections are in the New York State museum. In 1899, S. H. Jelliffe published "The Flora of Long Island” which listed 54 lichen taxa from various parts of the island. G. S. Wood (1905) published additions to the lichen flora adding 18 taxa to Jelliffe's list. In 1914, Wood published a list of the lichens growing in the vicinity of New York City and included many species from Long Island. The Cold Spring Harbor area was fairly well collected, not only by Jelliffe and Wood, but also by A. J. Grout in 1900 and Stanley A. Cain in the 1930's in.connection with the Long Island Biological Institute at Cold Spring Harbor; Since that time, however, no botanical field work has been done there. Lra 1;-ze are? a: CQIJZI 1.321.? 4.3. IQ? 0.- .... Jul...” 3 Some lichens collected by Stanley Cain as part of the "Flora of Cold Spring ‘Harbor".are represented in the New York Botanical Garden Herbarium, but no specimens collected by Jelliffe or Wood were seen. Unfortunately, the complete collection of the Cold Spring Harbor flora which did exist at one time (Cain, personal communication) could be located neither at the Biological Laboratories at Cold Spring Harbor itself nor elsewhere. Roy Latham, one of the most versatile, thorough, and knowledgeable of the Long Island naturalists, began collecting lichens in 1908. He confined his collecting to eastern Long Island, especially around Orient Point, and rarely went as far west as Hanorville. Latham's first concentrated effort was connected with his publication of the "Flora of the Town of Southold, Long Island..." in collaboration with S. H. Burnham (Burnham and Latham, 1914-1925). The Farlow Herbarium includes many of these old Latham specimens which had been sent to Riddle, Hesse, or Herrill for identification. Since the early 1900's, Latham has collected about 2000 lichen specimens including many rare species. His is by far the most complete collection of lichens made on Long Island previous to these studies. Hr. Latham kindly provided his, entire collection for my use. Approximately 2/3 of the collection are species.of Cladonia.a The Cladonia specimens were almost all determined in duplicate by Alexander Evans with whom.Latham carried on an active correspondence until Dr. Evans' death in 1960. Nhny of Lathamds collections represent the only specimens collected of some species rare on the island (see page 369). Hr. Latham continues to be active, and I have had the good fortune to accompany him on several collecting trips in eastern Long Island. Raymond Torrey had a strong interest in lichens, especially of the New York City area, and made many collecting trips to Long Island particularly I'l [g gr- .I u'~oo .I..t-~g.~ :1 37247 0.. 117‘" if; 4 to study the Cladoniae. His interests were not confined to the genus Cladonia, 'however, as is evidenced by his paper on Long Island rock tripes (Torrey, 1933). The New York Botanical Gardens Herbarium contains Torrey's Cladonia collections. These specimens were all identified by Evans and prepared for the herbarium by John W. Thomson (Thomson, personal communication). It is surprising that no other genus of lichens is represented in the Torrey collections. Although Babette Brown Coleman collected and published on some lichens from Montauk Point (Brown, 1948), no extensive collecting other than Latham's has occurred in recent years. (A complete list of Long Island collectors can be found in Appendix B). . -0- Q‘ A -31: .' .a: .1 7:! pa 4 DC 6 s II. DESCRIPTION OF LONG ISLAND A. Geography. Long Island makes up an eastern extension of the southern tip of New York State lying just to the south of the Connecticut coast and separated from the mainland to the north by Long Island Sound and to the west by the East iRiver and‘Nanhattan Island. Long Island is 116 miles long and, at its broadest point, 20 miles wide. There are several smaller associated islands just off the shores of Long Island, and these were visited and included in the study wherever possible. Included were Long Beach, Jones Beach, Fire Island, Westhampton Beach, Shelter Island, Gardiner's Island, and Fisher‘s Island; not included were Robbin's Island (a small island in Peconic Bay) or Plum Island, which is quarantined and not open to the public. The geographical unit, Long Island, is subdivided into four political units: Kings, Queens, Nassau, and Suffolk Counties. Kings County (more widely known by its borough name - Brooklyn) and Queens County are part of New York City. Brooklyn is very populous and, except for one or two large parks and some swampy areas to the south, is covered to a large extent with brick, concrete, and asphalt. Queens is not quite so built up and still has many areas of more or less natural woods and swamps. Forest Park, in the center of one of the most populated parts of Queens, and Alley Pond Park farther east, still show the magnificent red and black oaks (Quercus Egbgg} and Q, velutina) and tulip trees (Liriodendron tulipifera) which characterized the forests of that area prior to urbanization. Nassau county is a classical example of suburbia. Extensive housing 1 All phanerogamic nomenclature follows Fernald (1950) unless otherwise noted. v aft]. - m ..- .. . ,.- g . A n ‘1 ~... .. .. ‘o.- 0,, ...e - s.". .4 ‘.. ‘ 6 developments occupy its central portion and large estates are common on the- north shore» Inch of the area is still relatively undisturbed especially on the larger tracts of privately owned property to the north. The largest county in size and the smallest in population is Suffolk County. Although suburban developments are frequent along its western edge, the greater part of the area is made up of farmland and undeveloped pine barrens. Potatoes and cabbage are the chief crops produced. Resorts are common along the entire south shore. B. Geology. Prior to the Wisconsin glaciation, the entire area now Long Island, except for the western corner, was under water and was covered by a number of marine sediments (thClintock & Richards, 1936). Early Wisconsin glaciation (The Iowan-Tasewell complex) laid down two morainal ridges over this sediment. The first, the Ronkonkoma moraine, runs through the center of the island eastward to Montauk Point and then off the coast to Martha's Vineyard and Nantucket Island and probably resulted from the Farmdale advance (Flint, 1953). The second, caused by a readvance of the ice (the Iowan advance) after a slight withdrawal, formed the Harbor Hill moraine extending eastward to Orient Point, then to Fisher's Island, and finally to Cape Cod. A third advance, the Tasewell, overrode the Harbor Hill moraine (Flint, 1953) and produced many of the major topographic features we now see on the north shore such as the bluffs (figure ED, bays and inlets (Nichols, 1958). A broad outwash plain is associated with each moraine, and it is especially extensive south of the Ronkonkoma moraine where it forms a low, flat, sandy plain southward to the ocean. Wave action and ocean currents formed the off-shore barrier beaches, Fire Island being the longest. 7 Bedrock can be found only at the western edge of Long Island in Astoria (Queens). The topography of Long Island is entirely glacial in origin. With the exception of the moraines mentioned above, the land is extremely flat. The highest point on the island is 428 feet above sea level at High Hill near South Huntington. Kettle holes with associated bOgs or lakes are scattered throughout the island (Fuller, 1914; Nichols, 1958). The soils are formed on glacial parent material and are more or less sandy, very well drained, and usually fairly acid (figure 1). The morainal areas are characterized by medium to moderately coarse textured glacial till (Plymouth- Haven association) often bearing large glacial erratics. Acid sandy-loams with fairly good moisture capacities (Bridgehampton associations) lie to the south of the moraines in most places. Very well drained and very acid coarse textured gravel and sand of the glacial outwash (Colton and Adams associations) make up a large part Of the southern edge of the island. In central Nassau County the soil morphology is much like that of a typical prairie (Hempstead- Bridgehampton association) The soil is well drained, highly acid, and with a dark-colored surface layer (Cline, 1957). C. Climate. The precipitation over the greater part of the island is approximately 40 to 50 inches per year, or about four inches per month, except for the dry months of June and July (figure 3). Droughts are not uncommon in central Long Island. More than once a year, on the average, there is a "dry spell" (a period of at least 15 consecutive days, none of which receives 0.05 of an inch or more of precipitation). Approximately once every two years there is an “absolute drought" (15 consecutive days, none of which receives 0.01 8.1: 7' I - ‘ ‘ . i ' I .. - - - o A 3! 171?: if“ :1 It 8 of an inch of rain or more). East of Three Mile Harbor, the rainfall averages 30-40 inches per year. (Data and definitions kindly furnished by Brookhaven National Laboratory Meteorology Group.) Temperatures on Long Island are rather mild and differences are slight from one part of the island to another. 0n the average, the winter temperatures are about the same dhroughout the island, but are milder than farther inland due to the oceanic effect. Summer temperatures grade from warmest in the New Ybrk City area to coolest at the eastern half of the island (U.S.D.A., 1941) (figure 4). At Brookhaven National Laboratory, in central Long Island, the average recorded temperature was 65° F. between October lst and September 30th, and 40° F. between October lst and April 30th. Temperatures in that area rarely go below 10° F. or above 90° F. Winds are quite brisk all over the island. In the central portions, over half the days of the year have winds between 11 and 18 miles per hour. Nontauk Point on the eastern tip of the island is well known for its high winds. Prevailing winds are from the southwest during the summer and from the northwest during the winter. Fog and mist are common phenomena on the eastern tip of Long Island, particularly in the Montauk area (see figure 5). Depressions in the downs and between the dunes where fog can form create local pockets of extremely high humidity in the Montauk region (see also page 37). Almost every autumn, Long Island is subjected to violent storms which originate as hurricanes in the Caribbean and sweep up the east coast. Host of the storms do only minor wind damage to the plant communities but occasionallyr severe stormm cause extremely high tides, violent winds, heavy salt spray, and driving rains which do considerable damage along the coast and even farther inland, particularly on the eastern tip of the island. Roy Latham .11 3.1L "‘ e .—. s... . 0.- ..- ‘~ ..... s . K s 9‘... N 43!: ‘t 9 (in a letter) relates how the hurricanes of 1938 and 1944 completely flooded the beach at Orient Point (Long Beach) and swept away a great quantity of vegetation including all but traces of the lichen flora. Tides rose 12 feet and even the corticolous lichens were washed into the ocean. Trewartha (1961) in his modification of K8ppen's classification of climatic regions placed the Long Island area into his "Daf" category indicating a humid, continental climate, with warm summers. In summary, the climate of Long Island is characterized by periodically droughty, warm summers and rainy, mild winters. To the normally warm and droughty summers are added high winds and excessively drained soils greatly increasing vegetational drought. The situation is locally alleviated somewhat by moist on-shore winds and fogs in the extreme eastern part of the island, where the rainfall is the least and the winds are the highest. D. Vegetation Type . When one speaks of the "vegetation of Long Island," it must be understood that in many areas, there are two vegetations to be discussed... the present, and that of the presettlement period. This is especially true in the New York City area and adjoining Nassau County where urbanization virtually eliminated once important and conspicuous vegetation types leaving only fragmentary remnants. For example, Forest Park on the Brooklyn-Queens boundary is the only surviving remnant of a forest described as having been "heavily wooded with large timber of an aspect similar to the timber of the Connecticut coasts" (Svenson, 1936). As late as 1917, Harper (1917) reported some remnant forests in the Queens area as constituting rich woods broken with streams and meadows. Some of the larger trees Harper listed as being most abundant were 92ercus velutina, Q, 5125, Hicoria alba (1: Carya tomentosa), and Castanea dentata, with 92ercus coccinea being important in the drier woods I11: as i 211.12: 10 and Liriodendron tulipifera being conspicuous in the rich woods. Another excellent example of this massive vegetational obliteration can be seen in Nassau County in the "Hempstead Plains" region. Originally, this area was a 16 mile long botanical oddity... a natural true prairie on Long Island. The land was not good for farming because of the dense hard sod, but it was used extensively for pasture (Svenson, 1936). Hicks (1892) wrote a detailed account of the flora of the Hempstead Plains. The broad, unforested, gently rolling landscape provided a perfect situation for mass produced housing, and after the great expansion in suburban living just after World ‘War II, many housing developments arose on the "plains" such as those in Levittown, Garden City, and Mineola. At this date, the only remnants of this fascinating vegetation type can be found on fragments of the property adjoining some parts of the Meadowbrook Parkway and parts of Mitchell Air Force Base. It will later be pointed out that the lichen flora occurring on these fragments is amazingly rich for such a far western position on Long Island. The original vegetation of Suffolk County on the other hand, although fragmentary and relegated to parks in some areas to the west, remains in a more or less recognisable state (figure 2). Canard (1935) presented a vegetational analysis of the vegetation types of central Long Island giving them phytosociological binomials. Among the most conspicuous comnnnities can be recognized the well developed oak forests seen mostly on the north shore (Harper, 1917; Cain, 1936), the pine barrens which are well developed in central Long Island eastward to Riverhead (Harper, 1908; Britton, 1880), and heathlike ”downs” (as described by Taylor, 1923) which are very conspicuous in the Montauk area. Also important are the communities characteristic of :2 51:: :.:es ' 11 the sand dunes (Brodo, 1961a), the Chamaecyparis bogs (Bicknell, 1908; Harper, 1907; Nichols, 1907; Taylor, 1916), the red maple swamps (Cain & Penfound, 1938), and the Hempstead Plains (Hicks, 1892; Harper, 1911, 1912; Cain, Nelson, & MCLeen, 1937). A more detailed breakdown of the plant associations has been made by many authors (Miller a Young, 1874; Jelliffe, 1899; Taylor, 1915, 1922; Grier, 1925; Conard, 1935; Svenson, 1936; Brodo, 1961a). The names used in the following descriptions are those most widely accepted and used by the above authors and other naturalists in the area. The categories I used in a previous paper (Brodo 1961a), although well suited for describing central Long Island stands, ‘had to be somewhat expanded to be of use in depicting the vegetation types throughout the entire island. 1. Dune grass - Beach Hggther - Shrub Savanna and Sand Plains (formed on dune sand; excluding pine barren glades) (figures 7, 8 ). Dominant trees: Piggy rigida, Prunus serotina (both sparse and usually stunted). Dominant undergrowth and ground cover: Ammophila breviligulata, Hygica pgnnsylygnicg, Prunus maritime, Arctostgphylos uva-ursi, Hudsonia tomentosa. Soil: quartz dune sand with little or no organic *matter. Lightz: unlimited. Most of the barrier beach on the south shore and a few small areas on the north shore are composed of long, rolling dunes, some still moving. The best developed dunes and their corresponding vegetation can be found along the 2 Light: unlimited = almost entire area in open sunlight; excellent:= at least 1/4 the area in open sunlight, the rest in moderate shade; good:: less than 1/4 the area in open sunlight, the rest in moderate shade; fair;= no open sunlight falling on ground, but some sunlight filtering through the trees; poor:; tightly closed canOpy with virtually no sunlight reaching the ground. I 12 entire length of Fire Island, and magnificent, huge, moving dunes can be seen in the Napeague-Promised Land area and on the western edge of Hither Hills State Park facing Napeague Harbor. Trees are very sparse and occur mainly in boggy depressions between the dunes. Mere exposed trees are almost always dwarfed into a "krummholz" form. Depressions and hollows between the dunes are termed "slacks" or "lows" by Salisbury (1952) for those with or without standing water, respectively. They have local conditions of high moisture and cool temperatures due to receiving runoff from surrounding dunes and persisting morning fogs coupled with cool air drainage and protection from drying wind action. Salisbury (1952) also points out that such areas may be rich in soil nutrients (as compared with surrounding dunes) due to leaching and drainage into the hollows of minerals and some organic matter. Dune grass (Ammophila breviligulata) is the most vigorous of the dune plants and is found throughout the area, with shrubs such as Myrica pennsylvanica, Prunus maritime, and Toxicodendron radicans growing mainly on the lee sides of dunes. Bearberry (Arctostaphylos uva-ursi) and false heather (Hudsonia tomentosa) are often conspicuous on more exposed areas between the dunes (Brodo, 1961a). Canard (1935), whose Ammophiletum breviligulatae, Hudsonietum tomentosi, Prunus maritime-Myrica carolinensis ('3: g. pennsylvanicg) association, and g;§23_rigida scrub association all fit into this vegetation type, noted the close similarity of this community to the dune communities of Europe. Martin (1959) describes this vegetation type, as it occurs in New Jersey, in detail (see especially his communities 1-2, 8-11, 15-18, 24-29, & 44). 2. Pine barrens (:: part of continuum segment A in Brodo, 1961a). Dominant trees: Piggg_rigida, Quercus albg, g, coccinea. Dominant undergrowth: Quergus ilicifolia, Gayiussacia baccata, vaccinium Isl-'- .m—w—v— -- -01 ..).. -—-———-— 72¢ 311cc :‘J.:;:':'.;1 v- ~ mic: f2: : 1'47? on :re 5 o ..x r.’-‘. \ 5-“.“ . .I I s. ‘Jg. Iasze:e?. ‘x q s "ri; 5%.!» ‘ o N I‘ s l3 aggggtifglium, X, vaccilans, Pteridium aquilinum. Soil: Dune sand or Colton and Adams sandy loam. Light: good to excellent. The wide expanses of pitch pine (Pinus rigida) and scrub oak (Quercus ilicifolia) which are characteristic of most of central Long Island have existed for centuries virtually unchanged. George washington wrote in his diary on the 22nd of April, 1790, a description of the area he saw as he rode from Patchogue to Coram and Setauket. He described the area as "too poor for cultivation being low scrubby oak, not more than two feet high, intermixed ‘with small and ill thriven pines" (Taylor, 1922). Canard (1935) states that this basic community extends from Newfoundland (where it is fragmentary) south to Georgia and Texas, with Pinus taeda and g, palustris replacing g, Eigig§_as the dominant. Both his Pinetum rigidae and Quercetum ilicifoliae communities can be placed here. 3. Pine-oak forest ( ==continuum segments A & B in Brodo, 1961a) (figure 11). Dominant trees: Quercus alba, Q, coccinea, Pinus rigida. Dominant undergrowth: as in pine barrens with Q, ilicifolia sparse except in glades. Soil: Bridgehampton sandy loam. Light: good. This vegetation type is little more than an older, more mature pine barren. The three dominant trees are the same in both but the order of abundance is different in the pine-oak forest with Quercus velutina making its appearance. The soil is better developed with more organic matter, although the ground vegetation is essentially the same. The trees are generally older, taller, and straighter. Sparrow and Hoodwell (1962) have presented a good description of this vegetation type in their description of a radiation study area at Brookhaven National Laboratory in central Long Island. The Quercetum velutinag_ as described by Conard (1935) belongs here and can also be applied to the scarlet-black oak woods discussed below. l4 4. Scarlet-black oak forest (::_continuum segment C in Brodo, 1961a) (figure.12). Dominant trees: Quercus coccinea, 9,,velutina, Q, alba, Dominant undergrowth: as in pine-oak forest. Soil: Bridgehampton sandy loam. Light: good. Again, we have a slightly older, more mature forest of basically the same structure as the previous vegetation types. Pinus rigida becomes relatively unimportant here with Quercus velutina becoming important. 5. Red oak forest (::.continuum segment D in Brodo, 1961a). Dominant trees: Quercus velutina, 9,52253, and locally, 9, pgiggg. Dominant undergrowth: Viburnum acerifolium, Smilax glauca, Vaccinium sp., Parthenosissus gginquefolia. Soil: Plymouth-Haven loam generally with a good humus accumulation, on glacial till. Light: fair to poor. The red oak forest extends all along the north shore and includes parts of the Sag Harbor region. It is this vegetation type which originally covered much of the New York City area and which was described by Harper (1917). Ground cover in the present stands is usually sparse except in some local spots where Smilax species and ggbgg_species grow in dense thickets. Conard's (1935) Quercetum kalmietosum and Quercetum prini both seem to fit best here. Where the soil is moist, Eggg§,begins to come in and replace the oaks (Canard, 1935). 6. Beech-oak forest. Dominant trees: Egggg.grandifolia, Quercus £3255, Aggg.rubrum. Dominant undergrowth: very sparse. Soil: Plymouth-Haven loam with much hums on till. Light: poor. A few small isolated areas near the eastern tip of Long Island bear remnants of some of the oldest vegetation on the North American east coast. These forests of old beech and oak trees can be found on Gardiner's Island, near antauk Point, and on Shelter Island (Taylor, 1923). ‘H . . Inw- a on. I' .— 15 7. 29322, Dominant trees: Eggngg.serotina, Amelanchier intermedia. Dominant undergrowth: Hyrica pennsylvanica, Prunus maritime. Dominant groundcover: Andropogon scoparius. Soil: Colton and Adams sandy loam. Light: unlimited. Norman Taylor (1923) wrote a detailed account of the grasslands of the antauk region. The area seems to have been a grassland devoid of any substantial forest cover for as long as we have records. Prunus serotina is the only conspicuous tree in the entire grassland area, and it is of very scattered occurrence. Amelanchier intermedia also occurs in a few groves. Shrubs are scattered throughout the area. Taylor (1923) stated that "wind is unquestionably the most important (factor) in maintaining the area as a grassland." This community is called the Andropogon scoparii in Conard (1935). 8. Hegpstead Plains grassland. Dominant tree: 252325 serotina. Dominant shrub: Hygigg.pennsylvanica. Dominant ground cover: Andropogon scoparius. Soil: Hempstead-Dridgehampton sandy loam. Light: unlimited. A great deal of work has been done on the vegetation of the Hempstead Plains (see pagell). It is considered by most workers to be a true "natural prairie,” i.e., a stable grassland community. The long stretches of Andropogon scoparius are only occasionally broken by isolated black cherry trees or bayberry bushes. Wind was probably not an important factor in the development of the prairie here, as it was with the very similar antauk downs, since Hempstead Plains, in central Nassau County, is not an especially windy area. Hicks (1892) claimed that excessive drainage plus the thinness of the surface soil and general climate determined the character of the flora of the plains. The soil is made up of tight, matted sod with sandy eroded areas occurring l6 wherever the sod had been broken. This dense sod, almost too hard to plow through and too dense to allow tree roots to penetrate, probably prevented subsequent forestation by local trees (Svenson, 1936). Conard (1935) called this community the Andropogon Hempsteadi. 9. figg_, Dominant trees: Chamaecyparis thyoides, Acer rubrum, Nyssa sylvatica. Dominant shrubs: vaccinium corymbosum, Toxicodendron vernix (L.) Kuntze. Dominant ground cover: Sphagnum spp., Vaccinium macrocarpon, y, oxycoccos, Woodwardia virginigg, Soil: wet acid sand grading into acid peat. Light: excellent to poor, depending on canopy development. White cedar swamps at one time were abundant all along the south shore at the heads of tidal streams and salt marshes (Harper, 1907; Nichols, 1907; Bichnell, 1908; Taylor, 1916), Heusser (1949), who presented the history of such an "estuarine bog" from.the nearby New Jersey coast, stated that rising sea level, ditching (with the subsequent influx of brackish water) and fires caused the disappearance of the cedars in that area. Similar conditions probably occurred on the Long Island coast. In addition, with the spread of suburbanization, almost all the cedars in Nassau County were harvested and most of the swamps filled inin order to provide space for the ever-extending highways. Although there are still some fragmentary estuarine bogs in the Babylon area, the best developed bogs are those farther east and inland which were formed in glacial depressions and are surrounded by pine or pine-oak forests (figure 13). In the Manorville region, some bogs were extensively cultivated for cranberries but few are still in use. The soil is very acid and provides good habitats for bog plants such as vaccinium.macrocarpa, 2, oxygoccos, Drosera spp., Lycopodium spp., Sarracenia pugpurea, and Utricularia spp. VJ 17 The white cedar swamps in various stages of development make up the Chgmaecyparetum.thyoidis, Chamaedaphnetum calyculatae, and vaccinietum corymbosi of Conard (1935). An otherwise similar community but without white cedar has been called the Aceretum rubri, and is discussed next. 10. Red maple swamp. Dominant trees: Acer rubrum, Nyssa sylvatica. Dominant shrubs: Clethra alnifolia, Viburnum dentatum, Vaccinium corymbosum. Ground cover: sparse; Osmunda sp., Sphagnum spp. Light: fair to poor. In wet areas not suited for white cedar, red maple swamps become established. They are common throughout the island. Cain and Penfound (1938) described and discussed this vegetation type in considerable detail, referring to it as the Aceretum rubri (including both the Aceretum rubri and the Aceretum osmundaceum of Conard [1935] ). It can easily be seen that vascular vegetation and soil type are strongly correlated (compare figures 1 & 2). The red oak forests are largely confined to the Plymouth-Haven soils, the pine-oak forests remain closely correlated with the Bridgehampton sandy loam, and the pine barrens are best developed on the Colton and Adams coarse sands. The Hempstead Plains grasslands are confined to the Hempstead-Bridgehampton soil association which is considered to have been formed under grassland vegetation (Cline, 1957). The dune and down vegetation of the south shore occurs largely on windblown dune sand. .“ m.- III. HABITAT ECOLOGY A. General Methods. 1. Collection date. Many important ecological notes on particular species were gleaned from label data of individual collections. For very rare species, these were often the only data available other than my field notes. On my own collections, substrate was noted as accurately as possible for each specimen (e.g., the phorophyte species in the case of corticolous lichens). If the phorophyte species could not be determined in the field, I collected a portion of a branch or twig with the lichen for later identification. Height above ground was noted along with other parameters if they were thought to be locally important, such as exposure in relation to a body of water, a road, or a farm. With each locality, the general light conditions were recorded as well as the dominant tree layer, shrub layer, and ground cover. Since collections are not made in an unbiased way, label data are of no ‘ use in statistical studies and only limited information can be gathered from this source concerning host specificity, vertical distribution, and so on. Label data are of greatest use in determining where a species can occur, i.e., the substrate potential, and never where it cannot occur, i.e., the substrate limits. Label data can indicate trends, and where the number of collections is large and the ecological limits small, certain conclusions can be drawn, although they should be considered tentative. 2. Statistical Studies. There are many ecological phenomena which can be studied adequately only through the use of unbiased sampling and statistical analyses. Questions pertaining to substrate specificity, the range and frequency of species in different wooded stands, the effects of New York City on lichen distribution, and the vertical zonation of corticolous lichens were 18 19 all approached statistically with the following methods. Lichen sampling was carried out in two areas, one in central Long Island in 1959, and one on the north shore in 1961. Different sampling methods were employed in the two studies, but since both involved unbiased samples of small areas, the data should be comparable. In the first case, eleven stands in central Long Island were sampled using a modification of the "random pairs" method of Cottam and Curtis (1949). The ‘method has been fully described in previous papers (Brodo, 1961a; Culberson, 1955a; Hale, 1955a). Briefly, the method consisted of selecting pairs of trees at prearranged intervals along a randomly selected transect line until 20 pairs (40 trees) were examined. On each tree, two quadrats were studied, one from.the ground level to a height of 30 cm, and another 40 cm high, centered at 1.3 meters (breast height). Each quadrat encircled the trunk. The stands sampled in the 1959 study ranged from pine barrens to red oak forests. The second sampling study was done in 1961 in the red oak forests along the north shore. Twelve stands were sampled along an east-west transect starting at Forest Park in New Yerk City and going eastward to Shoreham in central Long Island (figure 18). For the purpose of this study it was desirable to limit the survey to red oak stands of fairly uniform composition. Due to the uneven topography of the morainal north shore, vegetation appeared very patchy and areas of more or less uniform tree composition were small. For this reason the random pairs method used in the previous study was unsuitable since it covered too much territory and would have included too diverse a vegetation within each sample. Instead, a spiral sampling technique was employed. The system simply consisted of choosing a point in the center of the area to be sampled and *working in an ever increasing spiral, examining all encountered trees until 20 50 had been studied. This then is essentially a 100% sample of a very small area. The selection of the starting point in each stand was made to center specifically in the greatest concentration of red oaks regardless of the lichen population. This entirely non-random way of selecting the stands is valid since it is not the tree vegetation which is under study but rather the epiphytic vegetation of those trees. By selecting stands for a certain tree composition, the important variable of forest type is largely eliminated and the epiphytic vegetation within the stand can still be sampled in an unbiased manner. On each tree, two cylindrical quadrats, delimited exactly as in the 1959 study, were examined. Neither dead trees nor any that were less than 10 cm in diameter at breast height (dbh) or inclined at an angle of more than 10° ‘were considered. Data sheets were constructed to include about 25 common lichens all of which could be identified in the field without question. The species and dbh of each tree as well as the presence in each quadrat of any listed lichen were recorded. Cover was not noted but the direction of exposure of each species was recorded by noting its presence for each of eight compass points. The lichens, as a rule, were easily identified in the field with a hand lens, although, on occasion, chemical tests were performed on the thalli with potassium hydroxide, p-phenylenediamine, or hypochlorite solution for confirmation. The phorophyte species were often more difficult to determine, perhaps owing to the apparent wide occurrence of hybridization in the area among members of the black oak group (Quercus velutina, Q, Egbgg, and Q, coccinea). If the tree under study was judged to be a hybrid, the two putative parent species 'were listed in place of a single species name (e.g., Qgercus 52253_X coccinea). Previously (Brodo, 1961a), these three members of the black oak group were 21 considered collectively under the name of Quercus velutina. As will be pointed out later, the epiphytic lichen populations on the three species are very similar. 3. Transplant experiments. In an effort to clarify some of the ecological factors governing lichen distributions, some transplant experiments using corticolous lichens were carried out. The methods employed were fully described in a previous paper (Brodo, 1961b) but a brief account will be presented here including a few modifications and improvements which were used in the latest experiments. Using a steel punch (hereafter referred to as a "bark-borer") consisting of a hole-saw blade bevelled on the outside to a sharp cutting edge, and a holder (figure 22a,b), a bark disk bearing a portion of a lichen thallus could be removed from a tree with little injury to the lichen (figure 23). The disk could then be transferred to a hole made in the bark of any other or the same tree using the same bark-borer. The death of the inner tissues of the bark disk was found to have no noticeable effect on the attached thalli. The cut edges of the lichens themselves also showed no degeneration and, in the case of the control disks, continued to grow after transplantation. With continued use of the bark-borer, the blade tended to overlap at the point where the edges met (figure 22b). This resulted in uneven disk edges and occasionally prevented an easy removal of the disk from the tree. To prevent this, a wooden disk was made 3/8 inch in thickness, and cut so that it fitted snugly on the inside of the blade and against the holder. This disk effectively prevented the overlapping of the blade during the cutting operation and still left sufficient room inside so that no damage to the lichen thallus occurred. Two methods of fastening the disk into its new position were tried, both employing grafting wax as an adhesive. The first (Brodo, 1961b) was to apply? 22 the wax to the back of the disk, and the second was to apply the wax to the inside of the hole receiving the disk. Due to the much larger number of disks lost in the second year run, it is recommended that the former technique be used. The transplant experiments were used primarily to study vertical distribution and east-west distribution (New York City effects) and will be discussed further under those headings. In all cases, the lichens were examined at least twice after transplantation, first, after four months, and second, after one year. In addition to the general methods described above, certain special techniques and procedures are discussed in their appropriate sections below. Results of individual studies are also discussed within the sections. B. Substrate. Although lichen thalli have usually been considered as neither saprophytic 'nor parasitic, it has long been known that certain lichens are more or less restricted to certain substrate groups. Keys to crustose lichens almost always make use of substrate early in the separation of groups of species on a gross level, such as the choice between "horticolous" and "saxicolous." The degree of substrate specificity, particularly of corticolous lichens, has been the subject of several studies (Hale, 1955a; Culberson, 1955a; Barkman, 1958; Brodo, 1959, 1961a). In an earlier study of Long Island lichens (Brodo, 1961a), eight corticolous species were categorized according to their associations with each of three tree species, Pinus rigida, Quercus alba, and Q, velutina (including 9, coccinea and Q, 5223;) in four segments of the pine to oak forest continuum. various relationships were seen: a) significant positive association of the lichen with the tree species over the entire continuum, b) significant positive association in some segments, but not in all, c) no significant positive or negative association with the tree in any segment, d) significant negative association 23 in some segments but not in others, and e) significant negative association in all the continuum segments. The above relationships were interpreted as follows, respectively: a) the lichen shows constant substrate specificity indicating possible substrate requirements, b) the lichen shows some specificity for the tree but exhibits no clearcut requirement for it, c) the lichen shows considerable flexibility in substrate requirements, varying in degree of association with any particular tree species as the bark characteristics such as texture, chemistry, and moisture relations change in the different stands, d) the lichen shows some tolerance for the normally unfavorable substrate, but will occur more abundantly on other more favorable trees if they are available, and d) the lichen has some sort of physical or physio- logical inability to inhabit that substrate. The results of that study placed Parmeliopsis placorodia in category §_with respect to {$921 ri ida; Graphis scripts and Lecanora caesiorubella were in category 5 with Quercus velutina and in categories g_and g, respectively, with 92ercus alba. The other species, Parmelia caperata, P. rudecta, g. ub- aurifera, and 2, sulcata had little difference in their associations with the two oak species, although all showed greater tendencies toward positive association with black oak (possibly due to bark stability). Thus, caution is necessary in interpreting association tendencies since association values vary somewhat between stands and vegetation types (see Brodo, 1959). From field observations and collection data, a number of other lichens can be considered narrowly substrate-specific although in the absence of unbiased sampling, no quantitative statement can be made concerning them. Some of these species are listed below with their substrate placed in parentheses. 24 Corticolous: Alectoria nidulifera (Pipp§,£igigg), Cetraria fendleri (Pippg rigida), Leptorhaphis epidermidis (Betula populifolia), Lecidea anthracophila (Pinus rigida), Lecidea scalaris (Pinu§,rigida), Trypethelium virens (Ilex spp. and Faggs ggandifolia). Saxicolous: Caloplaca citrina (mortar and concrete), Q. feracissima (concrete), 9, flavovirescens (concrete), Candelaria aurella (concrete), 9, vitellina (granite), Lecanora disperse (concrete), Lecidea erratica (granite pebbles), Rhizocarpon obscuratum (granite), Rinodina oreina (granite), Sarcogyne clavus (granite). Terricolous: Baegpyges roseus (eroded sandy loam), Cladonia submitis (acid sand), Q. m (acid sand). Lignicolous: Chaenotheca pppeocephala (white cedar stumps), Lecidea aeruginosa (planks), Micarea prasina (rotting wood). The statistical studies cited on page 22 attempted.to clarify the basic factors involved in specificities by relying on the correlation of lichen presence with measureable bark characters. Some of these characters are listed and discussed below. 1. Texture: The external texture of the substrate can be important in trapping diaspores and protecting developing thalli, in providing entrance to other layers or tissues of the substrate, in capturing and retaining moisture and chemical substances, and in supporting other organisms which may remove potential lichen sites, or aid in any of the above. Different parts of a tree trunk may have different textures and consequently may bear entirely different lichen floras. For example, in fissured bark the rough fissures often bear hygrophytic species such as Lepraria incana whereas adjoining plates support only hardy species such as the alga Protococcus viridis 25 (see Barkman, 1958, p. 33). LeBlanc (1962) cites bark moisture capacity as causing the differences between the rich lichen flora of red oaks and the poor flora of beeches. It is probable, however, that the conspicuous differences in bark texture between the two trees were important in producing the different epiphytic vegetations. Rough boulders normally bear more lichens than smooth ones, and some lichens undoubtedly have adapted to growing on very smooth surface in response to the competitive advantage of such an ability (e.g., Lecidea erratica, Rhisocappon obscuratum, Verrucaria microspora, and 2, silicicola). 2. Moisture-holding capacities: Many studies dealing with.epiphytic vegetation, particularly cryptogamic vegetation, have included moisture capacity measurements of bark substrates (Billings and Drew, 1938; Young, 1938; Hale, 1955a; Culberson, 1955a; Barkman, 1958; Brodo, 1959; LeBlanc, 1962). Barkman (1958) has reviewed this subject in detail. Although the methods employed by the various workers varied somewhat, in general, it was found that moisture capacities are greatest with soft flaky barks, near the tree bases, on windward sides of tree trunks, and in humid areas. Except in a few cases, moisture capacity was expressed as the ratio of water absorbed to dry weight of the sample. LeBlanc pointed out that by using dry weight in the expression, barks with the same actual moisture capacity per unit of exposed surface may appear to have different moisture capacities if their densities are difference. For example, bark sample g; with a surface area of 10 cm2 and weighing 10 grams may absorb 5 grams of water when submerged. Sample 2, also with a surface area of 10 cm? (and of the same volume) but weighing 20 grams may also absorb 5 grams of water. Since sample §_is twice as dense (and weighs twice as much) as sample A, it appears to have only half the moisture capacity, when in reality, the 26 capacities are equal. LeBlanc (1962) attempted to correct for this error by expressing water gain on a "per unit surface area" basis. Unfortunately, it is extremely difficult to accurately measure surface area with any but the smoothest of bark types and serious errors may thus be introduced into expressions derived in this way. Barkman (1958) stated that moisture capacity is more meaningful if presented in terms of sample volume. This would be excellent to compare barks which are known to become totally and uniformly saturated with water. However, if only surface'layers are wetted, as might well be the case with some of the hard-barked oaks, moisture capacity per unit volume is unusable. The measurement of the rate at which a given bark sample returns to dry weight after being saturated (either by vapor or by liquid water) for any given period of time is,under uniform conditions of humidity, a direct function of its surface area, moisture capacity, and water binding capacity, all parameters of importance to epiphytes. Hale (1955a) and Billings & Drew (1938) presented some data on water loss and found that bark samples returned to approximately dry weight in about the same time for all trees studied, but that the initial rates were greatest in tree barks having the highest moisture capacity (expressed as a per cent dry weight)., Although these figures are important, they still do not reveal exactly how much water remains available to the epiphyte during this water loss. That is, a bark which absorbs four grams of water per unit surface area may lose water twice as fast, during the first hour, as a bark which absorbs one gram of water per unit area, but at the end of that hour, the former still contains twice as much water as the latter and in a less strongly bound and hence more available state. Thus, it would seem that water loss rates alone could not be used as a substitute for some sort of a moisture capacity expression. ‘— L.—“-——-‘4—‘ 27 There seems to be no way to entirely overcome the density or the surface area problems. It is possible to introduce a correction factor into the dry weight expression to eliminate the density error, but the result is an expression in terms of volume and as stated above the samples would then have to be of the same volume or be proven to become uniformly saturated with water. The following example will illustrate this point. salaries Swag Dry weight: 5 gm 3 gm Volume: 10 cm3 10 cm3 Density: 0.5 gm/cm3 0.3 gm/cm3 Water absorbed: 6 gm 6 gm Moisture capacity: 6 gm/ 5 gm - 1.20 6 gm/ 3 gm - 2.00 Density correction: m.c. X d - 1.20 X 0.5-9pgp m.c. X d - 2.00 X 0.3sQLQQ These final values represent grams water absorbed per unit volume, ... and are functions of moisture capacity which are comparable and give relative positions of the bark types. The principle is valid, but is replete with difficult problems some of which were mentioned above. In addition, sample volumes are almost impossible to keep constant since some bark types are thin (e.g., Egggp grandifolia, A£g£.rubrum) and others must be taken in Ehick slices (e.g., 92ercus £3255, ngpp.americana). The measurement of volume is somewhat inaccurate in certain bark types since bark samples often contain spaces which trap air in water displacement procedures. Inaccurate volume determinations, of course, make density figures of little practical value. Errors in volume measurement also decrease the value of moisture capacity expressions based on volume alone. . Despite the shortcomings of some of the methods discussed above, moisture 28 capacity measurements were performed on bark samples from a number of common Long Island trees. The methods employed were essentially those of Culberson (1955). Bark samples were obtained with the use of the bark-borer used in the transplant studies (see pages 21-22) wherever possible. This method provided samples of very similar size and volume except for the thin bark trees. Some bark types were not amenable to bark disk removal due to their instability and flaky nature (e.g., Quercus alba and Pinus rigida). Bark samples of these trees were collected without a borer and were cut down to approximate the surface area of the disks. The samples remained unstudied for a year and a half and so were quite dry at the beginning of the observations. They were oven dried at 1000 C. for a period of 20 hours to ensure uniform desiccation. After cooling, the samples 'were weighed, then coated with a layer of paraffin on all cut surfaces and then reweighed to derive the weight of the wax. The volume of the wax on each sample was calculated with a knowledge of the wax's density. Volume was measured by water displacement and was precise to 0.3 cubic centimeters but volume measurements were somewhat exaggerated in certain bark types having large amounts of air retention, e.g. Pinus rigida bark. The exposed area was measured as follows: a small piece of aluminum foil was carefully fitted to the contours of a bark sample. The excess foil was then cut off at the limit of the exposed colonisable surface. The fitted foil piece was pressed flat, numbered, and weighed. The weights of the various foil replicates were then fitted on a standard curve constructed from the weights of foil samples of known surface area to find the surface areas of th£2 bark samples. In this way, very irregular, rough_surface features of the bar1( samples could be accounted for in the surface area measurements. 29 Water absorption expressed as per unit dry weight, per unit volume, and per unit of colonizable surface is presented in table 3 along with the other bark features of the common Long Island trees. In ranking the trees in order of bark moisture capacity, we can see that the method of expression is very important in the relative positions of the various species. Dry weight and volume expressions matched most closely, with Pippa rigida being a notable exception. Volume measurements of pine bark are complicated by considerable air retention between the bark plates during water displacement as noted above. This error would make the volume appear larger than it actually is and would thus effectively "lower" the moisture capacity expression based on volume. Quercus rubra appears more mesic than Eggpg_in the surface area expression, whereas the opposite is true with the dry weight and volume expressions. LeBlanc (1962) noted the same change in relative position of the two trees in his studies. One important difference in the area, dry weight, and volume sequences is in the relative position of Quercus 3123; it appears relatively more xeric in the former and more mesic in the latter. In view of the strong emphasis which has been placed on the difference in moisture capacities between black and white oaks in the past (see Hale, 1955a), it may be well to recheck these findings in other areas with larger samples. The sample size (1-8) in the data presented here was too small to warrant the formulation of strong conclusions pertaining to the relative positions of various trees with regard to their bark types. 3. Stability: The rapidity with which a given substrate surface is removed or changed in some way has a strong influence on the lichens that can inhabit the surface. Only rapidly growing and maturing species can become established on unstable surfaces. 30 No species can colonize shifting send as is found on sand dunes. Dune species usually become established on relatively less active dunes on plant remains (Brodo, 1961a) or under the protection of trailing or low growing vascular species such as Arctostaphylos uva-ursi or Hudsonia tomentosa. The thalli may later become detached and continue development independently on the relatively stable sand surface. Where the sand is protected from strong wind action and becomes covered with an organic film as in scrub oak thickets, certain species such as Lecidea uliginosa, L. guadricolor, or Cladonia cristatella can become established and actually servein binding the sand particles together (see page 46). Where the sand is even more stabilized, many more terrestrial species may gain foothold. Baeomyces roseus and Pycnothelia papillaria can apparently grow fast enough to grow over eroding surfaces. Rapidly sloughing bark severely limits the number of species which can inhabit a tree (Hale, 1952a; Barkman, 1958), and it is likely that this is one of the reasons for the relatively small number of species found on 21225 rigid . The best development of any species growing on pine occurs on the edges of the bark plates deep in the fissures where the bark is most stable. The poorest development is on the plate surfaces which lose outer flakes of bark almost continuously. The role of bark stability in limiting species coverage is made strikingly clear when a dead standing trunk of pine appears close to a living tree. The stable bark of a dead tree is covered ‘with lichens whereas only spotty coverage is seen on the living bole, even though both trunks have equal light and are standing side by side. It is possible that the absence of a canopy may have some effect in changing the moisture relations (via increasing drainage) on the dead tree or in failing to contribute inhibitory organic material, but these are probably not as important as the stabilized substrate. 31 Pebbles and small stones often shift and roll with changes in weather, and thereby expose or cover lichens which may be growing on their surfaces. I first considered this problem in a study of the lichens of an old field at the American Museum of Natural History Biological Laboratory at Dix Hills, Long Island. Yearly observations of numerous pebbles indicated that Lecidea erratica develops very quickly on exposed stones (see pages 47~48). Since these pebbles and stones undoubtedly shift or even turn over with frost action and heavy rains, rapid growth may be an important factor in the maintenance of populations of these species. Typical members of the exposed boulder communities such as Parmelia arseneana and g, conspersa have been found on some stabilized pebbles adding strength to the supposition that the instability of pebbles may be a factor in eliminating these over- shadowing but slowly growing species from competing for space with the small but rapidly developing Lecideae. Small stones continuously roll and shift in the littoral zone of the shallow bays and inlets, and it is not surprising to find that the marine verrucariae (1, microspora and 1, silicicola) often are found growing on all sides of these pebbles regardless of their position when collected. 4. Chemical Composition: Bark chemistry as with bark moisture capacity has been studied by most epiphyte ecologists. Barkman (1958) again provides an excellent summary of the information published on the subject. 0f the many facets of bark chemistry, acidity has been the most widely studied. Great emphasis has been placed on bark acidity in explaining the distribution of some lichens (Billings and Drew, 1938; Hale, 1955a; Culberson, 1955a; Barkman, 1958; DuRietz, 1945 in Almborn, 1948). Barkman (1958) and Almborn (1948) have pointed out some of the oversimplifications to which some 32 authors have fallen victim, but pH remains an important factor to be considered in epiphytic ecology. The pH of bark samples of several of the common trees were measured. A few grams of bark material were obtained by slicing the surface layers from a bark sample and chopping them into a mealy consistency. Between five and seven ml of distilled water were added to each chopped bark sample, enough to form a thick slurry, and the mixture was allowed to equilibrate at room temperature (approximately five hours). Acidity was measured using a glass electrode Beckman pH meter. The acidity of soil samples was umasured in a similar way. A soil slurry was formed using one part water and two parts soil (approximately 20 cc soil and 10 ml distilled water). The mixture was allowed to equilibrate (15 minutes) and pH was measured using the same apparatus as mentioned above. The results of these measurements are given in table 3a. It would seem that acidity either affects the lichen vegetation directly or indirectly, or reflects a condition which does, because definite correlations can be seen between lichen presence and substrate pH. The very low pH of Pinus rigida bark could explain its poor and restricted flora, and the high pH of 2123;, and Robinia provide clear associations with the so-called "nitro- phytic" (Xanthorion) comunity. It is especially significant that a black oak once found bearing Xanthoria thalli had neutral bark although this species of tree normally has very acid bark. That particular oak was growing in the center: of a large Long Island duck farm, the atmosphere of which was very obviously filled with ammonia and other gaseous and fine particulate materials. Trees along farm roads exposed to farm dust have long been known to bear rich “honiophilous” communities (Barkman, 1958; Almborn, 1948). The very high moisture capacity of Ulmus may be a significant factor in the specificity of 33 roadside species, although the oak mentioned above which supported a rich Xanthoria community had a low moisture capacity comparable with other oaks. The problem of separating nitrogen concentration from acidity in correlations of this kind has been discussed by both Almborn (1948) and Barkman (1958). Both authors point to the possibilities of other factors being involved, especially phosphorous concentrations. For example such typically "nitro- philous” species as Caloplacg cerina, g, pyracea and g, flavovirescens are are also found on turtle shell and bone, substrates known to be high in phosphorus. Since calcium concentrations are often high in alkaline substrates, calcium may be important in these specificities as well. Many so-called “hitrophilous” lichens, especially Xpnthoria parietina, X, £31155 and Physcia adscendens are commonly found on mortar and concrete which have a high pH and calcium concentration, but are certainly’not rich in nitrogen compounds. The presence of Cladonia submitis and associated lichens on the south shore and inland and their absence on the north shore is strongly correlated with soil acidity. The south shore and inland sands are all distinctly acid, whereas the north shore sands are neutral (table 4). Exactly what is involved in this correlation is still not clear (see page 264). The eroding soil supporting Baeomyces roseus has the same pH as the dune sand and therefore acidity cannot explain the differences in the terrestrial communities of the two soil types. The higher moisture capacity and organic content of the eroded sandy loam possibly are the deciding factors in this case. Other substrate minerals not studied here are undoubtedly important in lichen distributions. Although some data are available on the mineral contents of substrates (Barkman, 1958) and mineral nutrition of lichens (Smith, 1960a etc.) the subject is still far from adequately understood. 34 The extreme specificity of Trypethglium virens for several species of I135 (Johnson, 1959) suggests the presence of some genetically controlled metabolite in that genus which is essential for the establishment or survival of the lichen. Fagus grandifolia, another common phorophyte for Trypegheliu! virens would then also have to possess the ability to produce this substance or a substitute. There is some evidence that Tgypethelium actually does utilize some bark material (Johnson, 1940). Fink (1913) suggested that other endophloedal crustose species also derive some nutritional benefit from their substrate. The fact that Trypethelium virens has been found in healthy condition on Long Island only on living trees, an observation also made by Johnson (1940), and the fact that all the host trees of this species have thin living bark is added weight to the possibility that a specific class of metabolite is involved. It is easy to imagine a lichen living on a nutrient-rich substrate making use of these nutrients, especially when all the mechanisms for their absorption are available and efficient (Smith, 1962). More work on substrate specificity is needed to clear up these important problems. C. Climate. Atmospheric humidity is involved in the water budget of a lichen thallus to a greater degree than it is in the water budget of a rooted vascular plant in the same general habitat. This is due to a lichen's ability to pick up water vapor and use the absorbed moisture in photosynthesis and metabolism in a relatively short period of time, as compared with the green parts of vascular plants (see page 36). Thus a habitat which might be dry for a terrestrial vascular plant due to excessive soil drainage, may not be dry to lichens if air humidity was high enough during part of the 24 hour 35 cycle. It is the microclimate which one must measure in order to characterize the water budget in the ecological niche of a lichen. To perform such measurements was unfortunately beyond the scope of this work although such studies would be extremely interesting and valuable. Vertical and horizontal zonation (see pages 38-43), and patterned distribution in bark fissures or on bark ridges, are probably at least partly manifestations of different microclimates. l. Illumination and Temperature: Light intensity is a very complex factor having both direct and indirect effects on microclimate. As Barkman (1958, page 57) states "... it is often difficult to decide whether a given species is photophilous, thermophilous, or xerophilous," since strong light 'will raise the temperatures of both bark surfaces and the lichens themselves (especially if they are dark colored), and will, therefore increase the evaporation rate increasing drought conditions. The role of illumination in raising temperatures and thus evaporation rates and drought was an important consideration in Barkman's (1958) summary of the causes of horizontal zonation (zonation according to direction of exposure) of epiphytes on tree trunks in Holland. Lichens derive their principle nutrition from the photosynthetic products of their algal components, and so the lichen thallus is dependent upon light for survival. Since, in the lichens that have been studied, the net rates of photosynthesis per unit surface area are much lower than those of the leaves of higher plants (Smith, 1962), it is not surprising that most lichens are fbund in moderately or well lighted habitats. Deeply shaded forests, dry or moist, are, in general, lichen poor. Lichens exposed to full sunlight, however, are often subject to extreme drought. Many species have developed adaptations such as cortical pigment accumulation and cortical thickenings 36 (Barkman, 1958) which cut down light intensity and transpiration. Some Long Island lichens which seem to be distinctly photophilous are Cetraria islandica subsp. crispa, Cladonia submitis, g, bogyi, Xanthoria parietina, Egggelia sulcata, Usnea strigosa, Ramalina fastigiata, Pertusaria xanthodes, and Lecanora caesiorubella subsp. lathamii. These species are most often found in well illuminated habitats even though their general substrate types extend into more shaded areas. The first three species mentioned above are found almost exclusively on exposed sand plains and downs. Xanthoria parietina has long been known to be photophilous (Barkman, 1958). The remaining species occur most frequently in well lighted but dry, mature pine-oak forests of central Long Island. This vegetation type can be thought of as a compromise habitat between optimum light and optimum moisture (Brodo, 1961a). In more humid localities such as the Montauk region on the southern fluke of Long Island (see figure 5) these species all reach their maximum development in completely exposed situations. 2. Moisture: Lichens are classically thought of as among the most drought-resistant plant types. Although it is true that many species can survive in habitats much too dry to support any but the most xeric of bryophytes, many lichens are clearly limited to rather moist environments and many others are very sensitive to changes in environmental moisture. The role of moisture in the photosynthetic efficiency of lichens has been reviewed and summarized by Smith (1962). He points out that in the nonaquatic lichens which have been studied, photosynthetic efficiency is greatest at moisture contents below saturation. Nonaquatic lichens rarely are saturated in nature. Although most nonsorediose lichens absorb liquid water rapidly, they lose water almost as fast. Absorption of water vapor is a much slower process, but constantly humid areas are undoubtedly less droughty than dry, 37 exposed areas with frequent rains (Barkman, 1958). This is especially the case since it has been shown that lichens can absorb water from nonsaturated air (Pavillard, 1939; Barkman, 1958). Thus, the misty thickets and shrubby groves of the depressions in the Montauk area are wet habitats whereas just outside these groves on the exposed dunes where constant strong winds make evaporation high, the habitat is extremely dry (Taylor, 1923; see also Salisbury, 1952). Ried (1960) pointed out that lichens are most seriously damaged when they are subjected to intermittent wet and dry periods, which may explain why some lichens thought to be drought resistant through laboratory experimentation actually appear sensitive to low moisture conditions when observed in the field. He suggested that it is the ability of various species to recover from a drought which might determine the distribution of certain species. Moisture also has an important indirect influence on lichen growth. Inasmuch as microbial activity is highly dependent on moisture levels of various habitats, any lichen distribution dependent on the products of fungal or bacterial growth or on the changes in the physical characters of substrates subjected to such activity would necessarily follow moisture changes as well. Moisture comes to the corticolous lichen thallus from precipitation and from air humidity (both directly and through the wetted substratum) and rarely by inundation. In the tropics, moisture may be made available directly from the living tissue of the thin barked trees (Imshaug, personal communicatiOn)° The evaporation rate in any particular habitat and the moisture capacity of the substrate determines how efficiently and for how long this moisture is available to the lichen. The availability of rain to epiphytic lichens is influenced by canopy type and canopy density mainly through their effect on the flow of waker froul the leaves and twigs down the branchlets and branches and finally down the 38 trunk. This flow of water (”stemflow") is often a major route for the entrance of moisture to the forest interior (Kittredge, 1948) and is of course of major importance to corticolous plants. Stemflow is greatest with trees having ascending branches ("centripetal crown") as in figs; and 33325, and is least with trees having drooping branches ("centrifugal crown") as in gigggj 92ercus and giggg are intermediate in this respect (Barkman, 1958; Geiger, 1965). It should also be borne in mind that precipitation which has passed through a canopy ("throughfall") is much richer in certain minerals and ions than unintercepted rain (Tamm, 1951). In the pine-oak forest of Long Island, much of the rain reaches the bole directly through the loose canopy as well as by stemflow. In the dense red oak forest, light rains never reach the ground or tree trunks, being evaporated directly from the canopy. Heavy rains filter down through the canopy but only reach the hole via rain tracks (the channels of most liquid stem runoff) and general stemflow. However, once the rain has wet the ground and bark in a shaded forest, the precipitation is slowly converted to increased air humidity which slows evaporation from the wetted thalli and supplies additional moisture for a long period. The rain in a pine-oak forest, on the other hand, is quickly lost in the very well drained sandy soils and dried from the bark with no substantial increase in the local humidity for more than a very short period of time. It is therefore in the relatively open habitats that hygrophilous species occupy substrates with high moisture capacities (see pages 59-52), D. Verticgl Distribution. The vertical zonation of corticolous epiphytes has intrigued many cryptogamic ecologists (Plitt, 1924; Billings and Drew, 1938; Hale, 1952a; 39 Culberson, 1955a; Barkman, 1958; Brodo, 1959, 1961a, 1961b). Methods of study varied from detailed investigations of a few trees from base to crown (Plitt, 1924; Hale, 1952a) to studies of hundreds of trees only at basal and breast height quadrats (Hale, 1955a; Culberson, 1955a; Brodo, 1961a). Barkman (1958) made numerous observations concerning vertical zonation and thoroughly reviewed the previous work. Several approaches were taken in the study of this phenomenon on Long Island: a statistical evaluation of species presence in breast height and basal quadrats, experimental transplant studies, and field observations of lichen communities. As a result of the statistical investigations described previously, several common species could be characterized as to their vertical zonation affinities (table 5 and Brodo, 1961a). From an examination of the vertical distribution of certain common corticolous lichens in the pine-oak forests as compared with the red oak forests (disregarding phorophyte species) (table 5), one can see that the frequencies in the basal quadrat in the former are consistently higher than those in the latter. This tendency of species normally dwelling at breast height to be confined to the basal area in dry pine oak woods is consistent with the statements made by several authors (Billings and Drew, 1938; Plitt, 1924; Potzger, 1939; Barkman, 1958) concerning vertical microclimatic gradients. That is, bark moisture is greater, and evaporation is slower at tree bases as opposed to microhabitats higher on the trunk. Barkman (1958) has pointed out how different moisture conditions in different vegetation types can influence epiphytic vertical distributions. He states that in moist woods, typically base-dwelling communities sometimes cover entire trunks. 0n Long Island, this phenomenon is particularly striking with bog tree epiphytes. Within the humid, cool 40 bogs, Lgbg;1§_pu1monaria and Lobaria guercizans grow at all levels, but just outside the bogs, in the drier oak forests, the same lichens are confined to tree bases. Transplant experiments done in 1960-1961 dealing with vertical distribution of Lecanora caesiorubella and glgdonia chlorophaea (Brodo, 1961b) showed that the Lecanora could survive when transplanted from breast height to the tree base but that the Cladonia, upon being transferred from the base to breast height (1.3m), soon decayed. Lichen frequency data bear out the supposition that the Lecanora is somewhat more facultative in its vertical distribution than is the Cladonia (see table 5). Since the lichens were transferred on their original intact substrate to points on the same tree, the experiments also indicated strongly that it is microclimatic conditions rather than bark surface features or differences in organic or inorganic nutrients on a vertical gradient which largely determine where on a particular tree a lichen can survive. Since the degree of fungal-bacterial breakdown of bark appears to increase towards the tree base it is possible that the microclimatic gradient may be operating through a biological link to influence the lichen. In various local habitats not sampled in the statistical work, some noteworthy types of vertical zoning were observed. On the windward sides of trees growing close to bays and lakes, basal lichen communities often extend far up the trunk (see also Billings and Drew, 1938; Barkman, 1958). For example, Parmelia rudecta, Parmelia caperata, and Physcia orbicularis, all dominantly base-dwelling under normal conditions, were found growing high on the bay-facing side of an oak tree on Shelter Island. The lee side of the trunk had a normal basal zone. Inclination of the phorophyte trunk greatly changes its moisture conditions 41 and permits basal vegetation to grow much farther towards the crown (Barkman, 1958). A tree growing on a steep hillside essentially has the ground brought closer to the crown on the uphill side of the trunk, and this side, then, has a more "basal" epiphytic flora. Crustose species are almost entirely confined to areas above the base except for the normally basal epibryic and leprose crusts. Although light has classically been cited as one of the main causes of vertical zonation of epiphytes (Plitt and Pessin, 1924; Barkman, 1958; Hale, 1952s, 1955s; etc.), light probably was not significantly involved in the results of the Long Island studies since illumination does not seem to be a controlling factor in either forest type at the basal or 1.3 m levels. In pine-oak forests light appears to be abundant and uniform over most of the tree due to the low, loose canopy, and in the shaded red oak forest, light appears to be uniformly low until one reaches the upper portions of the trunk and canopy far above the level examined. 0n the few felled or wind-blown trees that were examined, an obvious trend toward a greater number of crustose species at the tree tops and a greater lichen cover in general points to a light effect. Photophilous Q§g§§.strigosa is most abundant in forest glades and forest edges, as well as on tree tops pointing to light as being the controlling force in its vertical distribution. Moisture, of course, is of major importance in all types of lichen distributions (see pages 36-38). Vertical moisture gradients of many kinds have been reported including evaporation rate (Plitt and Pessin, 1924; Potzger, 1939), bark moisture (Billings and Drew, 1938; Hale, 1952a) and relative humidity (Barkman, 1958). With the ground being a major water reservoir it is evident that the farther one moves away from this reservoir, the drier the microclimate will be. The more humid an area is, the less will be the 42 difference between humidity at ground level and humidity at greater heights and therefore, the less pronounced will be the vertical vegetational zonation which responds to this moisture gradient (see also Barkman, 1958, p. 39). This is indeed what is observed in the Long Island studies and we can therefore conclude that moisture is probably a controlling factor in most cases. Temperature as well as bark characters such as color, hardness, and porosity all have their effect on substrate moisture relations via evaporation rate, or moisture capacities, and all show vertical changes. Since epiphytes are sensitive to moisture changes, it is easily seen how a vertical zonation of epiphytes can be influenced by these physical features of the substrate. Organic and inorganic nutrients, either having been blown on to the bark surface, carried down by stemflow or throughfall (Tamm, 1951), or produced there by local microbial activity, are distributed along a vertical gradient and may play an important part in the distribution of certain species, particularly those species that normally grow on the ground. The possible role of nutrient accumulation in the maintenance of established colonies of Cladonia cthaea has been disproven by transplant experiments (Brodo, 1961b), but its possible importance in the establishment of certain species cannot be eliminated. Since virtually no work has been done on the factors involved in the establishment of different species in nature, little can be said about this important aspect of lichen ecology at this time. Certain bark characters such as hardness and rate of exfoliation have selective effects on certain lichens and certainly cause some vertical lichen zonation (see also pages 24-25; 30). Crustose species tend to be most abundant on the smooth young bark at the top of the tree, possibly responding just as much to the physical bark feature itself as to the increased light at those levels. Hale (1950, 1952a) discusses the importance of bark texture 43 in the maintenance of certain types of lichens according to their anchorage abilities. Some species have greater abilities to reinvade exfoliating bark than others and would cause vertical zonation along a bark age gradient. One then might view a single tree trunk as demonstrating all stages of a corticolous succession with all stages in time frozen at different levels of the trunk. E. Succession. Both directional and nondirectional changes in species composition were seen within certain lichen communities (see Hanson and Churchill, 1961, for a fairly detailed discussion of ecological changes of different kinds). The tracing method used in the growth rate studies (see Appendix A) provided a means for demonstrating the fluctuations in local lichen populations and the constant change in composition and coverage of lichen communities. Reports of no change in lichen communities in up to 50 years (A. L. Smith, 1921; Cooper, 1928) are to be viewed with some skepticism in the absence of precise measurements (as pointed out by Smith, 1962). Figure 16 presents one of the many examples of fluctuations which were observed. Here the thalli of Parmelia sulcata are shown to grow at one point while in other places they fall away and allow the invasion and extension of Physcia millegrana. When populations change in a directional fashion, succession can be said to be taking place (Hanson and Churchill, 1961). The changes described below may be truly successional or may be the first stages of a cyclic fluctuation. These changes were observed in the growth rate tracings of a community on Quercus glbg.in a moderately lighted oak woods (figure 17). The quadrat was at a height of one meter and was facing away from the prevailing wind directicu1. 44 1959: Parmelia sulcata - all thalli healthy, robust. ghyscia millegrana - all thalli healthy and vigorous; many very small thalli present. Lecanora caesiorubella - one thallus, vigorous with many large apothecia. 1961: Parmelia sulcata - some thalli showing evidence of decay; most healthy. Physcia millegrana - all thalli healthy, vigorous. Leggnora caesiorubella - vigorous; apothecia unchanged. Dynamics: a. nggnora caesiorubella is being encroached upon and covered on all sides by ghysci§_mi11egrang, although both appear to be healthy. vb. Wherever Parmelia sulcata and Physcia millegrana are both healthy and are growing adjacent to one another, the Parmelia is growing over the Physcia with one exception in a very local . area of {M 5313559. thallus. c. Wherever Parmelia sulcata appears to be dying, the Physcia is growing over the Parmelia. d. Small regeneration lobes can also be seen in the dying areas of Parmelia sulcata. 1962: Parmelia sulcata - most thalli showing considerable decay. Physcia millegrana - healthy, vigorous. Lecanora caesiorubella - one half the thallus whitened and decaying, Dynamics: . Physcia mdllegrana was encroaching considerably on the Lecanora. 45 Succession occurs in response to a change in the environment of the site without a change in regional climate or a change in the organism. The rate is dependent on the organism's rate of growth and the environment's rate of change (Barkman, 1958). The sequence of the successional stages depends on the characteristics of the participating organisms, often both physical and chemical. Billings and Drew (1938) described a microsuccession of epiphytic bryophytes due to the aging of the bark substrate. Not only does bark change in time due to the tree's own activity, but as Barkman (1958) points out, the epiphytes themselves alter the bark moisture capacity, and acidity. The forest of which the tree is a part changes in time also, especially with reference to light and humidity. With both corticolous and saxicolous lichens, the successional sequence is usually thought of as crustose to foliose to fruiticose and/or bryophytes, i.e., according to the growth form, although Rudolph (1953) has pointed out many exceptions to this scheme. The sequence described from the growth-rate tracings indeed fits into the more standard pattern. The one deviation involves poorly developed or depauperate specimens, in which case the succession may start to reverse. Physcia millegrana is normally overshadowed by Parmelia sulcata except when the latter is in poor health, at which time the Physcia will overgrow the Parmelia. In areas recently subjected to air pollution, fruticose and foliose lichens are usually more severely damaged than crustose species, and succession can therefore be reversed, i.e.)fruticose to foliose to leprose species. Barkman (1958) described such a reversal in Holland with Lobaria giving way to mosses which finally yield to Pleurococcu§_ as one approaches an industrial center. This is thought to be due to the fact that the freer an organism is from its substrate surface, the more surface area it exposes to the polluted air and the more susceptible it is 46 to air pollution. Thus, with increasing air toxicity, the larger foliose and fruticose lichens will be the first to go, then the smaller foliose species, and finally the leprose crusts and algae. In New York City, therefore, the algal communities covering the trees in the city parks can be considered to be in a disclimax stage maintained by air pollution. Successional stages on boulders have not been studied on Long Island. On the inland sand habitats, succession often proceeds from crusts such as Lecidea uliginosa (n: L. guadricolor to Cladonia spp. and finally to grasses and shrubs. 0n the windswept sand dunes of the south shore,however, the reverse is often observed. Dune grass (Ammophila breviligulata) gains a foothold, and upon its death and decay leaves a "stump" onto which some species of Cladonia, especially Q, 2231;, can become established. The clumps of Cladonia will close over the sand more or less stabilizing the surface and will decompose and provide substrate for other plants (Brodo, 1961a). A similar but more nondirectional cyclic change was described by Watt (1947) working with a very similar community: Callung,vulg§gi§, Arctostaphylos uva-ursi, and Cladonia sylvatica (. Cladonia arbuscula). In his scheme, the Cladonia stage can give way to bare soil again upon which the phanerogams will become reestablished. It is very possible that the same cyclic development may occur on the Long Island sand dunes, but I have not recorded any observations to that effect. Alvin (1960) in discussing lichens of an Ammophila-Calluna dune comlnnity (see page 73) considers sand stability and pH as prime factors in this succession with reproductive potential (soredia production) as possibly also important. On eroding sandy loam in the inland portions of the island, a type of cyclic change can be seen. VBaeomyces Eggggg_and/or Pycnothelia papillaria are the pioneers, effectively binding the soil particles together providing a 47 situation suitable for the invasion of many other species of Cladonia (particularly 9. strepsilis, Q. subcariosa, and g, clavulifera) which are followed by grasses and herbs. If the ground is disturbed and subjected to new erosion, the cycle will begin again. The role of lichens in a segment of old field succession was studied by me over a period of three years at the American Mhseum of Natural History's Kalbfleisch Field Research Station on Long Island. Even in that comparatively short period, some interesting trends were observed. The field under study (AP-5) was abandoned in 1954 and so was six years old when these observations were first made. In 1960, the phanerogamic vegetation consisted of a mixture of annual and perennial weeds with Andropogon becoming abundant by the third year of the observations. In general, the trends were as follows: Vasculgr Plants: Non-vascular Plants: Weeds sparse; l. Pebbles bare; soil exposed around weeds except light excellent for some clumps of moss. 2. Pebbles covered with pycnidia of Lecidea erratica; some non-podetiate Cladonia squamules appear on moss clumps and to a lesser extent on the bare soil. \/ 3. Pebbles covered with pycnidia and small apothecia Grass becoming important; of Lecidea erratica; podetial initials seen on light diffuse Cladonia thalli in moss clumps and, to a lesser extent, on the bare soil. 4. Pebbles covered with a mixture of pycnidia and mature apothecia of Lecidea erratica; podetia of Cladonia cristatella and Q, chlorophaea well developed in moss clumps and on soil. 48 5. Rapid decline of all species, especially Cladonia spp. Grass heavy; light very poor 6. Disappearance of Lecidea erratica from pebbles. Portions of this succession were seen in various quadrats and the entire successional picture is actually a composite of the many segments observed. Robinson (1959) in a paper dealing with old field succession in North Carolina, also noted the importance of Cladonia cristatella and g, ggayi ( O Q, chlorophaea) in the 6-9 year old stage. He stated that the lichens attain their greatest dominance after the decomposition of much of the moss and grass vegetation. If the same sequence follows on Long Island, the observations at the Kalbfleisch Station could represent a minor primary succession within the more over-all successional pattern which could only be seen over a longer period of time. Evans and Dahl (1955) noted that the most conspicuous lichen cover was in old field communities of mosses and perennial weeds ("Bryoid - Antennaria types”) although some species including Cladonia cristatella and g, pyxidata attained importance in the ”Poe - Aristida” community. The Bryoid - Antennaria community is well lighted, and becomes established on dry, unstable soil whereas the Poa - Aristida community is slightly more shaded and is found on more stable soil. Because of the common occurrence of ground fires in the pine areas of central Long Island, succession on burned ground and bark was studied in several areas. The types of pioneers on burned over barrens depend on the extent of the fire. If small areas, even very limited, are left unburned, a large number of species may be available for reinvasion. Fire can get very close to a lichen colony and not destroy it. In southern New Jersey near Tuckerton, I studied an area which was burned over not more than two years 49 previously. The fire swept through the area charring almost the entire ground surface as well as many tree trunks. The fire apparently was windswept and very rapid because on the lee sides of many trees, charred bark extended to a height of about four feet, whereas on the windward sides of the same trees, many lichens appeared unharmed. On the soil, a similar situation was seen. Tiny areas untouched by the fire supported healthy colonies of several Cladonia species, particularly Cladonia uncialis and g. subtenuis although the fire had devastated areas only a few feet away. Small moss clumps, especially of Leucobrygm spp. or Polytrichum spp., seemed to provide protection for small lichen thallus fragments, and some reinvasion of the surrounding area probably originated from these clumps. With a rich source of nearby species, succession seems to be rather haphazard with regard to pioneers, and is mainly dependent on which lichens have the best means of dispersal. Charred ground is soon covered by dust and then wind-blown soil and other plants, and is recolonized soon after the fire has left. Charred bark, however, remains uncolonized for a long period except by certain specialized species. Succession on an area almost totally destroyed by fire gives a better indication of a natural succession because invasion, with few exceptions, must occur frmm outside and the species ”selection” is much greater. I studied such an area in Yaphank, Long Island adjoining the Suffolk County Firematic Training Center. The fire had totally destroyed the ground cover and charred the ground over an area of about 50 acres or more. The trunks of pines were burned to a height of 10 to 12 feet and the oaks were charred to a lower height. Two similar areas were studied and yielded similar observations. One was in Centereach and the other near Selden. Prior to the last burn all the areas were pine-oak barrens of approximately the same age. 50 In all the areas, I noted that lichens invaded the soil before they invaded the bark of burned trees. The uncontested pioneer was the ubiquitous glgdonig_cristatella followed closely by g. bacillaris and Q, chlorophaea. All three species are extremely common on the island and all have very wide substrate tolerances. All three species are found on tree bark of many types, soil of many types, and even stones and gravel if they are present. Cladonia caespiticia was observed as an associated pioneer in the Yaphank and Centereach areas as well as on charred ground in two other incompletely burned areas. Sterile thalli of Lecidea uliginosa covered small patches of sand in the Selden locality. No reinvasion of the charred oak bark was seen, but the burned pine bark supported a number of species of crustose lichens. Lecanora subintricata, a very minute, athalline crustose species, was collected on a completely charred and almost destroyed pine; Lecidea anthracophila was on moderately charred bark near the edge of the burn. Other species such as Lecidea scalaris and the foliose Parmehopsis placorodig_were on unburned bark just above charred material and probably were remains of a preexisting population rather than a reinvading one. However, Lecidea scalaris, and sometimes .L. anthracophila,have been collected on charred bark on numerous occasions and undoubtedly can reinvade recently burned over forests. I did not study the long term effects on the terrestrial lichen flora after frequent burning. Buell and Cantlon (1953), however, observed an increase in the lichen cover with burning frequency over a period of years. Johnsen (1959) reported a slight increase in lichen cover with periodic burning, but declined from making a firm statement pending more cdmplete data. Both the above studies were made in pine forests, the first in the pine-oak region of New Jersey, and the second in a pure stand of,loblolly pine (gingg taeda) in the North Carolina piedmont. 51 F. Species gggposition within hgbitats. It has long been known to field botanists that certain plants tend to be found in association with certain other plants. It therefore soon became convenient to refer to these groups of species collectively as "communities" or "unions.” With the growth of the field of phytosociology, hundreds of plant communities were examined, analyzed, and named. It is my opinion that the use of latin epithets in naming biological communities of any kind implies an intricate predictable organization which does not exist. The principles underlying this opinion as they apply to lichen communities are listed below. 1. Each local lichen population has definite ecological requirements (i.e., a specific niche);:br certain species these requirements are narrow, and for others they are bread. 2. Lichens with similar gross ecological requirements will tend to be found together more frequently than lichens with dissimilar ecological require- ments. The more similar the gross requirements, the more frequently the occurrence of the two together. Since, according to the "ecological exclusion principle,” no two species having exactly the same niche requirements can exist together, as the ecological requirements of two lichen populations approach identity, a higher and higher degree of competition will develop between them. One species, once present in a habitat, could successfully exclude an ecologically similar species by: a) extremely rapid growth (preempting suitable available space), or b) chemically or physically altering the habitat preventing the establishment of the ecologically similar species. This, of course, could only be effective if the establishment requirements of a species were different from its survival requirements. Barkman (1958, p. 197) believes that the production of a growth inhibitor by Qpegrapha gg§i2_ and Q. cinerea prevent the two from occurring side by side although they seem 52 to have the same habitat requirements. 3. Any particular area or locality has a limited set of potential species available for colonization due to that locality's particular ecological and phytogeographic position. For example, a white cedar bog on Long Island does not have all the lichens in the world available for colonization (excluding the slim possibility of chance long distance dispersal). Only those species whose distributions include Long Island, which had means to arrive at Long Island (as via coastal plain swamps), and which require or can tolerate a high moisture, low light, cool temperature environment such as is found in these bogs, could occur there. Thus out of approximately 16,000 species known to science, we are left with about 30 which may be found in a Long Island bog. On any particular tree in the bog, such as a Chamaecyparis, the list is cut down even more (eliminating lichens which must grow on the ground, rotting wood and vegetation or smooth bark, etc) leaving only a dozen potentials. The chances are very high that some or all of these potential species will be on white cedars in that bog, their diaspores having arrived there and distributed themselves in a relatively random fashion. 4. Since the conditions in any particular habitat are not static, mainly due to changes in the substrate itself with time and age and to local changes in microclimatic conditions, neither is the community composition in these habitats static. Succession does not always occur with the same sequence of species or at the same rate. This often results in the establish- ment of mosaic communities of mixed development making any phytosociological classification extremely difficult. 5. The composition of lichen communities varies from one habitat to another in an unbroken continuum along physical, chemical, or microclimatic 53 gradients. Almborn (1948) cites an excellent example of such a continuum following an illumination gradient. The lichens involved were members of a community on Egggs. The concept of "community" as used here should not be confused with the integrated biological system, consisting of lichens, bryophytes, micro~ organisms, vascular plants, and animals, of which it is a part. Strictly speaking one can even think of each lichen thallus as a sort of "community" ... an intimate, highly integrated association of algae and a fungus. In conclusion then, we can consider a "lichen community" to be a group of species having similar gross ecological requirements and occupying a certain habitat together. This group of species is subject to directiohal and non-directional change with time resulting in a compositional continuum from one group to another. If we decide that lichen communities should not receive latin names, the problem of how to deal with communities still remains. One can "classify" a community according to its floristic composition, as do most phytosociologists, and merely refrain from giving it a specific name, or one can delimit communities according to their ecological affinities by classifying their habitats. The latter method is employed in this work. Each method has its advantages and disadvantages and a final choice depends mainly on the use of the ultimate product. Barkman (1958) in discussing epiphytic communities ably outlines the advantages of using floristic criteria for vegetational analysis. He states that phorophytes cannot be used alone since ”1. the kind of host tree is not of direct influence, 2. its significance varies from one region to another, 3. other ecological factors are thus ignored and, last but not least, 4. any logical system should be classified upon the characters of the objects to be classified, ig_casu 54 upon the vegetation itself...." His arguments are well taken and phorophytes alone are not used in the system outlined below. However, in my opinion 1) phorophytes as well as other lichen substrates often seem to have a distinct influence on their lichen vegetation as is evidenced by the number of species which are wholly or partially substrate specific (see pages 22-24), 2) the changes in substrates of certain communities from one area to another often give important clues pertaining to the causes of accompanying changes in vegetation, 3) ecological factors other than substrate can easily be included if needed, and 4) since lichen communities themselves are basically "unnatural," i.e., they are only fragments of true biological communities, the method of community classification one uses should depend on convenience and usefulness. The chief advantage of the classification of habitats over the floristic method described by Barkman is that the former does not eliminate any vegetational combination and therefore makes possible the classification of a total flora into communities; in the floristic method, representative associations are selected from the total flora, leaving many vegetational combinations not considered. The first major division of the habitat classification which follows is by the various vegetation types (in their broadest concepts). The next division is by substrate; first the general substrate type is considered and then any other narrower substrate classification that seems pertinent. Occasionally a sdcroclimatic division is made beyond that of the substrate. Under each habitat are listed, in approximate order of importance, lichens which have a high probability of being found in that ecological situation. These species comprise the ”lichen community.” One must keep in mind that the species lists represent potential communities and not actual ones. Rarely» will all of the species in any particular community occur together. 55 Bark characters for all the corticolous community phorophytes are sunmarized in table 3. To aid the reader in locating specific habitats, an outline of the habitat types precedes the discussions. 1. Upland Habitats A. Corticolous l. Pinus rigida 2. Qgercus alba 3- 9.92.1322 4. Q, velutina group 5. Egggg grandifolia 6-.A_c_e_t£u_l>au_a 7. Elm g americana B. Saxicolous 1. Mortar and concrete 2. Granite boulders 3. Pebbles and small stones C. Terricolous l. Mossy soil 2. Sandy soil D. Lignum 1. Stable, dry lignum 2. Unstable, highly decomposed lignum II. Bog and Swamp Habitats A. Corticolous l. Chamaecyparis thyoides Z-Assame 3. Ilex verticillata 56 B. Terricolous C. Lignicolous III. Maritime Habitats A. Aerohaline stratum (salt mist zone) 1. Corticolous a. Myrica pennsylvanica - Prunus maritima b. Prunus serotina c. Juniperus Virginians d. Ilex opaca 2. Saxicolous a. Concrete and mortar b. Granite boulders 3. Terricolous a. Stabilized sand b. Dune sand 4. Lignicolous B. Hygrohaline stratum (salt spray and storm tide zone) C. Hydrohaline stratum (littoral zone) I. UPLAND HABITATS A. Corticolous l. giggg,£igigg.(pitch pine). Species: (a) base - Cladonia bacillaris, g, incrassata, Q. cristate118, (b) breast height - Parmeliopsis placorodia, g, aleurites, Lecidea anthracophila, L, scalaris, Bgcidi§_chlorococcg. Comments: The best lichen development occurs on the edges of the bark plates, not on their surface (see page 30). A A4— __... L 57 Species in the basal community, especially Cladonia incrassata and 9, W are often found only on strongly decayed wood and on pine bases. The reasons may lie in the fact that both substrates are very acid (Barkman, 1958, p. 113) and usually moist. Pine needles and bark flakes often cluster at the bases of pines forming thick piles of material which retain moisture long after all other material is dry. Thus, pine bases have a particularly high local himidity. Pine bark species found at breast height are usually narrowly confined to pine alone, at least on Long Island, although pine-dwelling species which are found in bogs as well as pine forests often are collected on thggecyparis thyoides or even vaccinium corymbosum. 2. Quercus alba (white oak). Species: (a) base - Parmelia caperata, Physcia orbicularis, Cladonia coniocraea. (b) breast height - gggmelia caperata, g, rudecta, g, saxatilis, Physcig’orbiculggig, Parmelia subaurifera, Physcia millegrana. Comments: The relatively high moisture capacity and.low acidity of the bark of 99ercus 5122 renders it a unique habitat in the black oak and pine filled forests of central Long Island (see also Hale, 1955a). However, its lichen vegetation does not vary much from that of black oaks with a few important exceptions, notably among the Physciae which are rather common on white oaks and rare on black oaks. Distinctions between these two oaks are further developed under the discussion of the black oak group. 'LeBlanc's Parmelia caperata, g, rudecta, g, saxatilis, Physcia millegrana, and Eh. orbicularis unions (LeBlanc, 1963) resemble the Long Island white oak bark community at different points in the continuum of lichen composition. 58 3. Quercus 251325 (chestnut oak). Species: (a) base - ngdgnia coniocraea, ngmelia rudecta. (b) breast height - Parmelia sulcata, g, rudecta, g, caperata. Comments: The very hard, impervious bark of Quercus prinus makes it a rigorous habitat for all but the most xeric of species, especially above the base. Its relationship with the lichen vegetation of other trees of the red oak forest will be discussed under the Quercus velutina group. 4. Qggrcus velutina group (black oak group) including 9. velutina (black oak), Q, coccinea (scarlet oak), Q. £2253 (red oak), and all hybrids, especially 9, coccinea g rubra. Species: (a) base - glgdonig_coniocr§g§, g, chlorophaea, Parmelia caperata, g. rudecta, g, saxatilis. (b) breast height (partial shade) - Parmelia sulcata, g, rudecta, Bacidi§_chlorococca, Graphis scripts, Lecanora caesiorubella subsp. lathamii, Parmelia caperata, 2, saxatilis. (c) breast height (light good) - Parmelia sulcata, Lecanora caesiorubella subsp. lathamii, Pertusaria Eggghodes, Parmelia subaurifera, Lecanora chlarotera, Parmelia ggggtilis, Usneg_strigos§. Comments: Comparisons of the epiphytic lichen vegetation of members of the black oak group indicate that all species support very similar communities. Even the lichen vegetation on 92ercus glhg_bears many resemblances to that of members of the black oak group. Using data from the 1961 transect study of the red oak forests on the north shore, the epiphytic lichen communities of the principle tree species were compared. Only those on relatively common trees could be compared statistically. Kulcsinski's coefficient of community proved to be the most useful statistical tool. Only stands 7 through 12 (Sunken Meadow to ShorehauO 59 were used since those stands west of Sunken Mbadow were considered under the influence of the New York City atmospheric conditions. Where it seemed valuable and pertinent, the results of the red oak forests were compared with those of the pine-oak forests derived from data collected in 1959 in connection with the study of central Long Island (Brodo, 1961a). Continuum segments A and B taken together are considered as "pine-oak forest" for the purposes of these comparisons. It is possible to compare the epiphytic lichen floras of a number of trees using lichen frequencies at 1.3 meters and at the base as did Culberson (1955a), or disregarding vertical position. The latter was done for the lichen communities on oaks in the red oak forest. The communities were then arranged in a sequence with the most similar closest together and those most dissimilar farthest apart. Vertical distribution was disregarded in this case since there were few differences between the basal and 1.3 meter vegetations of the trees, and the additional lichen species introduced by the combined values aided in the computations. The matrix of coefficient values with the ranked communities is presented in table 1. The most striking aspect of the matrix is how high and how similar the values are. In Culberson's comparisons, the values ranged from 8 to 76 with only one pair of tree species having a coefficient over 70. In the Long Island study, all oaks, particularly Qggrcus veluting, Q, coccinea, and Q. coccinea X rubra were very similar in their epiphytic vegetation. Only 92ercus pgigg§_was distinctly apart from the others. Even Quercus 2123 with its soft porous bark was almost indistinguishable in epiphytic vegetation from the black oaks. These results agree well with what one finds in the actual red oak forest .. a rather monotonous and sparse epiphytic flora throughout the stands 60 regardless of the phorophyte. However, in the pine-oak forest, a field worker is struck by a subtle but distinct difference between the white oak and black oak epiphytic communities. Analyzing the coefficient of association of Q, gl§§_and Q, velutina communities in the pine-oak forest as was done with the communities on oaks in the red oak forest, we arrive at a figure of 78, which indicates they are similar in their epiphytic flora. Considering the basal and breast height lichen frequencies separately, the coefficient of association values are 74 and 75 respectively, still not reflecting any differences between the trees. Using only the presence of the number of lichen species unweighted by the number of trees examined rather than frequency, one arrives at a totally different and much more realistic picture (table 2b). The reader may come to suppose that this is merely a technique of statistical juggling to find results which fit preconceived notions. However, the policy of looking for a statistical means of revealing some more or less apparent ecological phenomenon actually can throw a great deal of light on the real factors involved in this phenomenon. This is a good case in point. An unweighted species presence analysis of the breast height vegetation of Quercus velutina and Q, alba reveals that the two are not at all similar whereas their basal communities are very similar. One can see that a few common and very frequent species can far outweigh a larger number of rarer, constant, and somewhat substrate specific species. For example, three species of Physcia and two of Pertusaria were found only on Q, glh§_and never on Q. velutina. But even if unweighted species presence is accepted as being a more instructive test, it cannot be used in comparing two trees of different frequencies unless a test is made to establish the extent to which the sample 61 size is affecting the number of epiphytic species observed. For example, if 50 white oaks and 25 black oaks were examined, one would normally expect many more epiphytic species on the former if, in a sample of 25 trees, the total lichen flora on that tree species is only barely represented. If it can be shown, however, that in this example, essentially all the epiphytic species on black oak were examined after a sample of 20 trees, and that the same was true of white oak, the two trees can be compared without regard for the sample size. Species-sample curves were constructed for each of the tree species and their lichen vegetation (both basal and breast height) in the red oak and pine-oak forests, and it was shown that with each tree species, the sample size (40-90 trees) was sufficiently large to allow direct comparisons with other trees. Coefficients of association were then calculated for Quercus velutina and Q, glba_using the basal vegetation and the breast height vegetation. After testing sample size and species number for the oaks in the red oak forests, Q, velutina (including Q, coccinea, as in the pine-oak forest), 9, 52255, and g,_gl§§.were found to be comparable on an unweighted species- presence basis (table 2). Two very interesting things can be seen from the table of coefficients. First, the breast height communities of Quercus 3123 and Q, velutina show greater difference than their basal communities. Secondly, this difference appears umch greaterzh the pine-oak forest than in the red oak forest. Concerning the first point, it should be noted that the bark of 93ercus glhg_is unlike that of Q, velutina in several respects, the most obvious being hardness, moisture capacity, and color. Bark hardness was not measured as was done by Culberson (l955a),but white oak bark is easily flaked off and gouged with a fingernail, and the bark of Q, velutina is 62 sometimes difficult to cut into even with a sharpened steel knife. The average moisture capacity of Q, glh§_was found to be 69% (dry weight), 411 (volume), and 141 (area), and that of Q, velutina was found to be 38% (dry weight), 30% (volume), and 17% (area) (see pages 25-29 for a discussion of moisture capacity measurements). Hale (1955a) emphasized the differences in bark characters, particularly the moisture capacities, between 9, albg_and Q. velutina. Color is more important than would first be suspected. Heat absorption and thus, indirectly, evaporation rate must be influenced by bark color. All these factors add up to a characterization of black oak bark as a very dry habitat (hard, low moisture capacity, high evaporation rate) and white oak as a comparatively moist habitat (soft, high moisture capacity, low evaporation rate). Many authors (Barkman, 1958; Billings and Drew, 1938; Young, 1938; Brodo, 1959) have noted the great importance of the physical characteristics of bark on the distribution of epiphytes. This difference in moisture relations between Q, yglgging_and Q; glbg_would naturally be less important at the tree base where the microclimate is normally humid (Barkman, 1958; Billings and Drew, 1938), than at breast height where microclimate is variable and usually drier. The larger number of species on Quercus alba may thus be related to the wetter microhabitat which can support a greater number of drought-sensitive species. This difference between 9, velutina and Q, 5123 disappears in red oak forests which are more moist and more shaded than pine-oak forests (Brodo, 1961a) and where the vertical zonation is in general not as distinct as in drier habitats. It should also be noted that many of the species which differentiated the two lichen communities in the well-lighted pine-oak forest were photophilous species and therefore were absent in the shaded 63 red oak forest. The lichen community found on Eggyg_spp., not included in the statistical studies, is very similar to that of the younger, smoother parts of the oak trunks, especially when well-lighted communities are compared. Depending on the state of development and position in the community continuum, (as reflected in the local dominant species), the following "unions" of LeBlanc (1963) can be referred to the community found on black oak on Long Island: Bacidia chlorococg§_union, Graphis scripta union, Parmelia rudecta, g, saxatilis and P, sulcata unions. 5. £532; grandifolia (American beech). Species: Trypethelium virens, Pyrenula nitida, Buellia curtisii, Graphis scripts, Phaeographis dendritica. Comments: The hard smooth bark of Fagus grandifolia effectively limits the lichen community growing on it to crustose species, and the dense, shade-producing canopy restricts the community even further. 6. Age; rubrum (red maple). Species: gypggygnig_physodes, ngmelia perforata, Pertusaria trachythallina, Parmelia subaurifera, Leganora chlarotera, L, caesiorubella subsp. lathamii. Comments: The community on maple in upland habitats closely resembles that of other smooth, hard barked trees, especially Quercus coccinea. gypggygnigtphysodes, however, is more abundant on maples than on oaks in oak forests. 7. giggg_americana (American elm). Species: Xanthoria parietina, g, fallax, Physcia millegrana, 2h, stellaris. Comments: Elms, especially as they occur along roadsides, have been 64 studied a great deal in relation to their lichen flora. The neutral pH of the bark no doubt has a direct or indirect effect upon the epiphytic flora because other neutral barked trees (e.g., Populus deltoides, Robinia pseudoacacia) have very similar floras. In the Braun-Blanquet system, this community would be included in the fignthorion pgrietinae Alliance (Barkman, 1958). In LeBlanc's treatment of Canadian epiphytic communities, (LeBlanc, 1963) the Lgnthoria fallax union would most closely apply here. B. Saxicolous. 1. Mortar and concrete. pH 7 - 10.8 Species: (a) Full sun - leoplaca feracissima, Lecanora disperse. (b) Partial or full shade - Caloplaca flavovirescens, Placynthium ni rum. Comments: Mortar and concrete being highly alkaline and calcareous are equivalent to limestone in their general characters and in their lichen vegetation. There is no natural occurrence of limestone on Long Island. 2. Granite boulders. Acidic, coarsely crystaline, very hard. Species: (a) Full sun - Rinodina oreina, Lecanora cinerea, Parmelia arseneana, L, conspersa, L, stenophylla, Sarcggyne clavus. (b)Phrtial or full shade - ngideg.glbocaerulescens, Lepraria sonata, Parmelia caperata, Lecanora cinereg, Buellia stigmaea. Comments: There is some species overlap on the granite communities of well illuminated and poorly illumdnated boulders, but a few species are absolutely restricted to one or the other (e.g., Rinodina oreina in the former and Lecidea albocaerulescens in the latter community.) 3. Pebbles and small stones. Usually smooth, but not always; high in quartz. Found in fields, on roadbanks, or in other open areas. Species: Lecidea erratica, L, coarctata, L, cyrtidia, AcarosEora fuscata, Rhizocarpon obscuratum, 65 Comments: Why these species develop on pebbles and not on boulders of similar hardness and chemistry is hard to determine. It is possible that the high mineral supply (derived from seepage and splashing from surrounding soil) or higher humidity (due to close proximity to ground) are involved. The lack of stability of small stones and pebbles is also probably a factor in limiting the kinds of lichens which can survive in this community (see page 31). C. Terricolous. 1. Mossy soil. Gravelly, but relatively rich in organic matter; in oak woods of various development; pH not measured. Species: glpdgnis subtenuis, Q, caespiticia, Q. cristatella, g. bacillaris, Q, pleurota, Q, chlorophaea, Q, furcata. Comments: This community is best developed in forest glades, or on moss-covered abandoned roads. It is almost always at least partially shaded. 2. Sandy soil. Little to no organic matter; pH 4.1 - 4.6. Species: (a) unstable eroded sandy loam (roadbanks, fire breaks, etc.; subsoil or very fine sandy loam) - Baeomyces roseus, Pycnothelia papillaria, glpdonia strepsilis. (b) More or less stable but bare sandy loam - slightly more sandy - Cladonia strepsilis, Q, subcariosa, Q. clavulifera, Q, atlantica, g. chlorophaea, Q, pleurota. (c) Very sandy soil - Lecidea uliginosa, ‘L. quadricolor, glgdonig_cristatellg, Q, macilenta, g, atlantica, Q, uncialis, Cetraria islandica. (d) Dune sand - shifting, sometimes grass-covered - Cladonia cristatella, Q, 2951;, 93 uncialis, Q, submitis, g, chlorophaea, Q, furcata (especially f. racemosa), Cetrarig.islandicg_subsp. crisps. 66 Comments: The sandy-soil communities as outlined above are largely arbitrary units derived from a continuum formed along a soil gradient from comparatively rich sandy loam to almost pure quartz dune sand. Although a few species are more or less confined to one unit (e.g. Baegpyces roseus and Pyonothelia papillaria), most terricolous species can be found throughout most of the continuum. D. Lignum. 1. Unstable, dry lignum (fences, planks, decorticate logs, decorticate branches and twigs). Species: Lecidea aeruginosa, L, botr osa, QQQdonia cristatella, g, bacillaris, Lecidea myriocarpoides, Bacidia chlorococca. Comments: This community occurs in both partially shaded and sunny habitats. A very frequent member of this community is the imperfect fungus Sirodesmium pezizoideum‘ Cooke 6: Ellis) Mason & Hughes. As logs and planks become heavily decomposed, the community composition changes giving rise to the community listed below. 2. Unstable, highly decomposed lignum (rotting logs and stumps). Species: Cladonia parasitica, Q, incrassata, g. bacillaris, Micarea prasina. Comments: The characters of high acidity and moisture capacity seen in rotting wood have much in common with the bases of Pinus rigida and the similarities in community composition are obvious (see page 56). The community is best developed in swamps and bogs where wood decays quickly, but it also occurs in oak forests on heavily decomposed stumps. 67 II. BOG AND SWAMP COMMUNITIES A. Corticolous l. Chamaecyparis thyoides (swamp white cedar) - Vaccinium corypbosum (highbush blueberry). Species: Parmelia hypotropa, Parmeliopsis ambigua, g. aleurites, Cetraria viridis, g, ciliaris, Hypogymnia physodes, Usnea trichodea, g.subfusca sensu Metyka. Comments: The corticolous lichen communities on these two woody bog plants are remarkably similar, especially in view of the fact that one is a conifer. Several members of the bog lichen community occur exclusively in bogs and only on these substrates (e.g., Cetraria viridis, Parmeliopsis ambigua, and Usne§.trighodea). The relationship between the bog habitat and the bog lichen flora will be discussed later. Several lichens commonly found on pine in pine forests are found on white cedar in bogs (Lecidea anthracophila, Ochrolechia parella, Parmeliopsis aleurites). Cetraria ciliaris normally found on white cedars was collected from Eggplg populifolia in two different maple swamps. The close similarity of L252L3_bark to that of conifer bark was noted by Barkman (1958), Skye (1958), and others. 2. gar; m (red maple). Species: Lobaria pulmonaria, L, guercizans, Parmelia rudecta, P, caperata, Pertusaria 52353, chidia chlorococca. Comments: The smooth hard bark of App; pppppp undoubtedly has a great effect on its epiphytic lichen vegetation. The maple community is almost totally different from that on bog trees and shrubs with looser more absorbent bark. 68 3. Ilex verticillata (black alder). Species: Trypetheliug.virens, Graphis scripts, Lecanora caesiorubella subsp. lathamii, Pertusaria pustulata. Comments: The affinities of this community to that of LL35 ppggg have already been mentioned. The presence of the other species mentioned may well be due to the better light conditions in Ilex verticillata thickets. The dense shade produced by the canopy of LL g gpggg_exclude all but the most shade tolerant of species. B. Terricolous (acid boggy sand, edges of bogs). Species: Cladonig calycantha, Q, atlantica. Comments: This community also is well developed on dry acid sand, especially as found in pine barrens. C. Lignicolous (rotting logs). Species: Cladonia parasitica, Q, incrassata, g, gigypg, g. vulcanica, Q, santensis, g, beaumontii. Comments: The community on rotting wood in bogs is basically identical with that of drier forests except for the occasional presence of the four rare species mentioned last. A marked geographic difference was seen in the community composition in disjunct localities of southern New Jersey and Cape Cod. The dominant species in the New Jersey bog-lignum community was Cladonia santensis which covered large areas of dead wood and cedar stumps. glgdonia vulcanica was not collected there at all. On Cape Cod, glgdgnia vulcanica was clearly dominant and Q, santensis was not collected. The lignum community in Long Island bogs showed neither Q, vulcanica nor 9, santensis as dominants; both are in fact very rare on the island. Instead Cladonia incrassata 69 and Q, parasitica, both common throughout the northeastern coastal plain, were most conspicuous. III. MARITIME HABITATS. Because maritime communities are so heavily influenced by their proximity to salt water, it is useful to classify them on the basis of their salt water exposure. Des Abbayes (1934) presented a detailed discussion of the zonation at the shoreline. Following Du Rietz (1925a in des Abbayes, 1934), he recognized three major divisions, 1) the aerohaline stratum, which is strictly terrestrial, receiving salt only as a fine mist suspended in the air and is never wet with salt water, 2) the hygrohaline stratum which receives salt water directly as salt spray or by immersion at very high tides or at the high spring tides, and 3) the hydrohaline stratum which is submerged with every high tide regardless of the season. Boyce (1954) and Oosting and Billings (1942) measured the salt spray concentrations at various distances from the mean tide. The latter authors found that salt spray is greatest on the exposed side of the fore dune, less at the hind dune summits, still less at the lee side of the fore dune and least at the lee side of the hind dune. Boyce reported salt concentrations of up to 2.2 mg. salt/sq.dm/ hr. at a distance of 270 m from mean tide with a wind speed of 11 m/sec. His data show that salt concentrations closely depend on wind speeds as well as distance from the salt source. 0n Long Island, wind speeds of 11 m/ sec. are very common (see page 8) and so one can safely say that the aerohaline zone extends at least 270 m from the water, and probably much beyond. Des Abbayes subdivided the hygrohaline stratum into three "echelons" based on the presence of certain indicator species. Since none of his 70 indicator species, except for Verrucaria microspora, are present on Long Island, only the three major strata will be used in this community classification. A. Aerohaline stratum (salt mist zone). 1. Comments: Corticolous. a. Myricg_pennsy1vanica (bayberry) - Prunus maritime (beach plum). Species: (a) Exposed to full wind and salt-mist (foredunes, bluff tops, beaches): (1) base - Parmelia sulcata, g, hypotropa; (2) breast height - Rinodina pillisris, Lecidea varians, Parmelia hypotropa. (b) Protected from full wind and salt mist (lee side of dune, groves of trees): (1) base ~ Parmelia sulcata, z, livida; (2) breast height - Parmelia hypotropa, L, perforata, Lgcidea varians, Ramalina fastigiata, Usngg strigosa. All the species listed are photophilous with high drought and salt resistance. It is evident, however, that wherever moisture is greatest in the dunes areas, the lichen vegetation is most luxuriant. This type of community is best seen in the Montauk Point area and behind the moving dunes at Promised Land. The lichen communities listed above occur on many shrubs along the shores, and almost unchanged on many of the dune and beach trees. Comments: (See discussions of black cherry and red cedar below.) b. Pppppp_serotins (wild black cherry). Species: Parmelia sulcata, P, subaurifera, Buellia curtisii, L, stillingiana, Pertusaria xanthodes, Lecidea varians, Usnea strigosa. This community has many similarities with the shrub communities, and differs chiefly in the inclusion of several additional photophilous crusts. 71 c. Juniperus Virginians (red cedar). Species: Physcia gillegrana, {,0rbicularis, Ramalina willeyi, Eggmelia hypotropa. Comments: This community is surprisingly "nitrophytic" (see page 32) perhaps from the neutralizing effects of salt mist (see Barkman, 1958). The absence of conspicuous crustose species is perhaps due to the instability of the substrate. d. ng§_22§g§_(American holly). Species: Trypethelium virens, Phaeographis dendritica. Comments: This community is very similar to that on Fagus grandifolia and llg§,verticillata which are the only other substrates for Trypethelium virens. The most striking similarity between the trees is that all three possess a very thin, hard outer bark with a living layer just beneath. This factor alone could not be the decisive one in determining the distribution of Trypethelium, however, since many other trees and shrubs have this character also (e.g., Acer rubrum and Amelanchier intermedia). 2. Saxicolous. a. Concrete and mortar. Species: Xanthoria parietina, Caloplaca citrina. Comments: Verrucaria muralis, X, nigrescens and Rinodina salina occur as rare members of the community having only been found at Orient Point. Xanthoria parietina and Caloplaca citrina are common aerohaline species although both are also widely distributed far from salt water (see pages 33% 339). Lecanora disperse and Candelariella aurella are also found in the aerohaline stratum as facultative members of the community. The former is listed by des Abbayes (1934) as a typical member of the aerohaline stage community. Alvin (1961) noted Catillaria chalybeia, Rinodina demissa 72 ( - g, salina), Lecanora disperse, ganthoria parietina and Candelariella vitellina (ecologically equivalent to Q, aurella?) as comprising a community found on the bricks of a sea wall on the east coast of England. This community is remarkably similar to the one on Long Island except for the absence of Caloplaca. b. Granite boulders. Comments: No lichens were seen which were at all confined to the aerohaline granitic rocks, although several species normally found farther inland were found growing in the salt spray zone. Parmelia caperata and Acarospora fuscata are conspicuous species in this category. Parmelia caperata was listed by des Abbayes (1934) as common in the aerohaline stage. 3. Terricolous. a. Stabilized sand (as on Orient Point). Species: Cladonia pygidata, g, strepsilis. Consents: On Long Island, Cladonia pgidata has only been collected on stabilized beach sand in the salt spray zone. It is interesting from the standpoint of the possible salt-preference of this species, that I have seen it growing in luxuriant abundance on beach sand on the shore of Lake Erie (Point Pelee, Ontario). Cladonia strepsilis is clearly a facultative member of the aerohaline community. b. Dune sand (as on Fire Island and at Napeague Beach) (figure 9:). Species: Cladonia submitis, Q, bggzi, Q, uncialis, Q, cristatella, Cetraria islandica, Cladonia_chlorophaea. Comments: ,Since the community on dune sand extends essentially unchanged into inland localities, salt spray can be eliminated as important in defining its distribution. It is possible, in fact, that heavy salt spray such as would occur on an exposed fore dune may inhibit the community's 73 development (see pages 264-265). A description of a coastal sand dune community is presented by Alvin (1960) in a study of lichen ecology of England's south coast at Dorset. He characterizes the dune lichen vegetation using a "cross-section" of a dune much as I did with a south shore Long Island dune (Brodo, 1961a). Although Alvin's dune system was more complicated (consisting of three ridges), his community is very similar structurally and even floristically to that of Long Island. Unbranched Cladoniae such as Q. coniocraea, Q, chlorophaea, and Q, macilenta were closest to the ocean on relatively unstable sand, giving way to the shrubby Cladinae (Q, s lvatica, Q, impggg, Q, gaggis), Q, furcata, Q, uncialis, and Cornicularia aculeata farther back in protected depressions behind the first main ridge, and finally appearing on the stable second ridge are Lecidea (Biatora) uliginosa, Cladonia crispata, and Q, sguamosa. With a few species replacements such as Cetraria islandica subsp. crisps for Cornicularia aculeata which in eastern America is much more northern, Cladonia submitis for the more northern and/or European Cladinae, and the North American endemic Q, atlantica and Q, sguamosa (which is more mesic on Long Island), the dune community is essentially unchanged in structure. This is but another example of closely related species in different geographic areas occupying similar niches in similar habitats to create remarkably similar communities. V 4. Lignicolous (windswept stumps). Species: Lecanora laevis. Comments: Species occurring in the windswept areas of the island (on beaches and sand dunes) often are very well developed. Lecanora laevis is a good example, often covering old, hard, windswept stumps, especially at Orient Point. 74 B. Hygrohaline stratum (salt spray and storm tide zone) (saxicolous). Species: Bacidia umbrina, Aggrospora fuscata. Comments: The species listed as “bharacteristic” of the hygrohaline community (which is almost non-existant on Long Island) are actually far from their normal habitats (farther inland) and seem to be displaying more of a tolerance for the zone than a preference for it. C. Hydrohaline stratum (littoral zone) (saxicolous) (figure 14). Species: Verrucaria microspora, X, silicicola. Comments: The members of the hydrohaline community are found in no other habitats. Degelius (1940) reported 2, microspora from the upper hydrohaline in Maine. Verrucaria erichsenii, which Degelius found abundant in the lower hygrohaline stratum was not found on Long Island at all. IV. CITY EFFECT A. Introduction. One of the most obvious and far reaching influences on the distribution of Long Island lichens is the proximity of New York City. The city itself (including Brooklyn and Queens as well as Manhattan, Staten Island, and Bronx) has a population of well over eight million people. The western edges of Brooklyn and Queens are heavily industrialized as well. The growth of a city can have a detrimental influence on lichen populations in two ways: direct and indirect (Barkman, 1961). The former involves the wholesale removal of available substrates on which lichens grow, i.e., by the draining of swamps, deforestation, and replacement of soil with concrete and asphalt. It is the less conspicuous indirect influence, the local and long distance effects of city-induced atmospheric changes, which warrants discussion in greater detail. Lichenologists have known of the detrimental influence of cities on lichen growth for almost a century, and within the past 30 years, detailed studies of some of the major European cities have appeared. Barkman (1958) and Skye (1958) have presented excellent reviews of the work up to 1958. Since that time, Beschel (1958) working in and around Innsbruck, Austria, Brightman (1959) studying an area near London, England, Fenton (1960, 1964) working around Belfast, Ireland, Magdefrau (1960) studying in Munich, and LeBlanc (1961) studying the area around Montreal, Canada as well as many others all have contributed to our knowledge in this field. In general, previous workers studying only corticolous lichens mapped the presence of various species around the town or factory in question marking the boundaries of lichen tolerances (sometimes also indicating the 7S 76 limits at which the species are normally developed) (see Beschel, 1958). Barkman (1963) recently presented a striking map of Limburg, Belgium showing the high correlation between the presence of coal mines and chemical factories and the absence of epiphytic lichens. Most studies differentiate between base and breast height presence and almost all present a summary listing the species encountered in order of their tolerance. Rydzak (1958) studied a number of Polish resort towns, treating each town separately and only noting whether each species occurred in the center or at the periphery of the town. He also summarized the data concerning each species but did not rank the species in order of town tolerance although it was possible to do so from his data. Beschel (1958) besides mapping lichen distribution zones around the towns he studied, placed considerable importance on comparative growth rates. working in graveyards from the town centers to the outskirts, he measured growth rates of various common species and discovered definite growth rate gradients. ,Brightman (1959) actually analyzed the substrate of some city tolerant species in an attempt to determine the factors influencing lichen tolerance to city atmosphere. Certain basic trends as one approaches a town were observed by almost all the workers. 1. Fruticose lichens are the first to disappear, followed by foliose, then smooth crustose, and finally leprose crustose lichens. Thus, leprose lichens are apparently most tolerant of cities. A leprose lichen, perhaps a Lepraria, was even noted growing as a greenhouse contaminant in the Paris Botanical Gardens in the middle of Paris (Culberson, 1963a). 77 2. There are definite geographic gradients along which lichens disappear; some are steeper than others depending on the size and extent of industrialization in the cities or towns involved. 3. The closer to a city one travels, the poorer the condition of the lichens encountered, and the less surface area they cover. 4. The so-called "hitrophilous" lichens (living on neutral or alkaline and often highly nitrogenous substrates) are by far the most city tolerant. 5. As one approaches a city, there is a gradual restriction of lichens to areas closer and closer to tree bases, parks, and bodies of water. 6. The disappearance of lichens and the appearance of industrialization are highly correlated, with the effects being greatest where fuel consumption and population size are greatest. 7. A decrease in atmospheric humidity and an increase in atmospheric pollution can be detected towards the center of a town or industrial center . Methods and Observations. On Long Island, three approaches were taken: 1) a transect study of lichens in red oak stands along the north shore at varying distances from the city, 2) the partial reconstruction of past distribution limits of some common lichens and a comparison with their present distributions, and 3) transplant experiments with corticolous lichens. l. Transact studies. The methods used in the transect study have been described in an earlier section (see pages 19-21). Eleven stands were studied in all, each selected to be representative 78 of the red oak vegetation type in as undisturbed a site as possible without regard for its lichen vegetation (figure 18). The advantages of this method over previous transect studies are that a) trees were selected in an unbiased sample, b) exact quadrat positions were established and c) entire stands were studied rather than individual trees enabling frequency per stand to be used as a parameter rather than coverage, the latter being more subject to error. Before one can make conclusions concerning trends in lichen frequency from east to west, it is first necessary to determine if any built in trends are present in the transect, especially in tree composition and tree age, both of which could influence lichen composition within the stands. Figure 19 showing the distribution of trees along the transect establishes that the stands studied are not uniform in composition, and, more important, that there is no significant east-west trend. A slight trend in tree size can be seen in figure 20, especially with Quercus rubra, with the largest trees being in New York City and the smallest in the eastern localities. This trend is probably due to the great age of the city park trees protected from fires and cutting for over a century by park authorities. In addition, the city rarely receives the brunt of the destructive autumn storms that sweep along the east coast and do much tree damage in eastern Long Island. With a tree size gradient of this type, however, we should expect the most abundant and diverse lichen flora in the western localities, and the opposite is true. we therefore cannot use tree size gradients to explain the gradual decrease of lichens from east to west. . ‘- 79 The frequencies of the six most common species along the north shore transect are plotted in figures 21a and 21b. On a standard unit plot (figure 21s), the lichen frequency peaks are clearly evident. If the data are plotted on logarithmic axes on the possibility that the city effect may be decreasing and the lichen frequencies increasing exponentially, the points fall in more of a straight line (figure 21b). Drawing a regression line through these points by eye, a slope can be seen which represents the behavior of a species along the gradient. By comparing the slopes of various species a better idea of relative rates of recovery from the city effect can be derived. At least five important observations can be made from the two graphs: 1) a definite trend exists in the distribution of the lichens, with frequencies increasing more or less continuously up to Stand 9 (with the exception of lichen-poor Stand 8), and then becoming erratic between Stands 10 and 12, 2) different species appear in the flora at different distances from the city, 3) most of the species make their appearance at Stand 4 approximately 25 miles from the center of Brooklyn, 4) maximum development is reached by ngdonia coniocraea by Stand 7, approximately 40 miles from the center of Brooklyn, and by the other species by Stand 9, 45 miles from Brooklyn, and 5) ngdonia coniocraea recovers more gradually than the foliose species, but still reaches its peak closer to the city, and is by far the most city tolerant species of the six. The abberant lichen frequency values in Stands 8, 11, and 12 are apparently due to some factor related to their being very close to the shore (within one mile). Although the local environmental characteristics of the last three stands (nos. 10-12) appear to influence certain species more than others, the localities are clearly east of the city effect zone. 80 2. Distributions. The results.of the transect studies are particularly interesting when compared with the distribution limits of many of the common species (see figures 86 through]jK)). Again, it is apparent that different species have different city tolerances with Cladonia coniocraea again the most tolerant. The sharpness of the distribution boundaries are very striking in some cases. The majority of the lichens have their distribution limit within five miles of the Stand 4 limit noted in the transect study. Since there are no natural climatic or vegetational boundaries at this zone which would explain the limits of so many cosmopolitari species, the city influence can be considered as causal. It should be remembered that at one time, the lichen flora in Brooklyn and Queens was similar to that of the north shore of Long Island today (see pages 367-368). 3. Transplants. Two transplant experiments were carried out using corticolous lichens in an attempt to see, directly, how rapidly city effects can be noticed and how far out on the island they can be detected. The general methods employed in the transplantation have been described on pages 21-22 (see also Brodo 1961b). In each experiment, 50 bark disks were studied, 25 bearing the foliose lichen Parmelia caperata and 25 bearing a crustose lichen (Lecanora caesiorubella subsp. lathamii in the first experiment and Graphis scripts in the second). Five disks with each species were placed in 4 new localities at varying distances from theycity, and five with each species were replaced in the original localities to act as controls. 1:1 each case, the new position of the disk was identical to its original position with regard to height and exposure direction. ‘The transect line for the first transplant experiment (set up in 1J960) — IIIIIIII 81 followed the Ronkonkoma moraine oak forest. Due to some inconsistencies in the vegetation from east to west along the transect, the second transplantation (set up in 1961) was made on a transect following the Harbor Hill moraine on the north shore using five of the same stands surveyed in the transect studies described before (Stands 1, 2, 4, 10, 12) (figure 18). The crustose thalli proved to be unusable in the evaluations due to their susceptibility to bark exudation damage and to the difficulty in assaying thallus damage, and so, only the Parmelia caperata data will be presented. The transplanted thalli were examined twice after transplantation: after four months and after one year. During the one year examination, the disks were rephotographed for comparison with photographs taken at the time of their original removal (figure 23). Color photographs were made with the second year experiments and proved much more valuable than the black and white photos taken with the first year transplants. Kodachrome-II film was used for the color pictures with an electronic strobe flash to insure uniform lighting. In the 1960 as well as the 1961 experiments, clear evidence of distanceu- related damage could be seen in the four month results (table 6). At the end of one year, the damage increased in almost all thalli except the controls, and was in general greater in the central island transplants than in those on the north shore (table 6). The fact that the normal distribution limit of Parmelia caperata occurs between Cold Spring Harbor- and Setauket (figure 95) is especially significant in view of the similar limits seen in the transplantation results. The limits should not be expected to coincide exactly since the thalli were transplanted to 82 positions on the tree trunk suited to thalli in a different stand and may not have represented the most suitable microclimatic niche in the stands in which they were transplanted. 4. Some additional observations. The species found farthest into the city was Lecanora disperse, which was growing abundantly over the concrete fences in the Brooklyn Botanic Gardens and Arboretum, part of Prospect Park. No other lichens were found in the park. This species, widely recognized in Europe as being very city tolerant (Erichsen, 1957), also was covering the upper surface of a concrete foundation near Rockville Center (Nassau County) in a locality where only one other lichen was found. Terrestrial and saxicolous lichens in general were found to be much more city tolerant than corticolous lichens, an observation also made by LeBlanc (1961). In Alley Pond Park (Queens) out of the seven species collected, four were terrestrial (Cladonia chlorophaea, Q, subcariosa, Q. caespiticia, and Q,.pitygea , two were saxicolous on pebbles (Lecidea erratica and Q, coarctata) and only one was corticolous (Cladonia coniocraea on the base of a black oak at the edge of a swamp). The Hempstead Plains locality at Meadowbrook had an almost luxuriant terrestrial lichen vegetatior. consisting of Cetraria islandica subsp. crisps, Cladonia caroliniana, Q, chlorophaea, Q, cristatella, Q, piedmontensis, Q, pleurota, Q, strepsilis, Q, subcariosa, and Pycnothelia papillaria. No corticolous lichens were found on the few trees in the locality. Table 7 presents an analysis of the lichen vegetation by habitat types for the ten localities closest to the city. There are almost three tinuas as many terricolous lichens as the next category. Localities adjacent t‘) long Island Sound supported the most well developed corticolous vegetati‘nn 83 City tolerance of lichens has often been correlated with growth forms (Barkman, 1958; Jones, 1952; Fenton, 1960, 1964) Table 8 shows how growth forms of corticolous lichens growing in the north shore red oak forests react to city influences. C. Results. 1. There is a clear east-west gradient in number of species, degree of cover, and frequency with some species approaching the city closer than others. 2. Results of the transect studies, transplant experiments, and the mapping of natural distribution limits of corticolous lichens all indicate that for the most common species, the New York City influence is felt up to 30 miles from the center of Brooklyn, or just east of the Nassau County boundary. 3. The most city tolerant lichens are terricolous species and saxicolous species. 4. The most city tolerant species is Lecanora dispersa, which is found on concrete or mortar. 5. The city effect is strong enough to kill a transplanted lichen within the four month period between August and December. 6. City tolerant corticolous lichens, when close to the city, are confined to tree bases close to swamps and are found only in their normal dry oak forest habitat farther east. 7. The growth form series of tolerance for corticolous species: leprose --- crustose --- foliose --- fruticose, seen by other workers (see page 76) does not strictly hold in Long Island's north shore, red oak forests. There (table 8), the series is: squamose --- foliose --- leprose -.n. crustose. Fruticose species (in Barkman's [I958] classification) such as 84 Qppgg_strigosa are not found close to the north shore at all, and first appear in more central portions of the island at Hauppage, 40 miles from central Brooklyn. Thus, fruticose lichens on Long Island are the least city-tolerant growth form and agree with the more classical growth form series of city tolerance in this respect. D. Discussion. A still unresolved controversy exists concerning the causes of the detrimental effects of town or city atmosphere on lichen survival and growth. One theory holds that it is the drought conditions induced by the towns' comparatively higher temperature and lower humidity that cause lichens to die in and near cities (Rydzsk, 1958). Other workers who have studied the problem closely, most recently Skye (1958), LeBlanc (1961), Fenton (1960, 1964), and Brightman (1959), argue that air pollution, especially $02 and particulate debris, is by far the more important causal agent. Barkman (1958) has reviewed the basic arguments and concludes that both factors must be involved but that air pollution probably has the greater effect in most areas (see also Barkman, 1963). Beschel (1958), while admitting that air pollution may be an important factor in some cities, feels that city climate is of major importance in restricting the growth of city lichens, at least in the areas he studied in Austria. In large cities air temperatures are significantly raised, runoff of rain is fast and complete, few trees are available for transpirational humidification, and in general there are distinct drought conditions (see Geiger, 1965). At the same time, the cities pour literally tons of solid debris, gases, and other pollutants into the air. The difficulty of separating these two factors is immediately evident. Both factors (heated, dehumidified air and polluted sir) are carried by the prevailing 85 winds, and the decrease in vegetation is reflected in these wind patterns (Barkman, 1958; Skye, 1958). Barkman states that drought effects are limited to the town ares proper whereas the pollution has an effect on areas more distant. Rydzak (1958) claims that so far, no city has been studied where air pollution was great enough to cause any detrimental effects on lichens. However, evidence to the contrary has been presented in a number of studies especially that of Skye (1958) who showed clear correlations between measured quantities of atmospheric pollutants, mainly 502, and the quantity and quality of lichen vegetation. He compared the lichen distribution around a slag mill with the distribution of the principal air pollutants in the same area. Skye found the two distributions to coincide almost exactly, even to the recognition of a double zone of lichen damage, one produced at some distance from the factory by smoke effluent from the stack, and the other, much closer to the factory, apparently caused by the slag heaps on the ground. More recently, Fenton (1964) showed the same sort of correlation with quantitative studies in and around Belfast, Ireland. If Rydzak's statement is true, then lichens would have to be regarded as much less susceptible to pollution damages than vascular plants and many animals including man. Halliday (1961) has reviewed the large amount of work done on the effects of SO2 pollution on man and vegetation pointing out how potent pollution can be on life processes. To cite just one such study, Thomas (1961) showed that a continuous application of $02 at concentrations of 0.14 to 0.18 ppm for three to eleven days is harmful or lethal to some vascular plants such as alfalfa, and somewhat higher concentrations are harmful within a few hours. It would also seem that lower concentrations would have their effect if longer periods of exposure were involved. 86 The question now becomes: to what levels of concentration do atmospheric pollutants reach in large and small cities, particularly New York City. Between 1958 and 1961 the New York City Department of Air Pollution Control recorded average concentrations of $02 of between 0.16 and 0.18 parts per million with maxima of between 0.96 and 1.20 ppm (City of New York, 1962). The United States Public Health Service in their national air sampling network (USPH, 1962) reported particulate concentrations of between 65 and 714 ug/m3 (average 189 ug/m3) in the city and ll-466 (average 80) ug/m3 in suburban Glen Cove in Nassau County. A more complete breakdown of the pollutants and their concentrations is given in table 9. Where does all this pollution come from? One must remember that 302 is mainly a byproduct of coal combustion. New York City consumes 32 million tons of coal annually, 1 1/2 times the amount used in Pittsburgh, and over twice the amount used in Detroit (Katz, 1961). There are 10 major coal- burning power stations in New York, plus an oil refinery in Brooklyn (U. S. D. H. E. W. 1962), to which must be added the incineration products of the refuse of eight million people and the products of the coal and oil burned to heat their homes. Hydrocarbon pollution cannot be overlooked either (see table 9). Katz (1961) points out that every short ton of gasoline consumed as auto fuel will produce approximately 141 pounds of organic contaminants (i.e., hydrocarbons), and 24.6 pounds of nitrous oxides plus smaller amounts of sulfur dioxide and other chemicals. ' Considering that there are over a million cars and 200,000 trucks operating in New York City, consuming over a billion gallons of gasoline annually (Greenberg & Jacobs, 1956) automobile exhaust pollution can rise to very high levels. 87 But even with all these sources of pollution, concentrations would probably not be as high as they are were it not for the climatic factor of "thermal inversion," very common in New York City, which acts in concentrating the atmospheric gases and particles to many times their normal levels (Greenberg & Jacobs, 1956). It is evident, then, that air pollution in and around the city does reach concentrations high enough to have a detrimental effect on vascular plants. Are lichens more or less sensitive than the higher plants? Biological evidence indicates that lichens are much more sensitive to pollution. Lichens tend to accumulate minerals efficiently and perhaps also accumulate pollutants (Gorhsm, 1959; Smith, 1960b). In addition, lichens are slow growing and long lived and can therefore accumulate large quantities of toxic materials over a long period of time whereas other plant types usually shed large areas of toxin-laden parts each season. Important too is the fact that in New York City, pollution levels are highest between November and March (Greenberg & Jacobs, 1956) at a time when lichen assimilation may also be highest (Smith, 1962). The only data we have on the actual concentrations required to destroy or at least exclude lichens are provided by Fenton (1964). He concluded that no lichen can tolerate smoke in excess of 250- 300 pg per cubic meter or 802 concentrations in excess of .035 ppm. Since Fenton did not measure the humidity levels along with pollution levels, it is still not clear how much lichen damage was due to drought and how much to the pollution; but if his data are nreasonably applicable, they would certainly help to explain the tremendous extent of the city-effect in western Long Island where pollution concentrations are often five to six times and sometimes 20 to 30 times this level. 88 However, Rydzak (1958) presented data showing that in small Polish resort towns where industry is virtually nonexistent and automobiles are very scarce, lichens still show gradients of diminished growth approaching the towns. He regarded the centers of these towns as essentially having no air pollution, or at least having no more pollution than do the outskirts of large cities where the lichen flora is normal. Although he neither presented nor cited measurements to prove that this relationship of $02 concentration was indeed a fact, we are asked to assume that fuel consumption (for heating homes), garbage disposal, and the small amount of industry present in these towns (a third of which have over 15 factories each) did not contribute ”significantly" to the air pollution in the towns. Making this somewhat tenuous assumption, however, we must agree with Rydzak and conclude that something other than air pollution is responsible for the city-effect on lichens in these towns. He then presents many meteorological data showing that in each of these towns, there is a distinct decrease in air humidity towards the town centers which correlates with the decrease in lichen flora. Based on these findings, he formulates the hypothesis that it is city- induced drought alone which limits lichen growth not only in small towns such as he studied, but also in larger cities. ”Impoverishment of lichen vegetation in small towns is an unquestionable proof that the analogical poverty of lichen flora in the studied large towns is not produced by toxic components of the air” (Rydzsk, 1958, p. 312). Whether we accept or reject Rydzak's gross overstatement, city induced drought undoubtedly has an effect on lichen distribution in and very near towns and must be considered in any attempt at explaining the city-effect. New York City has a somewhat drier climate than eastern Long Island, but the average humidity never drops below 60% in the city and only shows an 89 average difference of less than 5% relative humidity from one end of the studied transect to the other (see figure 6). It is conceivable, however, especially in view of Ried's results (see page 37) that it is the alternation of dry periods and humid periods which is the decisive factor in limiting the distribution of some lichens close to the city, and the Long Island lichens may also be responding to humidity effects. Most authors agree that the drought hypothesis and the pollution hypothesis are not mutually exclusive. Even Rydzak, who comes closer to claiming a unilateral cause of city-effects (via drought) than any other worker, admits, "If there existed an industrial centre emitting high concentrations of 802 and other toxic products into the air (no such centre has been studied so far) then, of course, the toxic and drought factors would co-operate in eliminating lichens from this place" (Rydzsk, 1958, p. 313). An analysis of the types of town effects reported so far by various workers, particularly Fenton (1964), Rydzak (1958), and Jones (1952), together with information gathered on Long Island, suggests a possible solution to the apparent discrepancies. This solution is mainly based on changes observed in the vertical distribution of corticolous lichens at various distances from a town, together with observations on the far outskirts of areas subjected to heavy atmospheric pollution where lichen growth is limited and yet there are no shifts in vertical distribution. It has been noted by authors cited above as well as by others that many corticolous lichens normally found at breast height occur at lower heights on trees or are restricted to trees occurring in humid localities as they approach closer and closer to a city. This has been explained by Rydzak (1958) as being due to the increased humidity compensating for the city-induced drought, and by Jones (1952) as due to humidity increase as well as a 90 decrease of windblown smoke at the tree bases. Barkman (1958) was of the opinion that the vertical zonation effect was due to moisture differences. Although it appears to me that moisture differences are by far the greatest cause of the vertical shift, there is the distinct possibility, even likelihood, that vertical pollution differences may be adding significantly to the effect. Differences in air velocity within the tree layer of a forest are known to occur, with the lowest velocity at the base and highest at about breast height on the trees (Geiger, 1965). Thus, the greatest volume of air would pass over the middle portion of the bole carrying with it the highest quantity of gas pollutant per unit time. In addition, as polluted air passes through ground vegetation, the air is undoubtedly "cleaned” to some extent and causes air moving along the ground to carry relatively less solid pollutants than air moving at a higher level (Geiger, 1965). There are, however, several additional points to take into account. First, toxic materials (such as sulfur and nitrogen compounds as well as hydrocarbons) are to a large extent associated with dust and aerosols (see Katz, 1961). It may even be that particulate sulfur compounds such as zinc ammonium sulfate and ammonium.sulfate are even more serious air pollutants than $02 (Corn and Deuaio, 1964). The slow moving air close to tree bases would be depositing more particulate matter in that region than the more rapidly moving air would be depositing higher on the bole. This might compensate somewhat for the "bleaner" air at the tree bases. Furthermore, it is not clear to what extent the air velocity differences may occur on trees along roadsides, and it is upon roadside trees that most of the European data rests. These problems can only be solved by careful investigation, and not by conjecture. 91 On Long Island, the only manifestation of the humidity compensation phenomenon (if this is indeed what it is) is in the restriction of Cladonia coniocraea to tree bases close to swampy areas when the species is in or near the city. I believe it is a response to increased and, perhaps more important, more constant humidity in the presence of drought conditions which governs this phenomenon although it may also be that the lush swamp vegetation "cleans" the air to some extent. However, all indications are that air pollution has a more powerful influence on lichen vegetation, particularly over comparatively long distances. If one accepts the premise that each city affects the lichen vegetation by means of two distinct city-produced factors, drought and pollution, the latter acting over longer distances than the former, a possible explanation for the city effects described by Jones, Rydzak, myself, and others emerges. The explanation is based on these suppositions: l) certain lichens have greater intrinsic pollution tolerances than others due to physiological or growth form characteristics,.i) downward shifts in the vertical distribution of corticolous lichens normally found at breast height will occur within an area where the drought effect is acting, and 3) no appreciable shift in vertical distribution will occur due to the effect of air pollution alone. These three suppositions, particularly the last, may be premature and seriously oversimplified. They certainly need substantiation through careful analysis of climatic and pollutant differences in relation to lichen growth. However, the suppositions are supported by the observations made so far, and they provide us with a starting hypothesis which is testable and can be accepted or rejected as future facts come to light. To illustrate the point, let us look at three studies of city effects, one made in towns having no appreciable air pollution (Rydzsk, 1958), one 92 made in an area having moderately high air pollution (Jones, 1952), and one made in an area with very high levels of air pollution (Long Island). Rydzak noted a lowering of vertical zonation with lichens decreasing in number and type but never dropping out entirely even in the center of his towns. The reason seems to be that air pollution was not a significant factor in his town effects. Since the lack of moisture was the only important lichen-limiting factor within the towns, only the most hygrophilous species (especially the fruticose species) dropped out with the others responding by becoming limited to tree bases. Jones, studying roadside trees in the Midlands of England, noted distinct vertical shifts with fruticose and foliose species as he approached industrial centers. He did not mention exactly how far from the center of town this shift was apparent although he seemed to indicate it was evident over most of the distance of lichen paucity. His observations on lichen growth might be explained by supposing that the drought zone and the pollution zone overlapped over much of his study area, and so some pollution-sensitive as well as the pollutionetolerant species were affected by the city-induced drought. 1 0n Long Island, air pollution acts over such a great distance, it effectively cuts the lichen population down to only a few highly pollution- tolerant corticolous species before the drought zone is encountered, and so few or no species show any change in vertical zonation approaching the city. The most city-tolerant corticolous species happens to be normally base-dwelling and although that species cannot change in its vertical distribution, close to the city, it does become restricted to humid localities. This suggested explanation is presented graphically in figure 24. The drought and pollution curves are drawn on a strictly hypothetical basis. The 93 drought effect, as represented, decreases slowly towards the edge of the town and then rapidly drops to zero near the outskirts (see Geiger, 1965). The pollution effect maintains itself over a longer distance but also drops off more rapidly outside the town. From Jones' data as well as mine (see figure 21), the lichen vegetation appears to slowly pick up as one leaves the town, then rapidly approaches normality. This progression is incorporated into the vertical distribution curve by letting the greatest width of the ”line” equal normal lichen abundance and allowing the line to taper according to the shape of the lichen abundance curve. In this way, the complete disappearance of the lichen vegetation can be represented at any point on the horizontal scale of distance from the town center. It must be understood that although they are based on actual observations of city effects, the representations are merely schematic. They can, however, be used as models for future work since all the curves can be derived empirically. The ranking of growth form types from leprose to fruticose with decreasing city tolerance correlates both with drought susceptibility (due to degree of transpirational surface) and pollution susceptibility (due to increase of absorption surface and increased distance from substrate with attendant greater gas flow) and so growth form alone cannot be used as an indicator of the cause of city-effects. Fenton (1960) goes into greater detail in analyzing the possible factors involved in the susceptibility of certain growth forms to air pollution. 0n Long Island, the fact that the squamose type appears to be most city tolerant (more so than crustose) is almost certainly due to either the generally poor lichen flora of Long Island and the outstanding city tolerance of the squamose species Cladonia coniocraea, or the small number of city tolerant species represented in the Long Island lichen vegetation. 94 It is possible that calcareous rock dwelling species gain their city tolerance from the ability of their substrate to neutralize the acid toxins from the air (Brightman, 1959). This then would explain the presence of Lecanora dispersa on concrete fences in the heart of industrial Brooklyn. The high tolerance of terricolous and noncalcareous saxicolous species is not easy to explain. Some of the terricolous Cladoniae were undoubtedly protected from much of the surrounding atmosphere by being situated deep in moss mats, but others were quite exposed in dry open fields as were the saxicolous species. Soils are known to be highly buffered (Russell, 1950) and it may well be that lichens adapted to live on normally acid soils can continue to survive if any excess acidity contributed by acidic pollutants is neutralized, much as it is on calcareous rock. In addition, it is very likely that these dry open field terricolous and saxicolous species have a high degree of natural drought resistance. E. Conclusion. Although city drought may well influence the distribution of pollution- tolerant lichens growing within or near the city, atmospheric pollution is believed to have the greater influence on the distribution of Long Island lichens. V. FLORISTIC ELEMENTS A. Introduction. In an area as small and geologically uniform as Long Island, historical factors cannot explain local distribution patterns since ample time has been available for the uniform distribution of any plants which arrived on the island other than very recently introduced adventives. The migration routes by which these plants reached Long Island are of considerable interest, however, and it is worthwhile to examine some of the probable sequences of events which fashioned the lichen flora of Long Island as we see it today. In attempting a floristic analysis of the lichen flora of Long Island, it has been necessary to analyze the major distribution patterns represented in eastern North America, and to view these patterns not only with regard to North American distributions in general, but also, to some extent, with certain aspects of worldwide distribution. There is a much greater need for a broad geographic perspective in dealing with distributions of lichens as compared with flowering plants since endemism on a species level is much more common in the latter (see Ahti, 1964). Approximately 26%3 of the lichens of Long Island are endemic to North America as compared with an estimated 652 of the vascular flora. (The vascular plant statistics were derived from an unbiased sample from Smith and Ogden's unpublished preliminary flora of Suffolk County, in conjunction with comments on endemism in Fernald [1959] .) Many authors have contributed to our understanding of the floristic patterns to be seen in eastern North America. Good (1964) provides a general pattern 3 Computed from a sample of 81% of the total lichen flora. 9S 96 of the major elements. The forest regions of the eastern deciduous forest as described and mapped by Braun (1950) although not delineated by floristic criteria reveal some of the basic floristic features of eastern North America, particularly the strong influence of the Appalachian Mbuntains. There have been few general treatments of lichen distribution in North America. Thomson (1963) in his monograph of Physcia, discussed American distribution patterns with an emphasis on extra-American relationships. Although his eight categories have limitations for the kind of floristic analysis I would like to attempt here, two of Thomson's categories are used in only slightly modified form. The phytogeographic system proposed by Hale (1961a) is very useful and many of his categories are retained essentially unaltered. B. Ihg,classification 2; elements. The floristic elements have been broadly classified according to general climate. An Arctic-Boreal, Temperate, and Tropical element can thus be recognized. The elements are each divided into two or more "subelements," and in one case, further subdivided into geographical units. The limits of these categories are presented below, and representatives of each in the Long Island lichen flora are listed in table 10. Element I; Arctic-Boreal. The Arctic-Boreal element is that element which has no climatic northern boundary. Since tree line would be a northern boundary for arctic corticolous species but not for arctic terricolous species, substrate was bypassed as a limiting criterion. we can recognize two subelements within the Arctic-Boreal element. The Arctic-alpine subelement corresponds closely with Thomson's (1963) "circumboreal 97 arctic-alpine" category. It is distinctly arctic in character extending into temperate United States only in the alpine zones of some of the eastern and western mountains. With this circumscription, it is obvious that no member of this subelement could be present on Long Island. The members of the Arctic-Boreal element which do extend into boreal and temperate climates are grouped together as the Boreal-ggmperate subelemgg£_(figure 25). These species are generally very widespread due to their broad climatic tolerances and access to circumpolar migration routes. Element II; Temperate. In the Temperate element are included all species with relatively distinct northern and southern climatic limits, usually close to the northern and southern boundaries of the United States. The temperate element can be divided into six subelements. The first is more or less intermediate between typically arctic and temperate distributions. This §g££h_Temperate subelement is not considered as arctic due to its relatively clear northern boundaries, but shows distinct boreal tendencies in many instances. It is best developed in northern United States and southern Canada, although it often extends southward to include most of continental United States (figure 26). The ”circumboreal north temperate" category of Thomson (1963) corresponds to this subelement which includes many of the more widely distributed common species. Three important physiographic features of temperate eastern North America are the coastal plain, the Appalachian and Ozark mountain systems, and the Mississippi valley. The Appalachian mountains form the core of the area occupied by the Appalachian subelement. Extensions and slight modificatjqnas of the basic Appalachian distribution permit us to recognize a number of) “units”‘within this subelement. The Appalachian unit includes only species 98 whose basic distribution is along the NE-SW mountain chain alone (figure 27). Extensions to include the Ozark Mountains, the Great Lakes region, and the southern Rocky Mountains define the Appalachian-Ozark, Appalachiag-Great Lakes, and Appglgchian-Great Lakes-Rocky Mountains units, respectively (figures 28-30). Species confined to any or all of the three segments of the coastal plain (i.e., Gulf, southern Atlantic and northern Atlantic) are included in the Eggstal Plain subelemggg (figure 31). This subelement often shows an extension into the Mississippi Valley. A large number of eastern temperate species are not restricted to the Appalachian or coastal plain regions but are found throughout the eastern deciduous forest from the Mississippi Valley (or even farther west) to the Appalachians or the east coast. These species comprise the East Temperate subelement (figure 32). There sometimes is a distinct northern or southern concentration within the subelement (see maps of Eggmelia galbina and g. livida in Culberson, 1961), but its division into two units is not warranted. Often, there is a narrowly restricted concentration of records in the northeastern states, and it is difficult to decide whether the species belongs to an eastern segment of the North Temperate subelement, a northern segment of the East Temperate subelement, or a portion of an Appalachian- Great Lakes distribution. Any or all of these may be involved, of course, and there is no value in recognizing separate categories. "Northeast Temperate“ species have arbitrarily been listed with the East Temperate subelement. There are a number of wide-ranging species which are apparently relics of ancient and worldwide distributions and which now are restricted in their distributions by their narrow climatic tolerances (see below). These specieas are grouped together into the Oceanic subelement. They are generally 99 characterized in North America by having east coast - west coast disjunct distributions (figure 32A). Other species with obvious oceanic tendencies (e.g., Lobarig.guercizggg, Collema subfurvum) but which have well defined distributions in one of the subelements already described, are considered only with the latter. In table 10, they are designated with asterisks. The oceanic type of distribution (usually considered as an element in its own right) has perhaps been studied more than any other, especially in Europe (Degelius, 1935; Mitchell, 1961; Faegri, 1958). Degelius (1941) also made some observations on oceanic species in eastern North America. The Oceanic subelement is characterized by occupying areas with high atmospheric humidity (although degrees of rainfall may differ from one place to another), and where temperature fluctuations are small from one season to another (i.e., having mild winters and cool summers). Though areas of this type are generally coastal, they need not be. A definite oceanic flora can be found in the Smoky Mountain region of Tennessee and North Carolina (Degelius, 1941). In the present study no distinction is made between "bu-oceanic” (strictly oceanic in distribution) and "suboceanic" (basically oceanic with a somewhat broader tolerance of other climates) as was done by Degelius (1935) since lichen distributional and ecological limits are still relatively poorly known within North America as compared with Europe. Members of the gritgmg subelement are restricted by habitat availability to the temperate maritime zones along the coast. Compared to the maritime flora of Europe, this subelement is very poorly developed in eastern North America (see Degelius, 1940). Although the Maritime subelement theoretically includes species restricted to the sea coast by the presence of salt water, salt spray, or some associated marine influence, no aerohaline species fronl Long Island appear to fit into this category. 100 Element III. Tropical. Species which show a basically tropical distribution are grouped into the Tropical element. Western hemisphere representatives are usually widespread in Central and/or South America and sometimes can be found in other tropical areas throughout the world as well. The element is manifest in eastern North America centered in the Appalachian mountain system and on the coastal plain, thus conveniently dividing the element into Appalachian-Temperate and Coastal Plain subelements. It is perhaps also proper to recognize an Oceanic subelement, although there appears to be only one example on Long Island. C. Summary pf significant features. Table 11 presents a summary of the categorization of the lichen flora into its phytogeographic elements and subelements, with figure 34 giving a graphic representation of some of the important facets of the major categories. The summary is based on table 10 which includes approximately 81% of the known Long Island lichen flora, all the species for which we have some good phytogeographic information. Some observations which deserve special attention are the following: 1. The Arctic-Boreal element is represented by 21% of the flora, all but two species being partially or entirely circumboreal. 2. Many of the most common species on Long Island (see table 11) are members of the Arctic-Boreal element, e.g., Cladonia chlorophaea, g, coniocraea, Parmelia sulcata and g, saxatilis. 3. The Temperate element is most abundantly represented (71% of the flora). 4. All North American endemic species are in the Temperate element, mainly in the East Temperate (6%) Appalachian (7%) and Coastal Plain (8%) subelements. In all, 24% of the lichens of Long Island are endemic. 101 5. Of the sampled species with an East Asia-East America disjunct distribution (16 in all), by far the greatest number (38%) are found in the East Temperate subelement. 6. Considering its northern latitude, Long Island has a surprisingly good representation of tropical species (8%). Most members of the Tropical element are confined to the coastal plain in eastern North America. 7. Most of the species having amphiatlantic distributions (12%) are represented in the Temperate element (the East Temperate and North Temperate subelements). D. Discussion. Braun's (1950) map of the forest regions and sections in eastern North America reveals that Long Island lies at the apex of three major forest types: the Oak-Chestnut region with its origin in the Appalachian foothills and southeastern piedmont, and the Southeastern Evergreen forest region which lies on the Gulf and Atlantic coastal plains. The Hemlock-White Pine-Northern Hardwoods region lies to the north but is separated from direct continuity with Long Island by an area of Oak-Chestnut forest in southern Connecticut. The new vegetation map by Kfichler (1964) shows a very similar pattern, but with the vegetation units more precisely delimited. For example, Kfichler's map clearly shows the change from the oak-hickory-pine forest of the southeastern piedmont to the northeastern pine-oak forest of southern New Jersey, Long Island, and Cape Cod, and shows more clearly the differences between the Gulf coastal plain and the central- and northeastern coastal plain vegetation. Thus, there are at present three unbroken biological "highways" along which species can migrate to Long Island from the south, and an almost uninterrupted conifer-hardwoods forest to the north which provides easy 102 access for northern species. These migration routes have existed essentially unchanged for thousands of years. The greater part of Long Island has only been available for colonization since late Pleistocene time after the last retreat of the Wisconsin ice in that area (ca. 15,000 - 20,000 years before the present). It is highly probable that a considerable portion of the northern continental shelf now underwater was exposed as a coastal plain during and just after the last glacial maximum (Fogg, 1930; Nichols, 1958). As the Long Island area became ice free this extensive coastal plain would have provided an opportunity for unhindered plant immigrations from the south and west. Soon after, sea levels rose due to the melting of the glaciers (Flint, 1957) flooding the Long Island Sound area, and separating Long Island from any closer connections it might have had with New Jersey, as well as submerging much of the south- eastern New England coast. Much of the submerged coastline north of Long Island reemerged with the up-doming of the area (Flint, 1957) but since Long Island was apparently on or just south of the "hinge line," it remained an island. The high percentage of circumboreal species in the Boreal-Temperate and Northern Temperate subelements (95%,and 77%, respectively) is not surprising in view of the extreme likelihood of late Tertiary and Pleistocene land bridges across parts of the arctic which allowed the free flow of plants from one continent to another (Fernald, 1931; Flint, 1957; Colinvaux, 1964). Graham (1964) cites evidence for North Atlantic migrations via the arctic islands during periods of temperate climate in the Cenozoic. Arctic species probably can still migrate via the northern islands in a circumboreal route (Li, 1952). 103 The theory of pre-Pleistocene continental drift is still very much alive and if true may explain many of the present day amphiatlantic lichen distributions (see Dansereau, 1957; Good, 1964). Hulteh (1962) suggested that amphiatlantic patterns are best explained by postulating eastern and western continental migrations from the Bering Strait region rather than trans-Atlantic migrations. Colinvaux (1964) has sketched the Pleistocene floristic activity over the Bering land bridge. Dahl (1950) considered present day American-European disjunct distributions of lichens and some other plants as having originated from arctic parental populations which survived the ice ages in unglaciated areas of the arctic. Among the Long Island lichens, 14% of the species have amphiatlantic distributions. North Temperate and some Oceanic species possibly migrated across the northern regions during pre-, inter-, cu: post-glacial warm periods and later retreated southward with a cooling of the northern regions and the glacial advance, returning only as far north as the northern conifer-hardwoods with the disappearance of the ice. Potzger (1952) presented palynological evidence to suggest that the pine barrens of southern New Jersey served as a refugium for many boreal communities which were displaced southward by the Wisconsin glaciation. These northern plants survived the ice ages side by side with southern communities, only to migrate northward again with the retreat of the ice. Long Island, therefore, was in an excellent position to be invaded by many of these northern species. Cladonia terrae-novae Ahti, though not a part of the Long Island lichen flora, probably derived its distribution pattern in this way (see page 259 ). Possibly some North Temperate species also were introduced into the North American flora in post- glacial times during the post-glacial warm period from Eurasia and were ‘ eliminated from the northern boreal and arctic latitudes following the recent 104 cooling in northern climate. Fernald (1931) and Braun (1955) present evidence showing that the Appalachian and coastal plain floras originated from pantropic connections that invaded the Appalachians at a very early time. With the uplift of the area during the Tertiary, some species moved out onto the newly exposed coastal plain leaving only fragmentary relics behind on the Appalachian plateaus. Much speciation appears to have occurred in the southern Appalachians during the long period of its isolation (Fernald, 1931), and the high percentage of endemics seen in the Appalachian and Coastal Plain subelements may date from this time. During the Pleistocene glaciation, coastal plain species were restricted to regions south of the ice, although probably not very far south (Braun, 1955; Potzger, 1952). With the retreat of the ice, the northeastern coastal plain became available for colonization from the south. Fernald (1931) cited much botanical evidence to support his theory that there was a post- glacial period of relatively warm climate when the entire coastal plain was connected by a continuous land formation perhaps as far north as Newfoundland (see also Braun, 1955). If this was the case, there was an excellent route available for the migration of the new coastal plain species northward to Long Island and beyond (see above and Fogg, 1930). The East Temperate subelement had at least two origins: one, as an eastern segmentation of a north temperate distribution, and the other as a broadening Appalachian distribution. Those East Temperate species which originated from the north are likely to show an amphiatlantic pattern, whereas those coming from the Appalachian center often are either North American endemics or show evidence of a widespread Tertiary (and East Asia disjunct) distribution (see below). 105 The historic relationships of the various elements, subelements, and units are summarized in figure 35. The similarity between the floras of temperate eastern North America and eastern Asia have long been recognized and discussed (Li, 1952). This classical disjunct distribution pattern is clearly evident within the temperate element of Long Island lichens. Eight per cent of the Long Island flora represents Eastern America - Eastern Asia disjuncts. Li (1952) states that Asian - Eastern Temperate floral similarities represent a relic distribution of a Tertiary flora which once covered the temperate to arctic northern hemisphere. The fragmentation of the flora was caused by many geological changes including mountain formations, continental submergence, climatic change, and glaciation (Li, 1952). It is especially interesting to note that we see these disjunct patterns on a species level with lichens whereas phanerogamic botanists rely on generic similarities (Fernald, 1931; Li, 1952). This sort of evidence can suggest extreme genetic stability and slow rate of evolution in many lichen fungi as compared with flowering plants (see also Thomson, 1963). Raven (1963) points out, in discussing amphitropic distributions, that disjunct distributions on a species level, especially when involving autogamous organisms (as would be the case with lichens) probably are due to long distance dispersal particularly by migrating birds and not to any once continuous populations which became extinct in intervening areas. While this may be true of amphitropic distributions of flowering plants along bird migration pathways, it is hardly possible that the east Asia disjunct distributions of dozens of species in the eastern American lichen flora could have their origin by long distance dispersal, especially when this disjunct pattern is well known in other plants at higher taxonomic levels. 106 Degelius' wide experience with the European lichen flora permitted him to recognize a number of European-American vicariant pairs in his studies of the lichen flora of Maine (Degelius, 1940). He proposed a new category, ”subvicarious species," to include species which do not entirely displace each other but rather, show different frequency ratios in the different areas. He suggested various alternate possibilities for vicariant and sub- vicariant combinations as follows (nos. 1-4). Capital letters indicate the species is abundant, and small letters indicate it is rare. Alternatives 5-11 have been added and will be discussed below. Alternative N. America Europe 1 A B 3 (true Vicariants) 2. A + b B 3 A a + B (sub-Vicariants) 4 A + b a + B 5. a b (?) 6. a B 7 A b 8. a + b B (not Vicariants) 9. A a + b 10. A + b A 11. A A + b In order to discuss these alternatives, we must first define "vicariant (or vicarious) species." Vicariants are disjunct, but closely related species which are similar morphologically and often ecologically. I think it is fair to say that most definitions implicitly or explicitly assume approximately equal abundance of the two vicarious populations. This would then immediately exclude alternatives 6 through 11, and especially 10 and 11 as vicariants. Since Degelius almost certainly wanted to emphasize relative abundance rather than absolute abundance of vicarious pairs, alternative 5 is superfluous (being equivalent to alternative 1) and can be eliminated. 107 Of course, there are many other possible combinations which could be listed, but they clearly do not represent vicariants. European-American vicarious species found in the Long Island flora are listed in table 12. Degelius' use of Parmelia ( Pseudevernia ) cladonia and g, furfuracea as an example of alternative 2 is not applicable. Pseudevernia cladonia is relatively rare in North America while 3. furfuracea is more widespread and often common. The pair would therefore more closely fit into alternative 10 (assuming the North American and European chemical populations of g, furfuracea are basically conspecific) and should no longer be considered as vicariants. It is interesting that alternatives 3 and 4 are entirely absent. Even Degelius (1940) could not give an example of no. 3, and his example of no. 4 (Lecanora carpinea - L. pallida) is no longer applicable in the light of recent studies (Imshaug and Brodo, in press). A consideration of the origin of Vicariants and their probable relative abundance is, therefore, of interest. Vicariants originate from speciation of an isolated portion of a widespread population. The geographic separation of a parent and daughter species may occur either before or after the initiation of the new species. L3ve (1955) makes a definite distinction between the two types of resulting vicariants. He regarded those species having arisen pips; geographic segmentation of a parent species as ”true vicariads," and those species which arose within a parent population by some immediate genetic isolating mechanism (such as polyploidization) and 13525 became separated from the parent population, as ”false vicariads.” Lave, after discussing the usefulness of the distinction, explains how in flowering plants, cytological studies can establish what type of vicaribm is involved in each particular case. Even if the distinction is useful, as it may well be in certain organisms, in lichen fungi, which 108 appear to be genetically "apomictic," the distinction cannot be made. In any case, if either type of vicariism mentioned above occurs, it is evident that alternatives 2 and 3 should be more common than alternative 1 because of the low probability of entirely displacing a parent population (i.e., with the parent population becoming totally extinct in one area). It is therefore significant that two of the tree examples of no. 1 cited by Degelius (1940) (Lobaria guercizans - L, amplissima; Ughilicaria papulomi * U. pustulata) now appear to be the more common alternative 2. (It should be pointed out, however, that the North American population of Lobaria agplissima is disjunct from that of L. guercizans.) It is therefore even more puzzling that there are no examples of alternative 3 in the lichen flora. One could hypothesize that all lichen Vicariants are "true” Vicariants (pgppp_L3ve, 1955) and have come fpgp_Europe (suggesting an interesting way of analyzing a migratory direction), but this would be an unlikely conjecture since it is also possible that "false vicariism" is involved and in the opposite direction. Alternative 4 which requires an original bi-directional migration, or occasional long distance imports in one or both directions with the maintenance of an equilibrium ratio between the two species (see MacArthur & Wilson, 1963), appears to be least likely of all. E. Summary. The affinities and possible origins of the various phytogeographic categories are presented schematically in figure 34. In general, there seems to have been two routes of worldwide distribution: arctic-boreal and tropical, with the Temperate element largely derived from one of these two origins. Long Island is approached from the north via the oak-chestnut forests which included parts of western Long Island before urbanization. The fragmentation 109 of what probably once were continuous European-American boreal or temperate distributions gave rise to many examples of amphiatlantic patterns including several vicarious pairs of species. Many northern species reached the island from the south, however, just after the last glacial maximum. The temperate species which originated in southeastern United States, in part, reached Long Island via the Appalachian Mountain system, which partially empties out into northern New Jersey and from there, into New England. The Atlantic coastal plain provided a coastal "highway" along which southern species, many of which originated in the southern Appalachians, could migrate northward to Long Island. These same two migration routes were used in the introduction of tropical species into the Long Island flora. Oceanic species, many of which had ancient origins and worldwide distributions, became isolated in various areas of eastern North America in late Tertiary and Quaternary times such as in the humid and comparatively mild Smoky Mountains of the Appalachian system and along the northeastern coast including parts of eastern Long Island, Nantucket Island, Cape Cod, Newfoundland, and Nova Scotia. VI. THE LICHEN FLORA A. Collections. In 138 Long Island localities, approximately 3200 collections were made. An additional 290 collections in southern New Jersey, 200 on Nantucket Island, and 400 on Cape Cod provided information on mainland and island floristic connections with the Long Island lichen vegetation. Floristic distribution maps often come under serious criticism because they are said to represent the perambulations of the collector rather than the distribution of the organisms. To overcome this shortcoming and to greatly increase the value of distribution data all species seen in a locality were collected no matter how common they are. In this way, a determination of where a species does not occur can be made almost as accurately as the determination of where the species does occur. As a result, a map can be prepared to indicate species absence as well as presence (as in Imshaug, 1957a). Of course, rare species will occasionally be missed and common ones will occasionally be forgotten, but, on the whole, an attempt at a complete-collection is a significant improvement over the more haphazard collecting methods of the past. This method was employed in all the Long Island, New Jersey, and Cape Cod localities. The Long Island localities are listed below and are represented by numbered dots in figure 15. KINGS COUNTY: (1) Prospect Park (Brooklyn Botanic Gardens). QUEENS COUNTY: (2) Forest Park, oak woods. (3) Alley Pond Park, oak woods and field. NASSAU COUNTY: (4) Sands Point, shaded maple-oak woods and open field. (5) North Hills, dry slope above swamp. (6) Valley Stream, Acer rubrum swamp 110 111 and oak clearing. (7) Rockville Center, Hempstead Lake State Park. (8) East Meadow, "Hempstead Plains." (9) Brookville, mature oak woods. (10) Glen Cove, mature red oak - beech woods. (11) Laurel Hollow. (12) Cold Spring Harbor, path and black oak woods. (13) Cold Spring Harbor, woods. (14) Syosset - South Huntington, young oak woods. (15) Bethpage, young oak woods, recently burned. (l6) Massapequa - Seaford, black oak woods. SUFFOLK COUNTY: (17) Centerport, red oak - chestnut oak woods and roadside. (18) Vernon valley (near Northport), red oak woods. (19) South Huntington - Half Hollow, oak - hickory woods. (20) Dix Hills, oak woods and mossy slope. (21) Commack, mature oak woods. (22) Deer Park, oak woods and pine woods. (23) Deer Park, woods, swamp, and field. (24) Near Babylon, pine - oak woods bordering acid bog. (25) Captree State Park, sand dunes. (26) Near King's Park, red - oak woods. (27) San Remo, beech - oak - ash woods. (28) Hauppauge, wet woods. (29) Central Islip, young oak woods. (30) Ronkonkoma, oak - pine woods. (31) Heckscher State Park south of E. Islip, oak - hickory woods. (32) Oakdale, young oak woods; West Sayville, roadside. (33) Fire Island, Cherry Grdve, Sunken Forest Preserve, IlsgDQppgp_grove. (34) Missequogue, chestnut oak - red oak woods. (35) St. James, red oak - black oak woods. (35 - 36) Nesconset. (36) Centereach, pine - oak barrens burned over. (37) Selden, roadside. (38) Farmingville, young oak woods. (39) Patchogue, young oak woods. (40) Patchogue, open pine barren recently burned. (41) Sayville, wet oak - pine woods. (42) Old Field, dry oak woods. (43) East Setauket, red - scarlet oak woods. (44) Port Jefferson Station, dry oak woods. (4S) Coram, mature oak woods. (46) Coram, burned over pine barren. (47) Middle Island, young oak - hickory woods. (48) Patchogue, field, young oak woods and maple 112 swamp. (49) Bellport, open field. (S of 49) Fire Island opposite Bellport, between dunes. (50) Miller Place - Mount Sinai, red oak - chestnut oak woods. (S of 50) Miller Place, oak - hickory woods. (51) Middle Island, mature oak woods. (52) Upton, pine - oak woods; Ridge, oak and pine woods. (53) Upton, Brookhaven National Laboratory, roadside boulder, oak woods. (S4) Upton, Brookhaven National Laboratory, pine woods and pine - oak woods. (55) Yaphank, oak woods, field and roadside elm. (56) Brookhaven, oak - pine woods and bog. (57) Yaphank, pine - scrub oak barren burned over. (58) Brookhaven station, pine barrens. (59) Shirley, oak woods. (60) Fire Island, S. of Shirley, sand dunes. (61) Shoreham, sand bluffs and black oak woods. (62) Shoreham - Wading River, shaded oak - hickory woods. (63) Wading River Station, old black oak woods. (64) Montauk Trail, young pine barren. (65) Upton, young black oak woods, oak - pine woods, pine woods and maple swamp. (66) Manorville, open quaking bog and surrounding oak woods. (67) Manorville, black oak woods and mature oak woods. (68) Manorville, oak woods, mature oak woods. (69) Manorville, pine - oak woods; South Manor, pine barren. (70) South Manor, recently burned pine barren and young oak woods. (71) Center Moriches, black oak woods. (NE of 71) Eastport, graveyard. (72) Wading River (Wildwood State Park), black oak woods and bluffs. (73) Near Riverhead, pine - oak woods. (74) Calverton, oak - pine woods and maple swamp. (75) Calverton, pine - oak woods. (76) 2 mi 8 of Calverton, Bald Hill, pine woods. (77) Riverhead, black oak woods. (78) Riverhead, bogs and adjoining oak woods. (79) Riverhead, pine barren. (80) Quogue-Riverhead Rd., SW of Flanders, oak - pine woods. (81) Riverhead, pine - oak woods. (82) Eastport, gravel pit bog. (83) Speonk, pine - oak barren, adjoining maple swamp and sphagnum bog. (84) Remsenburg, black oak woods. (85) Riverhead, pine barren and young pine - oak woods. (86) Flanders, 113 Chamaecyparis bogs and pine barrens. (87) Hampton Bays, pine - oak woods and Chamaecyparis bog. (88) Quogue Station, oak woods. (89) Quogue, sand dunes. (90A) Northville, deep black oak woods. (90B) Mattituck, sand bluffs. (91) Laurel, oak - beech woods. (92) South Jamesport, oak woods. (93) Hampton Bays (Squiretown), young oak woods; Canoe Place, roadside ggpxa tomentosa. (94) Shinnecock Hills. (95) Southampton, Hudsonia - dune area. (96) Near Cutchogue bluffs. (97) Peconic, oak - hickory woods. (3 of 97) Peconic Station. (98) Southold or Laughing Waters, oak - hickory woods. (99) Noyack, oak- hickory woods. (99 - 111) North Haven. (100 A) Noyack, mature oak woods. (100 B) Sag Harbor, oak - hickory woods. (101) North Sea, open oak woods. (102) North Sea, Chamaecyparis bog and oak woods above bog. (103) Tuckahoe, open grassy field. (104) Bridgehampton, red maple swamp. (105) Sagaponack, sand dune. (106) East Marion, oak - cherry - locust woods and bluffs. (107) Shelter Island, Silver Beach, oak - hickory woods. (SE of 107) Shelter Island, Rt. 114 & Smith St., roadside. (108) Shelter Island, Ram Island neck, cherry - locust woods. (109) Shelter Island, Ram Island Drive, red cedar thickets. (110) Shelter Island, Ram Island, oak - maple woods. (111) Shelter Island, N of Nichols Point, open oak woods, beech - oak woods, beach area. (112) Northwest, oak - hickory woods; Three Mile Harbor, oak woods, open fields and woody bog. (113) Orient Point, red cedar woods and shores. (116) Orient Beach State Park. (117) Springs, oak - hickory woods and oak woods. (118) Springs, roadside. (119) Amagansett, oak - hickory woods. (120) Napeague, dunes and sand flats and sand barrens. (121) Promised Land, sand barrens, oak grove, cherry grove. (122) Gardiner's Island, field and old oak woods. (123) Gardiner's Island, south and, grassland. (124) Napeague, sand dunes and pine barrens. (125) Hither Hills State Park, pine barrens and dunes. (126) Hither Hills 114 State Park, exposed ridge and fresh pond. (127) Hither Hills State Park, 'mature oak woods and wooded sand bluffs. (128) Montauk, white oak - scarlet oak woods. (129) Montauk, grassy downs. (130) Montauk, low sand dunes. (131) Montauk Point, sand and ridges. (132) Montauk Point, woods. (133) Montauk, shaded Ilex verticillata thicket. (134) Fisher's Island. It is evident from the map in figure 15 that comparatively few collections ‘were made in western Long Island. In Brooklyn, Queens, and Nassau Counties, collecting areas were almost exclusively parks, preserves, or highway borders. Even in many parts of Suffolk County, particularly along its ‘western edge and along the north shore, the only more or less natural areas available for study were on large private estates where the owners were kind enough to allow explorations of their property. B. Additional specipgpp_examined. Several herbaria known to have large or significant Long Island collections were visited including the Brooklyn Botanic Garden (BKL) (Brainerd and Hulst collections), the New York Botanical Garden ( NY) (Torrey Cladonia collections), the Farlow Herbarium (FH) (early Latham collections and some Ramalina material in the Howe collections), the New York State Museum (NYS) (earliest Latham collections, many reported in Burnham and Latham [19141 and Charles Peck collections), the Evans herbarium at the U. 8. National Museum (US: Evans), the University of Michigan Herbarium (MICH) (Latham collections identified by Fink), the Missouri Botanical Garden (MO) (Latham collections identified by Dodge), and the Herbarium of the Staten Island Institute of Arts and Science (Staten Island) which contains several old and interesting Long Island specimens. A few specimens were also seen from the University of Tennessee herbarium (TENN) and the Cornell University herbarium (CUP). Lathamis personal herbarium is given the designation: (Latham). 115 C . Taxonomy . 1. Species concept. The problem of "what is a species," difficult as it is with any group of organisms, is compounded and confounded in lichens by the fact that two organisms are involved. In discussing lichens, there are two facets to the problem: 1) what do we mean by "lichen species" ... the consortium, or merely the lichen fungal component, and 2) the common problem of where does one species end and another begin. The first facet was, in theory, solved in 1950 when the International Code of Botanical Nomenclature added the statement ”for nomenclatural purposes names given to lichens shall be considered as applying to their fungal components" (Lanjouw, 1961, Art. 13, Note 4). Culberson (1961a) seriously challenged that position and maintained in a convincing series of arguments that the name of a lichen should apply to the entire lichen thallus ... fungus plus alga. His main arguments center around the fact that almost nothing is known about unlichenized lichen fungi and that the little that is known points to the fungi as being quite different in morphology, physiology, and ecology from the lichen as a whole. Since theoretically, the classification and identification of an organism is based on its own morphology, etc., Culberson asks, how can one apply a name to an organism based on the totally different morphology, physiology and chemistry of a thallus of which the organism in question is only a part. Although his arguments are well taken, I still believe a lichen name should refer to the fungal component alone. To say that one can only classify an organism divorced from all other membersof its biotic environment is not valid. Obligate parasitic fungi are studied only in relation with their host, and yet the taxonomy of parasitic fungi has not come to a halt because of it. If perchance it is found that a particular parasite looks 116 different or has different reactions on different hosts, what may be thought to have been several host specific species at one time can be considered to be one species later with no particular difficulty. Why should it be any different with lichen fungi? I believe that very few different lichens will be found to have the same lichen fungus. Recently, Uyenco (1963) showed conclusively that the morphology of the Coenogoniug lichen thallus, a lichen in which the alga is the dominant component, is due to the fungal component alone. She showed that the same lichen fungus growing symbiotically with different species of algae in different regions, will produce identical lichen thalli. Thus even thallus morphology can be interpreted as a fungus character. To say that lichen chemistry cannot be used to characterize the fungal component of a lichen is to disregard the genetic basis for the ability to synthesize a lichen acid. The lichen fungus is involved in the production of the chemical, and in all probability at most derives certain essential chemical precursors from the alga (Hess, 1959). With a growing knowledge of the biochemical role of the alga in a lichen thallus, we will probably be able to establish a system in which the unlichenized fungus can produce characteristic substances in culture. Again we see that a thallus character, in this case, chemistry, can be and probably is indicative of the genotype of the fungal component. i It therefore seems entirely proper to use thallus characters in character- izing a lichen fungus. It also seems proper to use the name of the fungal component of the thallus in routine references to the thallus as a whole. There is no need to allow lingual gymnastics to.confuse and complicate the process of communication. If it is convenient to use a fungus name to refer to the thallus which it characterizes in nature, so be it. All those 117 involved know what the name actually stands for and there is no advantage to encumbering discussions with constant references to "Parmelia sulcata and its associated algae," rather than just "Parmelia sulcata," the lichenized state being understood unless otherwise specified. There still remains the problem of how broad or narrow a species we should recognize in lichenology. In the absence of evidence for heterothallism in lichen fungi, objective fertility "tests" as applied in phanerogamic systematics, are not feasible, and so more or less subjective analysis is the only means left for taxonomic decisions. It has been pointed out that regarding lichens as functional Wapomicts" may have some merit, especially in speciation and phytogeographic considerations (John Beaman, personal communication). I think it is fairly obvious that generalizations concerning the relative merits of specific characters cannot be made. The presence of soredia is sometimes important, sometimes unimportant; certain lichen substances are more important in some groups and less important in others. This problem is discussed in some detail by Imshaug and Brodo (in press) and will not be elaborated on here. It suffices to say that the more information we have about a species and its close relative, i.e., the distribution, morphology, chemistry, etc., the easier it is to decide whether it is a species, deserves only intraspecific rank, or does not warrant taxonomic recognition at all. Thus, for Lecanora caesiorubella, the rank of subspecies was selected for recognizable segments of the species based on a great deal of information of all kinds. With less complete information, chemical segregates may have been considered "strains" or perhaps full species. Some species recognized here are done so tentatively pending a more extensive and intensive investi- gation of their group. Such species-pairs as Cladonia didypp - g, vulcanica, and Q, sgpamosa - g, atlantica, as well as the Q, subcariosa group, and 118 others need more work, but until that time, the narrow limits are recognized. In all too many cases, there is a serious question as to the status of a particular taxon. If there is still relatively little information available on which to base a firm decision, the previous treatment which I consider 'most authorative is followed. The individual systematic problems of various taxa are discussed in detail in the section VI - D. 2.Eoologhufl.f0rms. One of the most difficult tasks of the taxonomist is to determine the status of forms found in differing habitats and showing different morphological or chemical characters. For example, since both 'moisture and light are needed for assimilation, some sort of morphological and physiological compromises must have been met by the lichens in their adaptations to particular niches. But has the change in ecology produced the changes in morphology, or does the morphological variant represent a genetically stable entity confined to one ecological habitat? Weber (1962) recently tried to answer this question in dealing with the ecological modifications of some crustose lichens in southwestern United States. He stressed the need for extensive field experience and the examination of large numbers of specimens in making objective decisions. In some cases, the situation is fairly clear. Xanthoria parietina, for example, has a tendency to lose (or fail to develop?) its anthraquinone pigment in highly shaded places (Thomson, 1949; Barkman, 1958). Thalli growing on concrete blocks at Orient Point appeared bright yellowcorange on the exposed upper surface of the block and equally vigorous but a pale yellowish-white on the shaded side of the block. The change in habitat from strongly insolated to shaded (or the accompanying changes of dry to humid, and salt-sprayed to protected) apparently influenced the quantitative chemical differences. 119 Cladonia cristatella presents a somewhat similar situation. When in shaded woods, this species is highly branched and squamulose with a very low concen- tration of yellow usnic acid. In open sunny habitats, the species is sparsely branched, almost without podetial squamules, and very yellow with a high concentration of usnic acid. Increased photosynthetic area is an advantage in shaded localities, with the increased transpiration from the increased surface area being insignificant to the well-being of the thallus. In exposed areas, since light is not limiting and moisture is, the extra surface area provided by numerous podetial squamules is not needed and in fact would be disadvantageous and so is selected against. The production of extra pigment in highly illuminated habitats applies to melanin formation as well as usnic acid or parietin formation. Several species of Cladonia, particularly Q, furcata and g, atlantica show distinct and often intense browning when exposed to strong sun. Cetraria islandica subsp. crisps shows exactly the same response in the same situation. Quispel (1959) and Barkman (1958) suggest that lichen pigments in dry thalli may have a role in the protection of algae from high light intensities and it would therefore be logical to expect them to be in higher concentration in open areas than in shaded areas. The Peltigera canina group provides an example of a much more difficult problem. There is basic disagreement on the status of ecologically differing members of this group, particularly 3, canina sens. str. and g, rufescens. A dry, open, eroded habitat is characteristic of g, rufescens whereas a more cool, moist, mossy habitat is typical for 2, canina. {Thomson (1950a) maintains that there are all gradations from one type to the other, and that g, rufescens is merely an ecological form. Lindahl (1953) insists that the two are clearly separate species. He performed some transplant experiments with mature plants 120 and found that the transplanted thalli did not survive well, and those that did survive did not develop into the type characteristic of the new locality. Lindahl's transplantations were, unfortunately, not controlled with thalli transplanted to similar habitats so that the failure of his plants to survive in the new environment is not significant in itself. The lack of morphological change in a.msture plant is also to be expected since patterns of growth, once having reached a "point of no return" (Cantino, 1961), may be difficult if not impossible to alter. Transplantations of isidia or tiny squamules may prove to be valuable in determining the role of ecological conditions on the thallus forms. , 3.1nfrupecif1c taxa. The use of infraspecific categories in this paper is admittedly erratic. In general, varieties and forms are not considered and no new infraspecific taxa are described, although a few new combinations are made involving varieties. The few exceptions involve references to more clearly defined taxa which sometimes are considered as full species (such as Rhizocarpon obscuratum f. reductum) or thoroughly studied taxa which fit well into an infraspecific rank (such as Lecanora caesiorubella subsp. lathamii). The numerous varieties and forms described in Cladonia are not recognized since the large majority are undoubtedly growth forms and ecological variants, and the rest have been insufficiently studied. . 4.1(eys and annotated list. The arrangement of the flora into subclasses, and fwmilies, follows Hale (1961a) with the following exceptions: the Nephromaceae (after Wetmore, 1960), the Baeomycetaceae (after Rashnen, 1943), the Candelariscese (after Hskulinen, 1954), and the Teloschistacese and Physciacese (after Nannfeldt, 1932). The arrangement of genera within the families follows Zahlbruckner (1926). Species have been placed in alphabetica1_ order within the genera except for Cladonia which was arranged according to 121 Mattick's (1940) treatment. The keys have been somewhat expanded to include brief diagnoses of each species. In many cases, however, additional descriptive comments concerning certain important or confusing taxa have been included in the annotated list. The number after each species epithet refers to the page in the annotated list where that species is discussed. All author abbreviations follow Sayre, etafl.(1964). In the annotated list, all specimens listed under"heterial seen" or elsewhere in the discussions which were collected by Imshaug or Brodo have been deposited in the Michigan State University Herbarium (MSC), unless otherwise noted. The locations of other specimens have been recorded using standard abbreviations (Lanjouw & Stafleu, 1964). Comments on North American and worldwide distributions were made to provide a framework for the floristic treatments presented in section V. No attempt was made to compile a complete listing of all known localities. Instead, a limited number of fairly reliable, and in most cases, modern treatments were consulted to provide information on the basic distribution patterns and affinities. undoubtedly some of the records are based on old concepts or misidentifications and are incorrect; I hope that the errors are few. Individual state and province records are presented where specific state- ments or maps of distributions are not available. Sources of the North American records, unless stated otherwise, are as follows: Nova Scotia (Lamb, 1950), Maine (Degelius, 1940; Davis, 1964a,b), Connecticut (Evans & Meyrowitz, 1926; Hale, 1950), Massachusetts (Ahmadjian, 1958; personal collections), New Jersey (personal collections), central New York (Brodo, 1959), North Carolina (Degelius, 1941; Culberson, 1958), Tennessee (Degelius, 1941; Phillips, 1963), Alabama (McCullough, 1964), Arkansas, Missouri (Hale, 122 1957), Oklahoma (Hale, 1957; Thomson, 1961), Indiana(F1nk,& 11303,],919), Arizona (Darrow, 1950; Weber, 1963), New Mexico (Rudolph, 1953), Michigan (Hedrick & Lowe, 1936; Thomson, 1951), Wisconsin (Hale, 1955a,Culberson, 1955a), Minnesota (Fink, 1910), Black Hills (Wetmore, in prep.), Idaho (Hedrick, 1946), Washington (Howard, 1950), British Columbia (Weber & Shushan, 1959), Alaska (Cummings, 1910; Thomson, 1950; Krog, 1963), Northern Saskatchewan (Thomson & Scotter, 1961), Manitoba (Thomson, 1953), Ontario (Thomson, 1955; Ahti, 1964), Quebec (Thomson, 1955), Baffin Island (Hale, 1954b), Canadian archipelago and East Arctic (Thomson, 1960; Lynge, 1935, 1947). Unless otherwise stated, European records are based on Grummann (1963), Poelt (1963), or Zahlbruckner (1922-1940). Statements concerning circumboreal distributions are based on papers by Lynge (1928, 1938, 1940a, 1940b, 1940c), as well as the papers on the North American arctic cited above. Asian references are'all presented directly in the distributional notes. Species regarded as “endemic" are found only in North America, including the West Indies. KEY TO GROUPS 1. Thallus crustose: attached to substrate at all points; lower cortex absent (if podetioid, see Group III; if squamulose, see Group II) . . . . . . . . . . . . . . . . . . . . . . . . 2 1. Thallus at least partially free from substrate . . . . . . . 3 2. Thallus bearing ascocarps . . . . . . . . . . . . Group I (12$ 2. Thallus lacking ascocarps .. . . . . . . . . . . . Group II (129) 3. Thallus foliose: lobes flattened, usually broad, clearly dorsi-ventral, attached to substrate either directly or by means of rhizines, or rarely, only by a central umbilicus; lower cortex usually present; apothecia sessile or immersed, thallus never podetioid . . . . . . . . . . . . . . Group III (135) 3. Thallus fruticose: lobes more or less terete,‘or less frequently, flattened; basally attached to substrate at one or several points; pendulous, caespitose, or podetioid . . . . . . . . . . . . . . . . . . . . . Group IV (139) 123 GROUP 1 - CRUSTOSE LICHENS (FERTILE MATERIAL) l. Phycobionts blue-green algae. Thallus dark brown to black, areolate to subsquamulose, isidiate, prothallus blue-green or blue-black; apothecia lecideine; saxicolous on concrete.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Placythium nigrum (201) 1. Phycobionts green algae . . . . . . . . . . . . . . . . . . . . 2 2. Ascocarps on short hair-like stalks; hymenium disintegrating and spores forming a yellow-to deep-brown mazaedium. Spores brown (in water), spherical, ca. l-l.5 u in diameter . . . . . . . . . . . . . . . . . . . . Chaenotheca phaeogephala (189) 2. Ascocarp sessile or immersed; hymenium remaining intact . 3 3. Ascocarp i elongated; irregular or oblong . . . . . . . . . . . 4 3. Ascocarp disk-shaped, hemispherical or spherical (sometimes imbedded within a stroma) . . . . . . . . . . . . . . . . . . 8 4. Spores nonseptate, hyaline, 3-4 x 7-13 u. Lirellae dark brown to red brown or black, oblong or elongate, rarely branched, 0.2 x 0.35-0.55 mm . . . Xylographa opegraphella (196) 4. Spores 1-7 septate . . . . . . . . . . . . . . . . . . . 5 5. Spores with cylindrical cells; ascocarp sscolocular . . . . . 6 5. Spores with lenticular cells; ascocarp ascohymenial . . . . . 7 6. Ascocarp.i enclosed in a heavy carbonaceous stromatic wall . . . . . . . . . . . . . . . . . . . ... Qpegrapha (142) 6. Ascocarp without a carbonaceous stroma or excipuloid margin . . . . . . . . . . . . . . . . . . . . Arthonia (141) 7. Spores hyaline, 5-7 septate, 32-48 x 6-9 p; exciple not continuous below, but well developed laterally and projecting conspicuously above hymenium . . . . . . . . . . . . . . . . . Graphis scripts (197) 125 7. Spores brown, 2-3 septate, 21-30 x 6-7 p; exciple continuous below, shallow, i.e., not projecting appreciably above hymenium . . . . . . . . . . . . . . . . . . . . . . . . . . Phaeographis dendritica (197) 8. Ascocarp spherical or flask-shaped with walls completely enclosing hymenium except for ostiole at apex; walls generally partly or entirely carbonaceous . . . . . . . . 9 8. Ascocarp disk-shaped or cup-shaped, with exposed hymenium; or, hymenium enclosed within thalline tissue in a wart-like structure as in Pertusaria, without carbonaceous walls of any kind . . . . . . . . . . . . . . . . . . . . . . . . l6 9. Ascocarps clustered in stromatic verrucae, more than one per stroma . . . . . . . . . . . . . . . . . . . . . . . . . . . . lO 9. Ascocarps single, scattered . . . . . . . . . . . . . . . . . . 11 10. Thallus, especially stromatic verrucae, covered with a rusty-red pigment which is KOH + purple. (Spores brown, 3-septate, but not seen in L.I. material). Very rare . . . . . . . . . . . . . . . . . . . Malanotheca cruenta (193) 10. Thallus brownish or olivaceous, smooth, KOH-; spores hyaline, 4-8 septate, 23-45 x 8-13 u. Frequent on 1125, and Egggg.. . . . . . . . . . . . . . Trypethelium virens (193) 11.3p0res muriform, hyaline. Asocarp sscolocular . . . . . . . . . . . . . ;.. . . . . . . . . . . . . . . . . Polyblastiopsis quercicola(132) 11. Spores non-septate or only transversely septate . . . . . . . 12 12. Ascocarp sscolocular . . . . . . . . . . . . . . . . . 13 12. Ascocarp ascohymenial . . . . . . . . . . . . . . . . . 14 13. Spores ellipsoid or fusiform, straight, 15-21 x (4) 5-7 p, 1-3 septate; on oak and beach plum . . . . . . . . . . . . Arthopyrenia (141) 126 13. Spores acicular, curved or sigmoid, 20-30 x 3-4 p, 1-3 (5) septate; on birch . . . . . . . . . . . . . . . . Leptorhaphis epidermidis l4. Spores brown, 3-septate, cells lenticular, 16-20(-25) x 10-13 u . . . . . . . . . . . . . . Pyrenula nitida l4. Spores hyaline .. . . . . . . . . . . . . . . . . . . .. 15 15. Spores nonseptate. Saxicolous . . . . . . . . . . . Verrucaria 15. Spores 3-16 septate, cells cylindrical. Corticolous . . Porina 16. Spores more than 50 per sscus, 4 x 2 n. Saxicolous . . . . l7 l6. Spores (1)8(20) per sscus, usually larger than 4 x 2 u . 18 17. Thallus epilithic, areolate to squamulose, brown; apothecia completely immersed in thallus . . . . . . . .Agarospora fuscata 17. Thallus mostly endolithic; apothecia sessile with a lecideine margin . . . . . . . . . . . . . . . . . . . . . . . . . Sarcogype 18. Spores muriform . . . . . . . . . . . . . . . . . . . . 19 18. Spores nonseptate or transversely septate . . . . . . . . 21 19. Corticolous. Thallus thin, hypophloedal; apothecia minute, punctiform, 0.1-0.2 mm across; spores hyaline, 32 - 46(-55) x 10 - 23(-27) u. Rare . . . O I I O O O O O O O O O 19. Not corticolous. Thallus well developed; apothecia usually larger than 0.2 mm; spores brown or sometimes hyaline . . . . . . . . . 20 20. Apothecia deeply concave, imbedded in thick thalline verrucae resulting in a double margin (thalline and proper); spores without any gelatinous epispore ("halo"), 22-40 x 10 - 14 p. Medulla C+ red, KOH + yellow. Saxicolous or growing on Cladonia. Rare . h . . . . . . . 20. thalline verrucae or arising between them; spores with a gelatinous epispore ("halo"). Medulla C+ red or C-, KOH + Apothecia flat to convex with proper margin alone, imbedded in (181) (192) (143) (143) (269) Arthothelium’taediosum(186) . .Diploschistes scruposus (198) 21. 21. 23. 23 25. 25. 27. 27. 29. 29. 127 or -. Saxicolous. Common. . . . . . . . . . . . . Rhizocarpon Spores brown, uniseptate . . . . . . . . . . . . . . . . . . . 22 Spores hyaline . . . . . . . . . . . . . . . . . . . . . . . . 23 22. Apothecia with thalline margin . . . . . . . . . . Rinodina 22. Apothecia without thalline margin . . . . . . . . . Buellia Apothecia with thalline margin or enclosed in thalline verrucae.24 Apothecia without thalline margin . . . . . . . . . . . . . . . 31 24. Spores vermiform or sigmoid, septate or nonseptate, length to width ratio 7-9:l, 45-62 x 5-8 u. Thallus PD+ orange and KOH+ yellow (thamnolic acid). . . Haematomma ochrophaea 24. Spores ellipsoid, oblong, or subspherical, length to width ratio ca. l.5-3:1 . . . . . . . . . . . . . . . . . . . 25 Spores over 40 u long, nonseptate . . . . . . . . . . . . . . 26 Spores under 30 u long, nonseptate or septate . . . . . . . . 28 26. Apothecia usually imbedded in thalline verrucae, or, if lecanorine, then spores over 200 u long: 1 per ascus.27 26. Apothecia lecanorine; spores 40-68 u long, 8 per ascus. Disks C+ red . . . . . . . . . . . . . . . . . . Ochrolechia Spores all hyaline, KO -; spore walls not radiately channelled. Common . . . . . . . . . . . . . . . . . . . . . . . . Pertusaria Spores sometimes brownish, KOH + sordid violet, 125-190 x 30-45 u; spore walls conspicuously channelled. Uncommon. Mglanaria macounii 28. Spores polarilocular. Disk KOH + red-violet or KOH - . . . . . . . . . . . . . . . . . . . . . . Caloplaca 28. Spores nonseptate. Disk KOH - . . . . . . . . . . . . 29 Apothecial disk and margin yellow. Saxicolous . . . Candelariella Apothecial disk black, brown, pale reddish-buff, or yellowish. Saxicolous or corticolous. (If disk is yellowish, then (149) (177) (176) (294) (167) (163 (277) (174) (168) 128 corticolous) . . . . . . . . . . . . . . . . . . . . . . . . . 30 30. Phycobiont Trente ohlia; apothecia immersed in thallus. Spores hyaline, ellipsoid, 11-16 x 5-8 p. Saxicolous. . . . . ... . . . . . . . . . . . . . . . Ionaspis odora (277) 30. Phycobiont Trebouxia; apothecia immersed in thallus or sessile . . . . . . . . . . . . . . . . . . . . Lecanora (164) 31. Ascocarp sscolocular. Spores usually septate, hyaline, ellipsoid to fusiform . . . . . . . . . . . . . . . . . . . . . . Micarea (1&2) 31. Ascocarp ascohymenial . . . . . . . . . . . . . . . . . . . . 32 32. Spores septate . . . . . . . . . . . . . . . . . . . . 33 32. Spores nonseptate . . . . . . . . . . . . . . . . Lecidea (145) 33. Spores uniseptate . . . . . . . . . . . . . . . . . . . . . . 34 33. Spores 3 or more septate, fusiform to acicular . . . . . Bacidia (147) 34. Spores polariloculsr, 13-16 X 8-10 u. Disks KOH + purple-red . . . . . . . . . . . . . Caloplaca discolor (335) 34. Spores not polarilocular . . . . . . . . . . . . . . . 35 35. Apothecia black, strongly convex to hemispherical; hypothecium dark brown; spores with cells of unequal size, 9-15 X 4-5 p . . . . . . . . . . . . . . . . . . . . . . . . Qgtillaria glauconigrans (218) 35. Apothecia pale pinkish-yellow or orange, deeply concave to t flat; hypothecium hyaline; spores with cells of equal size, 9-14 x 2~4 u. Asci extremely narrow, almost linear, thin-walled . . . . .. . 36 36. Apothecial disks pink-yellow (flesh-colored), deeply concave . . . . . . . . . . . . . . . . . . . . . . . . Dimerella 911255_ (198) 36. Apothecial disks pale orange to orange buff, flat . . . . . . . . seeeoeeeeeeoeseeeeeeoeoeWill-g1 (199) GROUP II - CRUSTOSE LICHENS (STERILE MATERIAL) 1. Terricolous . . . . . . . ... . . . . . . . . . . . . . . . . . 2 1. Saxicolous . . . . . . . . . . . . . . . . . . . . . . . . . . 5 1. Corticolous or lignicolous . . . . . . . . . . . . . . . . . . 15 2. Thallus black to dark brown, minutely verrucose to granulose. Thallus KOH -, PD -, C -. . Lecidea uliginosa (216) 2. Thallus pale grey, grey-green, or white . . . . . . . . 3 3. Thallus C + red, KOH -, PD - . . . . . . . . . Lecidea guadricolor (214) 3. Thallus C - . . . . . . . . . . . . . . . . . . . . . . . . . 4 4. Thallus PD + deep yellow (baeomycic acid) . Baeomyces roseus (229) 4. Thallus PD -, KOH + yellow (atranorine). Pycnothelia papillaria(227) 5. 0n calcareous rock or mortar . . . . . . . . . . . . . . . . . 6 5. 0n siliceous rock . . . . . . . . . . . . . . . . . . . . . . 7 6. Thallus dark brown to black, isidiate, KOH -; phycobionts blue-green algae. Prothsllus conspicuous, blue-green . . . . . . . . . . . . . . . . . . . . . . . Plgcynthium nigrum (201) 6. Thallus yellow or orange, ROB + dark purple; phycobionts green algae. Thallus granular to thickly areolate and only occasionally breaking into sorediate patches; margin of thallus diffuse . . . . . . . . . . . . . Caloplaca citrina (334) 7. Thallus yellow or yellow-green, KOH - . . . . . . . . . . . . 8 7. Thallus white, grey, or brown (no yellowish tint) . . . . . . 9 8. Thallus margin effigurate; thallus yellow-green. Medulla C + red, usnic and gyrophoric acids present . .Rinodina oreina (346) 8. Thalli smell, scattered, areolate to subsquamulose, deep yellow. Medulla C -, usnic and gyrophoric acids absent . . I O C O O O O O O O O O O O O O O O candelariella Vitellina (296) 9 . mdulla c + red O O O O O O O O O O O O O O O 0 0 O O O O O 10 130 9. Medulla C - . . . . ... . . . . . . . . . . . . . . . . . . . . 12 10. Thallus grey, smooth, with scattered patches of soredia. Medulla KOH - . . . . . . . . . . . . . . (unknown no. 1)4 10. Thallus grey to ashy, esorediate, verrucose to areolate. Medulla KOH + yellow to orange & PD + orange (stictic acid) . . . . . . . . . . . . . . . . . . . . . . . . . . 11 11. Thallus brown, verrucose. Medulla I + blue . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Rhizocarpon grands (226) ll. Thallus grey to ashy, aerolate to verrucose. Medulla I - . . . . . . . . . . . . . . . . . . . . . . . Rhiggcarpon intermedium (226) 12. Thallus leprose to granular sorediate, marginate and often zoned. Thallus PD + red or yellow, KOH - or + yellow (?) (fumarprotocetraric or barbatolic acid present) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lepraria zonata (356) 12. Thallus smooth to areolate or verrucose. Thallus medulla KOH + yellow or red (stictic or norstictic acid present) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13 13. Thallus dark cinereous or sordid green-grey, verrucose to areolate. Stictic or norstictic acid present . . . . . . . . . gecanorg cinerea (282) 13. Thallus white to very pale grey or ashy, smooth to areolate. . .14 14. Prothsllus white, often conspicuous. Stictic or norstictic acid present. Growing on stones or boulders in shaded woods . . . . . . . . . . . . . . . . . Lecidea glbocaerulescegg (208) 4 These unidentified sterile crustose species have been deposited in herb. MBC for future reference. 131 14. Prothsllus black, often conspicuous. Norstictic acid present. Growing on exposed boulders. Pycnoconidia short, straight, bacilliform, 4-6 x ca. 1 u . . . . . Buellia stigmaea (342) 15. Thallus squamulose, margins entire. Undersurface of squamules sorediate . . . . . . . . . . . . . . . . . . . . . . . . . . 16 (also see Cladonia key) 15. Thallus continuous or diffuse (not squamulose) . . . . . . . 17 16. Thallus PD + red (fumarprotocetraric acid), C - . Squamules dark green-brown to olivaceous, 0.5 - 0.75 (-l.0) mm broad . . . . . . . . . . . Lecidea anthracophila (208) 16. Thallus PD -, C + red. Squamules pale olivaceous, 1.0 - l. 5 mm broad . . . . . . . . . . . . . . . Lecidea scalaris (215) 17. Thallus leprose, sorediate, or coralloid-isidiate . . . . . . 18 17. Thallus smooth, areolate, or verrucose . . . . . . . . . . . 32 18. Thallus orange, yellow, or yellowish-green . . . . . . 19 18. Thallus grey, grey-green, brown, olivaceous, or black . 21 19. Thallus dark yellow to orange, KOH + dark purple. Thallus smooth, becoming coarsely sorediate in patches . . . . . Caloplaca discolor (335) 19. Thallus yellow to yellowish-green, ROE - . . . . . . . . . . 20 20. Lignicolous, on decorticate Chamaecyparis stumps in bogs. Thallus diffuse, leprose, pale yellowish or whitish-green . . . . . . . . . . . . . . . . . . . . Chaenotheca phaeocephala (189) 20. Corticolous. Thallus leprose-granular, deep yellow . . . . . . . . . . . . . . . . . . . . . . Candelaria concolor (297) 21. Medulla KC + violet. Thallus dark cinerous to grey-green; verrucae erupting into white sorediate mounds . . . . . . . Pertusaria 29555} (270) Zlowdullaxc-OI‘KC'l'rBd............o..... 22 132 22. Thallus effuse, leprose, or coralloid-isidiste . . . . 23 22. Thallus with t,distinct soralia at least at thallus margin, verrucose or iLcontinuous . . . . . . . . . . . 27 23. Thallus coralloid-isidiate; phycobiont Trentepohlia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Porina nucula (195) 23. Thallus effuse, leprose, phycobionts Trebouxioid . . . . . . 24 24. Thallus bluish-green or blue-grey, KOH + yellow (atranorine), PD -, or rarely, PD + red (fumarprotocetraric acid) . . . . . . . . . . . . . . . . . . . Lepraria incana (355) 24. Thallus whitish-green or dark-green to blackish-green . 25 25. Thallus KOH + yellow & PD + orange (atranorine and stictic acid). Thallus pale to whitish-green, with thick white prothalline mat . . . . . . . . . . . . . . . . . . . . . . . . . . Lepraria sp. (357) 25. Thallus KOH -, PD - . .. . . . . . . . . . . . . . . . . . . 26 26. Thallus coarsely granular, pale green to brownish-green. Lignicolous. Very rare . . . . . . . . Lecidea viridescens (217) 26. Thallus finely granular, dark green to blackish-green. Mostly corticolous. Very common . . . Bacidia chlorococca (219) 27. Thallus composed of scattered verrucae or areoles, some bursting into soredia. . . . . . . . . . . . . . . . . . . . . . . . . 28 27. Thallus *‘continuous and smooth, at least at the margins . . 29 28. Medulla C + red . . . . . . . . . . . . Qgcideg seruginoss (212) 28. Medulla C - . . . . . . . . . . . . . . Lecidea botryosa (209) 29. Thallus XOR + deep yellow & PD + orange (thamnolic acid) . . .30 29. Thallus XOR -, PD - . . . . . . . . . . . . . . . . . . . . . 31 30. Thallus pale grey to white with crowded hollow verrucae in the older portions many of which burst revealing coarsely granular soredia often leaving the center of the thallus —m 133 essentially leprose . . . . . . . . . . . . Haematomma sp. (295) 30. Thallus ashy or darker, with sorediate verrucae scattered evenly over the thallus . . . . . . . Pertusaria trachythallina(274) 31. Thallus greenish or brownish-green with maculiform greenish or yellow-green soralia scattered over the thin thallus . .Qpegrapha sp. (142) 31. Thallus grey or greenish-grey, bursting into scattered, granular- sorediate soralia. Hypophloedal; phycobiont Trentepohlia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (unknown no. 6)4 32. Medulla KOH + yellow or red . . . . . . . . . . . . . . 33 32. Medulla KOH - . . . . . . . . . . . . . . . . . . . . . 35 33. Medulls KOH + yellow. Thallus thick or thin, pale grey to dark ashy . . . . . . . . . . . . . . . . . . . . . . . . . . 34 33. Medulla KOH + red (norstictic acid). Thallus thin, smooth, becoming areolate or chinky, pale greenish-grey to white; pycno— conidia 4-7 u long, straight, bacilliform . . . . . . . . . . . . . .. . . . . . . . . . . . . . . .Buellia_curtisii or g. stillingiana (340) 34. Medulla KOH + deep yellow & PD + orange (thamnolic acid). Thallus densely verrucose and rugose, pale grey . . . . . . . . . . . . . . . . . . . . . . . . .Haematomma ochrophaeum (294) 34. Mbdulls ROB + yellow-orange, PD - (?), thamnolic acid and atranorine absent, (stictic acid present?) thallus very thin, smooth, greenish-grey. Pycnoconidia short, straight, 4-5 x ca. 1 u . . . . . . . . . . . . .(unknown no. 7)4 35. Thallus olivaceous to blackish-green, well-developed, rugose to verruculose; pycnidia common, brown, pycnoconidia 0.5 x 1.2 u . o s s o e e e e s e e s e e e e s e e e o e s e 0 38611111 Chlorantha (219) 134 35. Thallus very thin or hypophloedal, or, if thicker, ashy or pale greenish-grey; pycnidia common, black; pycndconidia over 4 n long..36 36. Pycnidia t clustered in small groups; pycnoconidia 4-5 x l u, straight, bacilliform. On Ilex and Fagus..Tgypethelium virens (193) 36. Pycnidia scattered evenly over the thallus; pycnoconidia curved . . . . . . . . . . . . . . . . . . . . . . . . . . 37 37. Pycnoconidia reniform, short, broad, 5-7 x 3-4 p . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Opegrapha rufescens (189) 37. Pycnoconidia sickle-shaped, slender (1 u broad). . . . . . . . . 38 38. Pycnoconidia 10-15 u long (measured end to end, in a straight line), very strongly curved . . . . . . . Opegrapha cinerea (188) 38. Pycnoconidia 15-20 m long (measured as above), slightly curved . . . . . . . . . . . . . . . . (unknown no. 5)4 7. 7. GROUP III - FOLIOSE LICHENS Thallus composed of aggregations of squamules individually attached to the substrate at one edge; (0.5-)1-3(-5) mm long or broad . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cladonia (149) Thallus centrally attached; squamules, if present, part of a broad thallus; thallus over 10 mm in diameter . . . . . . . . . . . . 2 2. Phycobionts blue-green algae . . . . . . . . . . . . . . . 3 2. Phycobionts green algae . . . . . . . . . . . . . . . . . .7 Thallus gelatinous when moistened . . . . . . . . . . . . . . . 4 Thallus not gelatinous when moistened . . . . . . . . . . . . . 5 4. Upper cortex absent; globular isidia present; thallus broad . . . . . . . . . . . . . . . . .Collema subfurvum (199) 4. Upper cortex present, paraplectenchymatous; isidia absent or coralloid - cylindrical; thallus narrow-lobed .Leptogium (144) Thallus small, lobes 2-3 mm broad; apothecia scattered over the surface of thallus; spores non-septate. Lower surface densely white or tan, tomentose . . . . . . . . . . . . .Pannaria lggida’ (201) Thallus large, lobes 3-30 mm broad; apothecia at tips of lobes; spores septate . . . . . . . . . . . . . . . . . . . . . . . . . 6 6. Lower surface ecorticate,usually conspicuously veined; apothecia on upper surface of lobes; medulla KOH - . . . . . . . . . . . . . . . . . . . . . . .Peltigera (p.p.) (144) 6. Lower surface corticate, glabrous, without veins; apothecia on lower surface of lobes; medulla yellow, KOH + pink to red-violet (anthraquinone: nephromin). . Nephroma laevigatum (203) Thallus bright yellow or orange . . . . . . . . . . . . . . . . 8 Thallus brownish, grey,grey-green, or yellowish-green . . . . . .9 135 ll. 11. l3. 13. 15. 15. 136 8. Upper cortex KOH + red-violet (anthraquinone: parietin) . . . . . . . . . . . . . . . . . . . . . . . . . Xanthoria (176) 8. Upper cortex KOH - (pulvic acid derivative). Candelaria concolor(297) Thallus attached to the substrate by central umbilicus . . . . . 10 Thallus attached to substrate directly, or by many fine rhizines.12 10. Thallus yellow-green. Ascocarps apothecia with orange disks, abundant . . . . . . . . . . . . . . . . . . Lecanora rubina (288) 10. Thallus brown, with no yellow tint . . . . . . . . . . . . ll Ascocarps (usually present) perithecia; medulla C - . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dermafpcarpon miniatum(191) Ascocarps (if present) apothecia with black disks, medulla C + red . . . . . . . . . . . . . . . . . . . . . . . . . Umbilicaria (162) 12. Cephalodia abundant, scattered over the upper surface of thallus. Phycobiont Coccomyxa. Very rare; on soil . . . . . . . . . . . . . . . . . . . . . . . . . Peltigera aphthosa (204) 12. Cephalodia absent . . . . . . . . . . . . . . . . . . . . 13 Thallus inflated, hollow; lower surface corticate, brown to black, smooth, naked. Granular soredia in labriform soralia; medulla PD + red (monoacetyl-protocetraric acid), KC + red (physodic acid) . . . . . . . . . . . . . . . . . . . . . . gypggyggig,physodes (314) Thallus solid; lower surface rhizinate, tomentose, or ecorticate.l4 l4. Hypothallus present, composed of a thick mat of interwoven black hyphae. Medulla PD -, KOH -, KC - . . .Aggig’colpodes (321) 14. Hypothallus lacking . . . . . . . . . . . . . . . . . . . 15 Lower surface felt-like or tomentose, without rhizines. Lobes broad, over 3 mm across . . . . . . . . . . . . . . . . . . . . '16 Lower surface rhizinate . . . . . . . . . . . . . . . . . . . . 17 137 16. Medulla PD -, C -. Apothecia common, immersed in depressions in lobes; spores brown, uniseptate, 4 per ascus; phycobiont Coccomygg . . . . . . . . . . . . . . . . Solorina saccata (204) 16. Medulla + orange (stictic acid), or, C + red (gyrophoric acid). Apothecia, if present, sessile; spores hyaline, 3-septate, 8 per ascus; phycobiont Trebouxia . . . Lobaria (144) 17. Thallus yellow or yellow-green . . . . . . . . . . . . . . . . . . 18 17. Thallus brown, grey, or grey-green . . . . . . . . . . . . . . . . 20 18. Lower surface bright yellow; usnic acid absent. Thallus smooth or rugose; soredia and isidia absent; black pycnidia common along thallus margins, sometimes becoming partially laminal. Medulla PD -, KOH -, C -, KC - . .Cetraria viridis (320) 18. Lower surface not yellow; usnic acid present . . . . . . 19 19. Thallus with lobes less than 1 mm broad; older portions covered with granular soredia; divaricatic acid present. .Parmeliopsis ambigua (298) 19. Thallus with lobes broader than 1 mm; soredia present or absent; divaricatic acid absent. . . . . . . . . . . . . . . . . Parmelia (p.p.)(168) 20. Medulla PD +-orange, ROB + deep yellow (thamnolic acid) . . . . . . . . . . . . . .. . . . . . . . .Parmeliopsis(p.p.)(l68) 20. Medulla not having that combination of reactions (thamnolic acid absent) . . . . . . . . . . . . . . . . 21 21. Lower surface white, pale buff, or yellow . . . . . . . . . . . . 22 21. Lower surface light or dark brown, or black (although marginal areas may have broad, irregular, white blotches . . . . . . . . . . . . 26 22. Thallus brown or olivaceous - brown. Medulla C + red . . . . . . . . . . . . . . . . . . . Parmelia subaurifera (312) 22. Thallus grey or grey-green . . . . . . . . . . . . . . . 23 23. Thallus lobes 3-7 mm broad . . . . . . . . . . . . . . . . . . . . 24 138 23. Thallus lobes 0.5 - 3 mm broad . . . . . . . . . . . . . . . . . 25 24. Pseudocyphellae on upper surface; medulla I - . .Parmelia (p.p.) (168) 24. Pseudocyphellae absent; medulla I + blue..Cetraria tuckermanii (320) 25. Cortical hyphae parallel with surface . . . . . . . . . . Anaptychia (180) 25. Cortical hyphae perpendicular to surface . . . . . . . . . . Physcia (173) 26. Rhizines black with white tips, very dense; lobes 1-2 mm broad . . . . . . . . . . . . . . . . . . . . . . . . . 27 26. Rhizines uniform in color, sparse to dense; lobes l~6 mm broad . . . . . . . . . . . . . . . . . . . . . . . . . . 28 27. Medulla mustard-yellow, KOH + dull red-brown; lobes pruinose, especially near tips, with granular marginal soredia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pyxine sorediata (348) 27. Medulla red-orange (KOH + purple) or white (KOH -); lobes not pruinose; soredia marginal and laminal . . . . . Physcia orbicularis (350) 28. Pycnidis,common, marginal; rhizines sparse . . Cetraria (p.p.) (172) 28. Pycnidia rare, laminal; rhizines usually 1 dense. .Parmelia (p4x)(l68) GROUP IV - FRUTICOSE LICHENS Thallus having erect, terete or subterete podetia or pseudopodetia. Mostly terricolous, but sometimes corticolous or saxicolous . . . 2 Thallus not podetioid; erect and shrubby, or, more or less pendent, corticolous or saxicolous . . . . . . . . . . . . . . . . . . . . 5 2. Podetia or pseudopodetia hollow . . . . .. . . . . . . . . . 3 2. Podetioid structures solid . . . . . . . . . . . . . . . . . 4 Primary thallus squamulose or soon absent; spores nonseptate . ..... .Cladonia (14‘?) Primary thallus crustose, persistent, white granular; spores uniseptate. Medulla KOH + yellow (atranorine). . Pycnothelia papillarh1(227) Primary thallus consisting of white granules; podetia short, each one terminated by a large pink apothecium. Podetia and thallus PD + yellow, KC - (baeomycic acid). 0n raw eroding soil . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Baeomyces roseus (229) Primary thallus consisting of prong-like phyllocladia; pseudopodetia sterile or with brown apothecia. Medulla PD -, KC + red (lobaric acid). Saxicolous. . . . . . . . . . . . . . . Stereocaulon ssxatile (228) 5. Thallus composed of terete filaments . . . . . . . . . . . .6 5. Thallus composed of distinctly flattened or at least basally angular lobes or branches . . . . . . . . . . . . . . . . . 7 Thallus dark brown; filaments having no central cartilaginous axis. Soralia present. Medulla PD + red (fumarprotocetraric acid).A1ectoria (172) Thaflus yellow-green to grey-green; filaments with a central, cartilaginous, elastic axis. . . . . . . . . . . . . . . . . . '92223 (173) 7. Thallus light or dark brown, shrubby; terricolous. Marginal pycnidia abundant; pseudocyphellae linear, submarginal; medulla PD'- - . - - - - . . - wwwbsp- 2122a (318) 139 140 7. Thallus not brown; corticolous . . - - - - - - - - - - - - . - 3 8. Thallus yellow or orange. Cortex XOR + red-violet . . . . . Teloschistes (176) O O O O I O O O O O O O O O O O O O I C O O O O 8. Thallus grey-green or yellow-green. Cortex K- or K + yellow.9 9. Medulla C + red. Thallus isidiate, grey-green (usnic acid absent); clearly dorsi-ventral . . . . . . . . . . . . Pseudevernia furfuracea (316) 9. Medulla C -. Thallus not isidiate, yellow-green to grey-green (usnic acid present); upper and lower surfaces not distinguishable . . 10 10. Thallus soft, flexible (without chondroid layer), sorediate. Medulla KOH - , PD - . . . . . . . . . Evernia mesomorpha (322) 10. Thallus stiff, (with chondroid layer), often caespitose, esorediate. Medulla KOH - or KOH + red . . . . . Ramalina (172) ARTHOPYRENIA l. Spores ellipsoid to subfusiform, 15-17 x 5-7 p, 1~3 septate, with cells usually unequal in size; pseudothecia 0.15 - 0.26 mm in diameter; paraphysoid threads persistent, distinct. Corticolous. . . . . . . . . . . . . . . . . . . . . . . . . . . . A. pinicola l. Spores fusiform, 16-21 x 4-5 (-7) p, 1-3 septate, with cells equal in size; pseudothecia 0.15 - 0.25 mm in diameter; paraphysoid threads distinct and persistent. Corticolous. . . . . .A. cerasi ARTHONIA l. Phycobiont Trebouxia. Thallus whitish-to yellowish-green, granular to verrucose; ascocarps round; disks ashy grey to black, heavily pruinose; spores 3-septate, (14-) 16-22 x 5-7 p . . . . . A, caesia l. Phycobiont Trentepohlia . . . . . . . . . . . . . . . . . . . . . 2 2. Ascocarps jet black or bluish-grey (even when moist) . . . 3 2. Ascocarps red-brown to dark brown or brownish-black, turning a distinct red-brown when moistened . . . . . . . . . . . . 5 3. Hypothecium (fruit base) brown. Thallus scattered, granulose to disappearing; ascocarps punctiform; spores 3-septate i clavate, (181) (181) (183) 10-17 x 4-6 p e e e e s e e e e o e e e s e o o e e A. Cfrs mediella (184) 3. Hypothecium (fruit base) hyaline or essentially absent . . . . . 4 4. Spores (3-)5 septate, penultimate cells much shorter than other cells, 17-20 x 5-7 u . . . . . . . . . . . A, sexloculares (185) 4. Spores 3-septate, all spore cells equal in size, 14-20 x (4") 5-7 '1 e s s e o e e o e e e e s o o e s e e e A. Bunctifomis (184) 5. Spores 2-4 septate, hyaline, one end cell much larger than other cells; ascocarps epruinose; spores 14-20 x 5-7 u . . . . . . A. siderea (185) 141 142 5. Spores constantly 3-septate, ashy brown, all cells equal in size; ascocarps heavily pruinose; spores 12-17 x 4-6 p . . . Arthonia sp. (186) MICAREA l. Saxicolous. Thallus greenish, minutely verrucose to granulose; ascocarps less than 0.5 mm in diameter, buff to light brown; spores 3-septate, (8-)12-l6 x (2-) 3-4 u . . . . . . . Bacidia cfr. trisepta (224) l. Lignicolous (on rotting wood) . . . . . . . . . . . . . . . . . . . 2 2. Spores mostly uniseptate, sometimes nonseptate, 6.5-8.5 x 3.0— 3.5 u; thallus blackish-green, minutely granulose; ascocarps very convex to hemispherical, brown to black. . . . .M. prasina (187) 2. Spores 1-3 septate, 16-19 x 5-6 p; thallus dark green to greenish-black, smooth or verrucose to t granulose; ascocarps very convex to hemispherical, pitch black . . . . . M, melaena (187) OPEGRAPHA l. Thallus thin, continuous to scurfy or hypophloedal; spores 8 per ascus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 l. Thallus thin, becoming sorediate in maculiform yellow-green soralia; spores 4 per ascus. Ascocarps short and broad, 0.5-0.65 x 0.1l-O.4 mm, somewhat branches; spores 5-7 septate, 18-23 x 4-5 u . . .Qpegrapha sp. 2. Spores 22-36 x 2-3 p, 3-7 septate; ascocarps (0.25-)0.5-2 mm long, somewhat branched, pycnoconidia 9-15 x 1-2 p, strongly curved or twisted. . . . .‘. . . . . . . . . . . . . Q,cinerea (188) 2. Spores 19-24 x 3.5 p, 1-3 septate; ascocarps up to 0.5 mm long, unbranched; pycnoconidia 5-7 x 1-2 p, curved . . . . . . 00000000'00°'°""“°"”Q°w_(189) 143 VERRUCARIA l. Spores 6-9 x 3-5 p; perithecia 0.1 — 0.2 mm across; thallus very thin, filmy, sordid dark brown. On quartz pebbles in littoral zone . . . . . . . . . . . . . . . . . . . . . . . V, microspora (189) l. Spores 15-26 x 6-15 u; perithecia 0.2 - 0.4 mm across . . . . . 2 2. On littoral quartz pebbles. Thallus smooth, extremely thin, continuous, black to dark brown; spores 16-25 x 6-10 u . . . . . . . . . . . . . . . . . . . . . . . .‘V. silicicola (191) 2. On concrete and mortar. . . . . . . . . . . . . . . . . . 3 3. Thallus thick, dark brown to brownish-grey, dispersed verrucose, areolate to almost squamulose; exciple carbonaceous; spores 15-18 x 8-9 p . . . . . . . . . . . . . . . . . . . V, nigrescens (190) 3. Thallus thin, pale grey to whitish-ashy, areolate to chinky, the areoles being f dispersed; exciple pale; spores 20-23(-26) x 10-14 u . . . . . . . . . . . . . . . . . . . . . . y, muralis (190) PORINA l. Perithecia, buff to tan, 0.2-0.3 mm across; spores 5-9 septate, 48-75 x 7-9 a. Thallus effuse coralloid-isidiate; exciple pale . . . . . . . . . . . . . . . . . . . . . . . . . 2, 523313 (195) l. Perithecia black; spores less than 6.5 u broad . . . . . . . . . 2 2. Spores 3-7 septate, 30-42 x 5-6 p; exciple pale; thallus greenish-black, chinky to almost granulose, well developed . . . . . . . . . . . . . . . . . . . . . . . . g, cestrensis (194) 2. Spores mostly 9-13 septate, 58-65 x 5-7 p; excipule carbonaceous; thallus dark or light grey-green, diffuse, very thin, almost absent in places . . . . . . . . . . . . . . . . g, hibernica (1194) —[ 1. 144 LEPTOGIUM Thallus very thin, isidiate, the isidia cylindrical, becoming coralloid and subsquamulose; apothecia absent. . . . . . . . L, cyanescens (200) Thallus relatively thick, not isidiate or sorediate, but rather, rugose and finely rugulose; apothecia common; margins smooth and entire; spores 20-23 x 9-12 n . . . . . . . . . . . . . L. corticola (200) LOBARIA Thallus olivaceous, pitted and reticulate, with soredia and sometimes isidia on the ridges and margins. Sterile on L.I. Medulla PD + orange & KOH + yellow (stictic acid), C -, KC + reddish (lobaric acid?) . . . . . . . . . . . . . . . . . . . . . . . L, pulmonaria (202) Thallus grey to light green, smooth, without soredia or isidia. Usually fertile. Medulla PD -, KOH -, C + red . . . . L, guercizans (203) PELTIGERA Phycobionts green algae, cephalodia scattered over thallus surface. Rare. . . . . . . . . . . . . . . . . . . . . . . . . . 2, aphthosa (204) Phycobionts blue-green algae; cephalodia absent . . . . . . . . . 2 2. Thallus surface glabrous (without tomentum). Spores acicular 75-103 X 4-5 u . . . . . . . . . . . . . . . . . g. polydactyla (206) 2. Thallus surface tomentose to some extent . . . . . . . . . . 3 Thallus producing minute regeneration squamules at edges and along wounds . . . . . . . . . . . . . . . . . . . . . . . . g, praetextata (206) Thallus not producing regeneration squamoles . . . . (2, genius) 4 S. 5. l. 1. l. 3. 145 4. Thallus with grey granular soredia produced in small, laminal, orbicular soralia. . . . . . . . . . . . P, gaggga var. gpggig_ (205) 4. Thallus esorediate . . . . . . . . . . . . . . . . . . . . . 5 Veins on lower surface white . . . . . . . . . g, ggnig§_var. rufescens(205) Veins brown to the edge of the thallus . . . . L, canina var. ulorrhiza(206) LECIDEA On soil . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 0n rock . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 On bark or old wood . . . . . . . . . . . . . . . . . . . . . . . 8 2. Thallus green to greenish-grey or greenish-white, verrucose, becoming sorediate, C + red; apothecia 0.6-1.3 mm in diameter; hypothecium hyaline; spores 6-10 x 3-6 u . . . . L, guadricolor (214) 2. Thallus dark olivaceous-brown to black, granulose, C -; apothecia mostly 0.3 - 0.4 mm in diameter; hypothecium dark brown; spores (6-)8-10 x 4-7 u . . . . . . . . . L, uliginosa (216) Apothecia white pruinose with conspicuous dark grey rims. Thallus light grey to whitish-grey, continuous to irregularly cracked, KOH + red (norstictic acid) or KOH + yellow (stictic acid). Usually on shaded rocks . . . . . . . . . . . . . L, albocaerulescens (208) Apothecia black or brown, epruinose. Thallus KOH - . . . . . . . 4 4. Apothecia 0.5 - 1.5 mm in diameter; disks black . . . . . . 5 4. Apothecia less than 0.5 mm in diameter; disks black or brm O C O O O O O O O O O O O O O I O O O O O O O O O O O 6 Spores16-18x8u. . . . . . . . . . . . . . . .L.macroca_1;pa (213) Spores 7-12 x 3-6 p . . . . . . . . . . . . . . L, cfr. cygtidia (210) 6. Spores 11-20 x 7-10 u; apothedial disks red-brown to dark brown to black; hypothecium yellowish to hyaline. . . . L, coarctata (210) 146 6. Spores 6-8 x 3-4 p; apothecial disks black; hypothecium dark brown . . . . . . . . . . . . . . . . . . . . . . . . . 7 7. Epithecium and outer edge of exciple dark green to greenish- black . . . . . . . . . . . . . . . . . . . . . . . L. erratica (211) 7. Epithecium and outer edge of exciple reddish-brown, not green (but hymenium may be pale olivaceous at times). . . . L. cyrtidia (210) 8. Thallus squamulose, composed of imbricate squamules. Apothecia rare . . . . . . . . . . . . . . . . . . . . . . . 9 8. Thallus not squamulose. . . . . . . . . . . . . . . . . . . 10 9. Thallus C + red, PD - (lecanoric acid). Squamules mostly 0.5 - 1.0 mm across, yellowish- or olive-green . . . . . . . . . L, scalaris (215) 9. Thallus C -, PD + red (fumarprotocetraric acid). Squamules mostly less than 0.5 mm across, olive- to brownish-green to dark olivaceous brown . . . . . . . . . . . . . . . . . . L, anthracophila (208) 10. Hypothecium dark brown or reddish-brown . . . . . . . . . .11 10. Hypothecium hyaline . . . . . . . . . . . . . . . . . . . 12 ll. Thallus thick verrucose-areolate, becoming sorediate, grey-green to brown; spores 6-12 x (2-)3-5 u . . . . . . . . . . . .L, botryosa (209) ll. Thallus not sorediate, very thin, dark green-black; spores 6-8 x 3-4 . . . . . . . . . . . . . . . . . . . . . L, myriocarpoides (214) 12. Spores narrowly ellipsoid to fusiform, 11-19 x 3-5 u . . . 13 12. Spores ellipsoid to spherical, 5-10 x 3-7 p. Apothecial disks red-brown to black . . . . . . . . . . . . . . . . . 14 13. Spores 15-19 x 4-5 p, sometimes uniseptate; apothecia often strongly convex and hemispherical, disks flesh-colored to darker brown 0 . . . . . . . . . . . . . . . . . . . . . L, vernalis (217) 13. Spores 11-13 x 3-4 u, never uniseptate; apothecia I convex but not hemispherical, disks yellow to pale orange. . . . (see Lecanora syggicta)(289) 147 14. Thallus yellow-green, areolate to chinky or somewhat granular, C + yellow-orange. Apothecia red-brown to dark brown, usually less than 0.3 mm across; spores 7-10 x (4-)5-7 u . .L, varians (216) 14. Thallus grey-green to brownish-green, granulose to sorediate, C + red or C - . . . . . . . . . . . . . . . . . . . . . . 15 15. Spores subglobose, 5-6 x 3-6 u . . . . . . . . . . . . . L. nylanderi (214) 15. Spores ellipsoid or oval, 6-10 x 3-4 u . . . . . . . . . . . . . . 16 16. Thallus C + red. Apothecial disks lead black, margins prominant . . . . . . . . . . . . . . . . . . L, aeruginosa (212) 16. Thallus C -. Apothecial disks black, margins disappearing . . . . . . . . . . . . . . . . . . . . . L, viridescens (217) BACIDIA 1. Spores narrowly ellipsoid or narrowly ovate; ratio of length to width not more than 7:1 . . . . . . . . . . . . . . . . . . . . . . 2 1. Spores acicular, very narrow, ratio of length to width usually more than 7:1. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 2. Saxicolous. Apothecia minute, buff to light brown, convex; margin disappearing; spores (8-)12-l6 x (2-)3-4 u . . . . . . . . . . . . . . . . . . . . . . . . . . . .'L. cfr. trisepta (224) 2. Corticolous . . . . . L . . . . . . . . . . . . . . . . . . 3 3. Hypothecium dark brown; hymenium brownish; spores 16-20 x 5-6 u . . . . . . . . . . . . . . . . . . . . . . . . . . . .(see Micarea melaena) (187) 3. Hypothecium and hymenium.hyaline; spores (19-)23-32 x 3-6 p . . . . . .. . . . . . . . . . . . . . . . . . . . . . . Lgcidig_chlorococca (213) 4. Spores strongly curved and spiral-shaped, 13-16 x 2-3 u (measured end to end, in a straight line). Saxicolous. Rare 0 C O O O O O O O O O C O O O O O O O O O O 2. umbrina (224) IIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIiIIIIIIIIIIIIIIIIIIIIIIIIIIIII' 148 4. Spores 1 straight. . . . . . . . . . . . . . . . . . . . 5 Saxicolous. Disks usually lighter than margins; epithecium dark greenish-black to black; spores obscurely 3-septate, 19-28 x 1-2 p . . . . . . . . . . . . . . . . . . . . . . . . L. inundata (223) Corticolous . . . . . . . . . . . . . . . . . . . . . . . . . . 6 6. Hypothecium dark brown or red-brown . . . . . . . . . . .7 6. Hypothecium pale, hyaline, yellowish, or very light brown . . . . . . . . . . . . . . . . . . . . . . . . . 8 Apothecia small, 0.25 - 0.60 mm in diameter; spores obscurely 1-3 septate, 20-33 x 293 u; phycobiont Trebouxia . . . . L. chlorostica (220) Apothecia large, 0.75 - 1.25 mm in diameter; spores obscurely 6-9 septate, 35-55 x 3-4 u; phycobiont Trentepohlia . .‘L. schweinitzii (223) 8. Polysporous; thallus thick, coarsely granular to verrucose, dark green to olive; apothecia irregular, up to 1.25 mm in diameter, often clustered and crowded; margins raised, conspicuous. Spores 5-7 septate, 20-26 x 2-3 p . . . . . . . . . . . . . . . . . . . . . . . . . L, chlorantha (219) 8. Octosporous; thallus thin, greenish-grey; apothecia smaller, round, not clustered; margins not raised. . . . . . . . 9 Disks red-brown to black; margins concolorous or lighter, disappearing with age; epithecium reddish-violet (intense in KOH); spores 7 to many septate, 39-68 x 4-6 u . . . . . L, atrogrisea (218) Disks light buff to 1' dark brown (never black); margins darker than disks; epithecium brown; spores obscurely 3-4 septate, 19-32 x 1-3p.......................L.intermedis (221) l. 1. 3. 3. 149 RHIZOCARPON Spores uniseptate, hyaline to slightly tinted, (ll-)l3-20 x (5-)6-10 u. Thallus sordid greyish-green to ashy, verrucose to minutely verruculose, KOH + red (norstictic acid) or KOH + yellow (stictic acid) . . . . . . . . . . . . . . . . . . L. cinereovirens (225) Spores muriform or submuriform, or thallus sterile. . . . . . . . 2 2. Medulla C + red (gyrophoric acid?), KOH + yellow (stictic acid).Spores soon dark brown, many celled. . . . . . . . . . 3 2. Medulla C -, KOH + red or KOH -. Spores hyaline for a long time, then brown . . . . . . . . . . . . . . . . . . . . . . 4 Medulla I -. Thallus whitish to light ashy or brownish-grey, subcontinuous to areolate, and finally verrucose; spores 26-38 x 10-15 u . . . . . . . . . . . . . . . . . . . . . . . L, intermedium (226) Medulla I + blue. Thallus dark brown or grey brown, verrucose with * round, t scattered verrucae; spores 25-29 x 10-13 u . . .L, gggggg_(226) 4. Medulla KOH + red (norstictic acid). Thallus t smooth, thin; apothecia without any indication of a thalline margin; spores 20-27x10-13u................. L.plicatile (227) 4. Medulla KOH - . Thallus verrucose or areolate, almost squamulose in places; apothecia immersed in small areoles giving appearance of a thalline margin; spores 19-29(-32) x 8-16 u . . . . . e e e e e o s e s o e e e s e e e e s e e e o _R_e Obscuratum (226) CLADONIA Primary thallus crustose, persistent, consisting of grey-green or grey to whitish verrucae or granules; spores uniseptate. PseudopOdetia usually under 0.75 mm tall, molariform to somewhat 150 branched, often inflated. Pseudopodetia KOH + yellow & PD - (atranorine) . . . . . . . . . . . . . . . . Pycnothelia papillaria (227) Primary thallus squamulose or absent in mature plant; spores nonseptate . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 2. Podetia forming a more or less complex branch system (shrubby); primary thallus disappearing in maturity . . . . . . . . . .3 2. Podetia simple or sparingly branched, or EEEEEEJ primary thallus squamulose, persistent. . . . . . . . . . . . . . . . . . 15 Podetia corticate, except where cortex is replaced by soredia in the sorediate species . . . . . . . . . . . . . . . . . . . . . . 4 Podetia ecorticate, esorediate (Subsection CLADINA) . . . . . . . 9 4. Podetia yellowish, usnic acid present (Subsection UNCIALES).5 4. Podetia grey-green to brownish, usnic acid absent (Subsection CHASMARIAE) . . . . . . . . . . . . . . . . . . 7 Csrtilaginous cylinder forming an unbroken inner lining of the podetia, with tiny white granules resembling pruina; cortex smooth and shiny; podetia slender (dry habitats) or robust (moist habitats). Medulla UV + blue-white (squamatic acid). . . . . . . . . g, uncialis (257) Csrtilaginous cylinder more or less discontinuous or fibrous; cortex not smooth nor shiny. Medulla UV - (squamatic acid absent). . . 6 6. Podetia inflated, contorted, perforate; cartilaginous cylinder composed of loosely interwoven strands; medullary hyphae (as seen in podetial cross-sections) loose and anastomosing, (5-)6-8 u in diameter. . . . . . . . . . . . . . . . . 9, L251; (256) 6. Podetia not inflated or perforate; cartilaginous cylinder composed of closely interwoven strands; medullary hyphae compact, 3-5(-7) u in diameter . . . . . . . . . . . . . Q, caroliniana (256) 151 7. Soredia present, especially at podetial tips; podetia usually sparsely branched . . . . . . . . . . . . . . . . . . . . . . . 8 7. Soredia absent; podetia intricately branched . . . . . . Q, furcata (255) 8. Soredia usually farinose, scattered in irregular patches over much of the podetium, gradually coalescing into a continuous sorediate area; squamules confined to the lower half or third of the podetium, or absent. . . . . . . Q, farinacea (253) 8. Soredia granular, mostly confined to the tip of the podetium; squamules commonly covering entire podetium. Rare . . . . . . . . . . . . . . . . . . . . . . . . Q, scabriuscula (253) 9. Branching more or less isotomic, distinct main stems absent or only exceptionally present; plants giving a rounded, tufted appearance. Thallus PD- . . . . . . . . . . . . . . . . 10 9. Branching anisotomic, distinct main stems usually present. . . .11 10. Thallus yellowish, KOH - (usnic acid present, atranorine absent); tetra- to pentachotomies predominating, usually star-shaped around an axillary hole . . . . . . g, alpestris (259) 10. Thallus grey, or rarely somewhat yellowish, KOH + yellow (usnic acid absent, atranorine present); di- or trichotomies predominating, axils generally closed. Surface appearing very rough, almost tomentose; algal layer not continuous . . . . . . . . . . . . . . . . . . . . . . . . . g, evansii (258) ll. Thallus PD - . . . . . . . . . . . . . . . . . . . . . . . . . 12 11. Thallus PDw+ red (fumarprotocetraric acid) . . . . . . . . . . 13 12. Branches very robust, often sprawling; axils broadly open; branching usually tetrachotomous with dichotomies rare; algal layer very smooth and compact appearing almost corticate. ‘ ,7 IIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIT 14. 14. 16. 16. 18. 18. 152 Pseudonorangiformic acid present, atranorine absent. Very common . . . . . . . . . . . . . . . . . . . . Q, submitis (263) 12. Branches usually slender, always erect; axils often closed or only slightly open; branching usually trichotomous with dichotomies common; algal layer smooth or decomposed. Pseudonorangiformic acid absent. Very rare . . . . L, EiEii (265) 13. Branching predominantly dichotomous, tri- and tetrachotomies rare; branchlets usually very slender, erect; axils infrequently open; main stems often indistinct; pycnidial jelly red . . . . . . . . . . . . . . . . . . . . Q, subtenuis (260) 13. Branching predominantly tri- and tetrachotomous around widely open axils; branchlets robust, falcate; main stem always distinct; pycnidial jelly colorless . . . . . . . . . . . 14 Thallus blue-grey. Usnic acid absent, atranorine present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . L, rangiferina (262) Thallus grey-green to yellowish-grey. Usnic acid present, atranorine absent . . . . . . . . . . . . . . . . . . . . . . . . Q, arbuscula (263) 15. Podetia and apothecia absent . . . . . . . . . . . . . . .16 15. Podetia and/or apothecia present . . . . . . . . . . . . .36 Medulla PD + red, orange, or yellow . . . . . . . . . . . . . . .17 Medulla PD - . . . . . . . . . . . . . . . . . . . . . . . . . 29 17. Medulla PD + red (fumarprotocetraric acid) . . . . . . . .18 17. Medulla PD + yellow or orange . . . . . . . . . . . . . . 23 Squamules sorediate on lower surface, broad, entire to broadly lobed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . l9 Squamules esorediate on lower surface. . . . . . . . . . . . . . 20 19. Squamules large, over 1.0 mm broad, ascending..§, coniocraea (244) 153 19. Squamules minute, 0.5 - 0.75(-1.0) mm broad, closely appressed . . . . . . . . . . . . . . Lecidea anthracophila (208) 20. Margins of squamules finely divided to T granulose . . . . . . .21 20. Margins of squamules entire to broadly crenate . . . . . . . . .22 21. Grayanic acid (or, very rarely, cryptochlorophaeic acid) present (in Long Island material) . . . . . . L. chlorophaea (241) 21. Grayanic and cryptochlorophaeic acids absent. .9, caespiticia (249) 22. Atranorine present. Squamules strap-shaped; margins somewhat revolute . . . . . . . . . . . . . . . . . . . . . . Q, apodocarpa (248) 22. Atranorine absent... . . '. . .‘. . . .'. .‘. . . . . . . . . . . . . - . . Q-WQo-W Q-vmnemtha. (1.2mm 23. Medulla C + green . . . . . . . . . . . . . .'§. strepsilis (236) 23. mdUIIC C - e s s e e e e e e s o e o o e o e o s e e e 24 24. Medulla KOH + blood red (norstictic acid) . . . . . g, subcariosa (237) 24. Medulla KOH - or KOH + yellow . . . . . . . . . . . . . . . . . 25 25. Medulla KOH + deep yellow and PD + orange (thamnolic acid) . . . . . . . . . . . . . . . . . . . . . . . . . 26 25.MedullaKOH-,PD+yellow. . . . . . . . . . . . . . . .28 26. Lower surface of squamules sorediate. Terricolous, corticolous, or lignicolous . . . . . . . . . . . . . . . . . . . .Q, magilggga (231) 26. Lower surface of squamules esorediate. Lignicolous . . . . . . 27 27. Margins of squamules granulose, sometimes reducing the primary thallus to a granular crust . . . . . . . . . Q, parasitica (249) 27. Margins of squamules finely divided, not granulose . . . . . . . . . . . . . . . . . . . . . . . . . . g, vulcanica (232) 28. Squamules entire or crenate. Squamatic and baeomycic acids absent, psoromic acid present . . . . . . . . . . . . . . . . . g, brevis (239) 28. 30. 30. 32. 32. 34. 34. 36. 154 Squamules finely divided. Squamatic and baeomycic acids present, psoromic acid absent . . . . . . . . . g, atlantica or Q, beaumontii (251) 29. Thallus C + red. Lower surface of squamules sorediate . . . . . . . . . . . . . . . . . . . . . . . . . . Lecidea scalaris (215) 29. Thallus C - . . . . . . . . . . . . . . . . . . . . . 30 Upper surface or lower surface of squamules yellow or yellowish (usnic acid present) . . . . . . . . . . . . . . . . . . . . . 31 Upper surface of squamules grey to grey-green, lower surface white (usnic acid absent or not detectable). . . . . . . . . . . . . 33 31. Lower surface of squamules sorediate. Squamatic acid present, barbatic acid absent . . . . . . . . . . . . . . g, incrassata(233) 31. Lower surface of squamules esorediate. Squamatic acid absent, barbatic acid present. . . . . . . . . . . . . . 32 Squamules very large, broadly crenate to strap-shaped; lower surface yellowish. Didymic acid absent . . . . . . . . g, robbinsii (235) Squamules small, usually finely divided; lower surface white. Didymic acid present . . . . . . . . . . . . . . . . . Q, cristatella (233) 33. Lower surface of squamules T'sorediate. Barbatic acid present, didymic and squamatic acids absent. . .g, bacillaris (230) 33. Lower surface of squamules esorediate . . . . . . . . . 34 Squamatic acid present, didymic acid absent . . . . . . Q. sguamosa (250) Squamatic acid absent, didymic acid present . . . . . . . . . . 35 35. On highly decayed wood in shaded bogs . . . . . . . Q, glgymg (232) 35. On soil, dry tree bases, or dry lignum in exposed areas. . . . . . . . . . . . . . . . . . . . . . . . . . g, cristatella (233) Apothecia essentially sessile on primary squamules or on very short decorticate podetia (less than 2 mm tall); squamules finely crenate; 155 apothecia brown, flat to strongly convex. Squamules PD + red (fumarprotocetraric acid) . . . . . . . . . . . . . g, caespiticia (249) 36. Apothecia, when present, on I well developed podetia at least partially corticate and over 2 mm tall; podetia often sterile . . . . . . 37 37. Podetia without cups or tiers . . . . . . . . . . . . . 38 37. Podetia with more or less distinct cups or tiers . . . . 63 38. Podetia without soredia or granules, although in some cases somewhat ecorticate. . . . . . . . . . . . . . . . . . . . . . .39 38. Podetia granular or with granular or farinose soredia. . . . . 53 39. Apothecia red; podetia and squamules yellowish-green to grey-green (usnic acid present). . . . . . . . . . . . . 40 39. Apothecia brown, tan of buff, or absent; podetia and squamules grey-green to olive-green or yellowish-green (usnic + or - ) . . . . . . . . . . . . . . . . . . . . 41 40. Primary squamules covered on lower surface with granular or farinose soredia. Common on decaying stumps and logs. .Q, incrassata (233) 40. Primary squamules without soredia. Very common on many substrates. Podetia usually grey-green, squamulose on bark in the shade, and yellow-green without squamules on the ground in the sun..L, cristate11a(233) 41. Podetia more or less abundantly branched. Podetia PD + red or orange . . . . . . . . . . . . . . . . . . . . . . . 42 41. Podetia usually simple, or, if branched, only once or twice near the summit. Podetia PD + or - . . . . . . . . . . .43 42. Podetia often growing in dense mats, 10-20 mm tall, often bearing brown apothecia; holes in axils often surrounded by proliferations giving the appearance of rudimentary cups. Podetia KOH + lemon yellow, PD + red-orange or orange-yellow (thamnolic acid). . . . . . Q, floridana(252) 156 42. Podetia not growing in dense mats, usually over 20 mm tall; apothecia rare; holes in axils never surrounded by proliferations. Podetia KOH - or brownish, PD‘+ red (fumarprotocetraric acid). . g, furcata (255) 43. Podetia PD + . . . . . . . . . . . . . . . . . . . . . . 44 43. Podetia PD - . . . . . . . . . . . . . . . . . . . . . . 51 44. Podetia PD + red (fumarprotocetraric acid). . . . . . . . . . . 45 44. Podetia PD + yellow to orange. . . . . . . . . . . . . . . . . .47 45. Thallus yellowish-green to gray-green; podetia 7-15 mm tall, minutely squamulose; apothecia minute, present or absent. Very rare. . . . . . . .j. . . . . . . . . . . . . Q, simulate (248) 45. Thallus dark or pale green-grey; podetia usually less than 10 mm tall, not squamulose; apothecia always present, large, at least equal to diameter of podetium. Common. . . . . 46 46. Podetia usually grooved and twisted, often decorticate, often longitudinally split or striate; apothecia buff to light brown, two to three times the diameter of the podetium. . . . . . g, capitata(236) 46. Podetia usually corticate, verrucose or areolate, not twisted or striate; apothecia dark or sometimes light brown, one to two times the diameter of the podetium. . . . . . . . . . . g, clavulifera (238) 47. Medulla C + green, KOH - (strepsilin and baeomycic acid). Thallus and podetia olive-green; podetia T inflated. . . . . . . . . . . . . . . . . . . . . . . . . . . . . Q, strepsilis(236) 47. Medulla C -, KOH + or - (strepsilin absent). Thallus and podetia grey-green or brownish-green; podetia usually slender. . . . . . . . . . . . . . . . . . . . . . . . . 48 48. Podetia with perforate tips or axils, covered with large or small squamules or verrucae. . . . . . . . . . . . . . . . . . . . . 49 157 48. Podetia not perforate, without squamules, or, slightly squamulose on lower half. . . . . . . . . . . . . . . . . . . . . . . . . 50 49. Podetia ROH - (baeomycic acid present). Podetia commonly over 10 mm tall, slender, grey-green. . . . . g, beaumontii (252) 49. Podetia KOH + yellow (thamnolic acid present). Podetia usually under 10 mm tall, robust, pale grey to almost white . . . . . . . . . . . . . . . . . . . . . . . . . .L, santensis (250) 50. Medulla KOH + red (norstictic acid) . . . . . . . . . g, subcariosa (237) 50. Medulla KOH - (psoromic acid) . . . . . . . . . . . . . g, brevis (239) 51. Thallus with a distinct yellow tint (usnic acid present).52 51. Thallus without any hint of yellow (usnic acid absent). Thallus grey or brownish-green; podetia commonly 10-15 mm tall, fissured. Atranorine present. Very rare . . .L, cariosa (237) 52. Primary squamules small (mostly less than 0.5 mm broad); podetia common; apothecia flat, reddish-brown, abundant. Rare . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Q, piedmontensis (235) 52. Primary squamules very large (1-4 mm broad); podetia rare, very short, arising from lateral edges of squamules; apothecia strongly convex, dark brown. Very rare.. . . . . . . . . . . . . g, robbinsii (235) 53. Podetia PD + . . . . . . . . . . . . . . . . . . . . . . 54 53. Podetia PD - . . . . . . . . . . . . . . . . . . . . . . 61 54. Podetia PD + yellow to deep yellow-orange, KOH +-lemon yellow (thamnolic acid). . . . . . . . . . . . . . . . . . . . . . . . 55 54. Podetia PD + deep red, KOH - or + dingy brown (fumarprotocetraric acid) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 57 55. Apothecia brown to purple-brown, common. Podetia and margins of primary squamules covered with large corticate granules. . . . . . . . . . . . . . . . . . . . . g, parasitica(249) — 158 55. Apothecia red, but sometimes lacking . . . . . . . . . . . 56 56. Primary squamules esorediate; podetial soredia coarsely granular; podetia often decorticate and transluscent with cartilaginous layer exposed. O C O O O O O O O O O 0 O O O O O O O O O I O 9.0 vulcanica (232) 56. Primary squamules sorediate; podetial soredia farinose or rarely granular, covering podetium; podetia often decorticate turning brown to black, but opaque. . . . . . . . . . . . . . . g, macilenta (231) 57. Podetia short, rarely taller than 6 mm, with blunt spices, covered with coarsely granular soredia on the lower 1/2 to 2/3 of podetium, and farinose soredia on the upper 1/2, ecorticate areas abundant. Grayanic acid present . . . e e e o e e o s e e s e s e e o o s o e s o e 0 £0 C211ndrica (246) 57. Podetia usually much taller than 6 mm, spices sharply pointed, * corticate at the base, corticate on upper parts. Grayanic acid absent . . . . . . . . . . . . . . . . . . . . . . . 58 58. Podetia partially decorticate, the decorticate areas becoming pellucid and dark; granular soredia covering large portions of the podetia. Podetia often bent or contorted. . . . . . . . . . . . . . Q, pityrea (247) 58. Podetia corticate for the most part, or the cortex is replaced by granular or farinose soredia. . . . . . . . . . . . . . . . . . . 59 59. Podetia unbranched, relatively stout, tapering T abruptly to a sharp point, each podetium.arising from the center of a primary squamule. Podetia and squamules with a vague yellowish- green tint; farinose sorediate on upper half or more of podetium; squamules large, sometimes sorediate. Common and variable . . . . . . . . . . . . . . . . . . . . . . . . . . C. coniocraea (244) 59. Podetia commonly branched, long and slender, not arising from the center of primary squamules. . . . . . . . . . . . . . 60 60. 60. 62. 62. 64. 64. 66. 159 Soredia usually farinose, scattered in irregular patches over much of the podetium, gradually coalescing into a continuous sorediate area; squamules confined to lower half or third of the podetium, or absent. Frequent. . . . . . . . . . . . . . . . . . . Q, farinacea (253) Soredia granular, mostly confined to tip of podetium; squamules commonly covering entire podetium. Rare. . . . . . . . g, scabriuscula (253) 61. Podetia corticate for most of length, some areas bursting into granular soredia; some granular soredia on lower surface of squamules near the margins; apothecia red. Usnic acid present or absent. Rare. . . . . . . . . . . . Q, floerkeana (230) 61. Podetia mostly sorediate, often with many decorticate areas; apothecia red. Usnic acid absent. . . . . . . . . . . . 62 Podetia covered with granular soredia, or soredia becoming farinose on upper half; decorticate areas becoming transluscent, then brown; primary squamules esorediate. On wood or bark. Rare. . . .9, gigygg_(232) Podetia entirely covered with farinose soredia, occasionally with a small corticate area at the base; decorticate areas white, opaque; primary squamules usually having granular soredia on lower surface near the margins. 0n various substrates. Very common. . .9, bacillaris (230) 63. Podetia without soredia or granules, but sometimes squamulose or minutely verrucose. . . . . . . . . . . . . . . . . . . 64 63. Podetia sorediate or granular. . . . . . . . . . . . . . . 72 Cups opening into podetia. . . . . . . . . . . . . . . . . . . . .65 Cups closed by continuous membranes . . . . . . . . . . . . . . . 69 65. Podetia KOH + deep yellow (thamnolic acid). . . . . . . . .66 65. Podetia KOH - . . . . . . . . . . . . . . . . . . . . . . .67 Cups very narrow, almost rudimentary, slightly perforated at tip. Rafa. . s e e o o s o o o o o o o o o c o o o o o o o o s 20 Banten818 (250) 160 66. Cups broad, with extensive proliferations. . . . . . . Q, gagggggggig (256) 67. Podetia PD + . . . . . . . . . . . . . . . . . . . . . . . 68 67. Podetia PD - (squamatic acid present). . . . . . g, ESEEEQEE. (250) 68. Podetia PD + yellow (baeomycic and squamatic acids present). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . g, atlantica (251) 68. Podetia PD + red (fumarprotocetraric acid). Podetia and cups irregularly perforate and lacerate. . . . . . . . . . Q, multiformis (253) 69. Podetia proliferating from center or edges of cups; cups shallow, flat, or slightly convex; podetia corticate. . . .70 69. Podetia simple, deeply goblet-shaped, not proliferating, extensively decorticate. Inside of cup lined with small or large scattered areoles or flat squamules. Grayanic acid absent . . . . . . . . . . . . . . . . . . . . . g, pygidata (241) 70. Proliferations irregular, mostly from cup edges; cups abortive, T squamulose; squamules large. . . . . . . . . . . . . . g, mateocyatha 70. Proliferations from center of cups, regular; cups well formed; podetia esquamulose . . . . . . . . . . . . . . . . . . . . . . . . . . . 71 71. Cups gradually expanding from stalk; podetia usually completely corticate. On neutral soils. . . . . g, verticillata(239) 7l. Cups abruptly expanding from stalk; podetia with a 1 continuous cortex becoming distinctly areolate or partially decorticate. On acid soils, especially in or near bogs. . . . . Q, calycantha(240) 72. Podetia distinctly yellowish-green (usnic acid present or absent). 73 72. Podetia grey-green or brownish (usnic acid absent). Apothecia brown. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 76 73. Podetia PD + orange & KOH + yellow (thamnolic acid); usnic acid absent. Podetia with narrow, shallow cups, corticate at base, soon becoming farinose sorediate and sorediate for 161 most of length. (On Nantucket Island, not on Long Island) . . . . . . . . . . . . . . . . . . . . 9, digitata 73. Podetia PD -, KOH - ; usnic acid present. . . . . . . . .74 74. Apothecia brown. Barbatic acid present, zeorine absent. Cups deep, goblet-shaped, covered with farinose soredia. . . .9, carneola (234) 74. Apothecia red. Barbatic acid absent, zeorine present . . . . . 75 75. Cups often elongate, somewhat split longitudinally; soredia farinose. Rare . . . . . . . . . . . . . . . . . 9, deformis (233) 75. Cups goblet-shaped, not split; soredia coarsely granular. Common. . . . . . . . . . . . . . . . . . . . . . 9, pleurota (234) 76. Soredia coarsely granular, covering entire podetium. Podetia PD + red (fumarprotocetraric acid) or PD - ; grayanic or, rarely, cryptochlorophaeic acid present. . . . . . . . . . . . . 9. chlorophaea(24l) 76. Soredia farinose. Podetia PD + red (fumarprotocetraric acid). .77 77. Cups shallow, deeply dentate, with short spur-like branchlets proliferating from edges giving a star-like appearance, or, infrequently, these proliferations are lacking. Homosekakaic acid present (but often difficult to demonstrate) . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 9, nemoxyga (246) 77. Cups usually deep, not proliferating from edges. Homosekakaic acid absent. . . . . . . . . . . . . . . . . . . . . . . 78 78. Podetia slender, trumpet-shaped; cups narrow; soredia covering entire podetium. Substance "H".absent. . . . . . . . . 9, fimbriata (243) 78. Podetia broad, goblet-shaped; cups wide; soredia usually absent on lower half of podetium where there is a continuous cortex. Substance WE” present. . . . . . . . . . . . . . . . . . . . . . 9, conista (244) 1. l. 1. 0 162 UMBILICARIA A Thallus pustulate; undersurface naked. Apothecia common; disks +'smooth, becoming somewhat gyrose with age with margins complete (leiodisc).. . . . . . . . . . . . . . . . . . . . . . 9, papulosa (267) Thallus smooth; undersurface rhizinate or lamellate. . . . . . . 2 2. Undersurface with flat, reticulate lamellae; rhizines absent; apothecia common; disks very gyrose with margins lacking (actinodisc) . .. . . . . . . . . . . . . 9, muhlenbergii(266) 2. Undersurface densely rhizinate with a mat of short black rhizines; apothecia not seen on L. 1. material, rare else- where. (Disks concentrically gyrose with a t complete proper margin Lgyrodisc] ). . . . . . . . . . . . . . . . . 9. mammulata (266) SARCOGYNE Apothecial disks rough, verrucose, carbonaceous; epithecium carbonaceous, thick, very uneven. Apothecia 0.3-1.0 mm across; hymenium (65-)lOO-120(-200) u (including the black epithecium). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . §, simplex (268) Apothecial disks 4 smooth, reddish-black (especially when wet); epithecium thin, brown, granular. (Note: Occasionally some carbonaceous material appears in epithecium, but always in very small amounts). . . . . . . . . . . . . . . . . . . . . . . . . 2 2. Apothecia 0.5-2.0 mm across; hymenium 85-120 u high; hypothecium usually yellowish or brownish. Common. . . . . . . 9, 219399 (267) 2. Apothecia less than 1 mm.across; hymenium 60-85 u high; hypothecium hyaline. Rare. . . . . . . . . . . . . 9, privigna (268) 163 PERTUSARIA 1. Fruit warts smooth, or at least not sorediate or granular . . . . 2 1. Fruit warts sorediate or granular. . . . . . . . . . . . . . . . . 8 2. Spores 8 per ascus (or rarely 4 per ascus). . . . . . . . . 3 2. Spores 2 per ascus or 1 per ascus. . . . . . . . . . . . . S 3. Spores uniseriate. . . . . . . . . . . . . . . . . . . . . . . . . 4 3. Spores biseriate. Fruit warts smooth; ostioles prominent, depressed. Fruit warts PD + orange and KOH + red (norstictic acid). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9, propingua (271) 4. Thallus epiphloedal, thick; fruit warts crowded, over 1mm across, eupertusariate, PD -, KOH - . . . . . . g, tuberculifera (274) 4. Thallus hypophloedal, thin; fruit warts scattered, under 1mm across, ampliariate, PD -, KOH - (?) . . . . . . . E, 912199 (269) 5. Apothecium lecanorine. Disk and thallus C + red . . . . . g, 321393 (275) S. Apothecium not lecanorine; fruit warts with one or more ostioles. Fruit warts and thallus C - . . . . . . . . . . . . . . . . . . . 6 6. Thallus grey, rarely yellowish; fruit warts eupertusariate, polycarpous . . . . . . . . . . . . . . . . . . . . . . . 7 6. Thallus yellowish-green, rarely grayish; fruit warts ampliariate (or infrequently, somewhat eupertusariate), monocarpous or dicarpous. Thallus UV + pink-orange; fruit warts PD 2 orange & KOH + yellow (stictic acid) . . . . . . . . . . g, xanthodes (276) 7. Fruit warts PD +-red, KOH + red (fumarprotocetraric acid + salacinic acid). Spores (85-)97-124(-l38) x 35-45 u, always hyaline, radial canals absent. . . . . . . . . . . . . . . . . . . . . g, subpertusa (272) 7. Fruit warts PD + orange & [OH + yellow (stictic acid). Spores 125-173 x 30-62 u, hyaline to brownish, radial canals and transverse wall 164 markings usually conspicuous. . . . . . . . . . . Melanaria macoun99' (277) 8. Sorediate warts KC + violet. Thallus dark ashy. . . .P. amara (27o) 8. Sorediate warts KC -. Thallus light or dark grey. . . . . .9 9. Soredia PD + orange & KOH + yellow (thamnolic acid); spores 2 per ascus. . . . . . . . . . . . . . . . . . . . . . . 9, trachythallina (274) 9. Soredia PD -, KOH -; spores l per ascus. . . . . 10. (9, multipuncta) (270) 10. Thallus thin, smooth; fruit warts scattered, bases 2 broad; spores 97-110 x 45-48 p. . . . . . . 9, cfr. multipuncta (#1) 10. Thallus thick, verrucose; fruit warts crowded, base constricted; spores 125-150 x (45-)55-7O u . . . . g, cfr. multipuncta (#2) LECANORA l. Thallus becoming distinctly lobed at the margins, or subfoliose. Saxicolous. (Subgenus PLACODIUM) . . . . . . . . . . . . . . . . 2 1. Thallus with margins not lobed or subfoliose . . . . . . . . . . . 3 2. Thallus closely adnate, crustose; apothecia greenish or brownish, 0.5-1.5 mm in diameter. On calcareous substrates..9, muralis (287) 2. Thallus ascending, subfoliose to peltate; apothecia yellowish or orange, up to 2.5 mm in diameter. On granite . . 9, 599135 (288) 3. Apothecia immersed in thallus (especially in young condition); disks black; spores (12-) 16-20 3 7-10 h. Saxicolous (Subgenus ASPICILIA). . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 3. Apothecia sessile (Subgenus EULECANORA). . . . . . . . . . . . . . 5 4. Thallus KOH -. Pycnoconidia (9-)lO-14 x l u . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9, caesiocinerea (279) 4. Thallus KOH + yellow (stictic) or KOH +-red (norstictic). Pycnoconidia (lO-)13-18 x l u. . . . . . . . . . . 9, cinerea (282) S. 165 Disks pitch black. Apothecia up to 2 mm across; epithecium tinted violet, especially in KOH; spores 12-16 x 7-9 p. . . . 9, atra (278) . Disks yellowish to brown or dark brown (never black) . . . . . . . 6 6o spores 6-8 H Wide 0 e e s s o e s e s s s s o e e s o s o s 7 6. Spores 2-6(-7) u wide. . . . . . . . . . . . . . . . . . . ll . Disks heavily pruinose, C + orange; apothecia lavender. Apothecial sections KOH + blood red (norstictic acid) . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9, caesiorubella subsp. lathamii (279) Disks epruinose or very light pruinose, C -; apothecia brown. Apothecial sections KOH + yellow (atranorine alone). . . . . . . . 8 8. Amphithecium containing large colorless crystals; epithecium inspersed with granules. . . . . . . . . . . . . . . . . . .9 8. Amphithecium without large colorless crystals; epithecium not inspersed with granules . . . . . . . . . . . . . . .. 10 Epithecium PD + red-orange with the production of small orange acicular crystals. Epithecial granules persistent in KOH; apothecial disk dark brown, epruinose, strongly convex; margin crenate, soon becoming thin and bead-like; spores 12-14 x 7-8 u. Very rare. . . 9,.cfr. insignis (286) Epithecium PD - or PD + yellow (with no crystals formed). Epithecial granules dissolve in KOH; apothecial disk yellow-brown to red-brown, often slightly pruinose, flat to convex; margin thick, smooth to crenate; spores 10-13 x 6-7 p. Very common. . . . . . 9, chlarotera (280) 10. Thallus very thick, verrucose and chinky, neither granular nor sorediate; apothecia up to 2 mm in diameter, often twisted and "urn-shaped"; disk reddish-brown; apothecial cortex thick, 45-50 a. On cedar stumps and old wood. . . . . . . . 9, 1323;9(287) 10. Thallus thinner, smooth to granular and sorediate; apothecia 0.5-1.0 mm, circular, closely adnate; disk deep mahogany-brown; 166 apothecial cortex 16-25 u thick. On bark. . . . .iLecanora sp. (290) ll. Spores 5-7 x 2-4 p. Apothecia minute, 0.2-0.4 mm in diameter. 0n old wood. . . . . . . . . . . . . . . . . . . . . . 9, subintricata (288) ll. Spores 8-16 x 3-7 p. . . . . . . . . . . . . . . . . . . . . . . 12 12. Saxicolous (on limestone and mortar). Thallus almost lacking; apothecia 0.25 - 0.50 mm in diameter; disks yellow-brown to olive-brown; margins white or ashy, usually persistent; spores 9-10 x 4-6 p . . . . . . . . . . . . . . . 9, disperse (285) 12. On bark, wood, or bone. . . . . . . . . . . . . . . . . . l3 l3. Disks yellow pruinose, lemon yellow when young gradually turning red-brown. Thallus well-developed, grey, very rough; spores ll-l4 x 4-5 p. . . . . . . . . . . . . . . . . . . . . . . . 9, cupressii (284) 13. Disks epruinose or lightly white pruinose, yellow to brown. . . .14 14. Apothecial margin cortex indistinct, not gelatinous; thallus granulose to sorediate, yellow-green; apothecia scattered or crowded; disks yellow to buff. . . . . . . . . . . . . l6 l4. Apothecial margin cortex distinct, gelatinous, thick; thallus essentially absent, or if present, not granular or sorediate; apothecia very crowded, abundant. . . . . . . . . . . . . 15 15. Apothecial sections KOH - (atranorine absent). Disks buff to very pale brown, lightly white pruinose; spores 10-13 x 3-5 (-7) u . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9, 993299 (285) 15. Apothecial sections KOH + yellow (atranorine present). Disks yellow-brown to dark brown, epruinose; spores 9-12 x 4-7 p . . . . . 9cfrZ§L1£ (289) 16. Apothecial margins persistent, becoming thin and disappearing in age, soon becoming granulose; spores 10-16 x 3-5 p. Atranorine absent. Frequent. . . . . . . . . . . . . . . . 9, conizaea (284) l. l. 1. 167 16. Apothecial margins absent in all but the youngest apothecia, smooth when present and hardly distinguishable from the disk; spores ll-l3 x 3-4 p. Thallus C -. Very rare . . . . . . . . . . . . . . . . . . . . . 9, syggicta var. syggictera (289) OCHROLECHIA Thallus (cortex and medulla) C - and KC -. Disks often somewhat pruinose; spores 45-68 x 21-36 g. Vsriolaric acid present in apothecial margin. Common; corticolous.. . . . . . . . . . . . . . . . 9, parella (290) Thallus (cortex or medulla) C‘+ red . . . . . . . . . . . . . . . . 2 2. Cortex of apothecial margin C + red; amphithecial medulla C -. Thallus thick, verrucose; spores 40-60 x 25-26 u. Algae present in a layer (sometimes not continuous) below hypothecium; apothecial cortex relatively thin. Very rare; corticolous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9, rosella (291) 2. Cortex of apothecial margin C -; amphithecial medulla C + red, stipe C -. Thallus thin, rimose to verrucose; spores 38-59 x 21-26 g. Corticolous. . . . . . . . . . . Ochrolechia sp.(293) HAEMATOMMA Thallus covered with sorediate verrucae towards the center, becoming smooth at the edges; thallus eventually becoming a granular sorediate crust, blue-grey to greenish-grey; sterile. Thallus PD + orange & KOH + yellow (thamnolic acid). . . . . . . . . . . . . . 9, sp. (295) Thallus coarsely verrucose or almost granular, but not sorediate, whitish-green to yellowish-green; apothecial disks red-brown, common; spores (35-) 45-62 x 5-8 p. Thallus PD«+ orange & KOH + yellow (thamnolic acid). . . . . . . . . . . . . . . . . . . . 9, ochrophaeum (294) 168 CANDELARIELLA 1. Octosporous; thallus appearing mostly black, or pale to dull yellow in small areas, granulose to verrucose or subsquamulose. On calcareous rock. . . . . . . . . . . . . . . . . . . . . 9, aurella (296) l. Polysporous (spores about 20 per ascus); thallus yolk- to»greenish- yellow, never darkening, granular-verrucose, with granules or subsquamulose verrucae becoming crowded into flattened or rounded patches. On granitic rocks. Often sterile. . . . . . . 9, vitellina (296) FARMELIOPSIS l. Thallus yellow-green, surface bursting into irregular lamdnal soralia which coalesce into a mass of granular soredia. Thallus PD -, KOH -, usnic and divaricatic acids present. . . . . . . . . . . 9, ambigua (298) l. Thallus grey-green or grey, esorediate. Thallus PD + orange and KOH + yellow (thamnolic acid); usnic and divaricatic acids absent. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 2. Thallus isidiate, sdnate; sterile. . . . . . . . 9, aleurites (298) 2. Thallus not isidiate, often ascending; rarely sterile . . . . . e e e e e e e e e e e e e e e e e e e e e e e e _P_e BlacorOdia (299) PARMELIA l. Thallus yellowish-green (usnic acid present) . . . . . . . . . . . 2 1. Thallus greyish, olive-green, or brownish, no trace of yellow (usnic acid absent) . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 2. Soredia in punctiform soralia, or tiny verrucae scattered over upper surface of thallus; lobes broad, 4-6 mm, or rarely less. Medulla PD + orange, KOH -, KC + red (protocetraric and caperatic acids present). Corticolous or saxicolous..9, caperata (302) 169 2. Soredia or tiny verrucae absent; lobes less than 4 mm broad. Medulla PD + yellow or orange, KOH + red. Saxicolous. . . . 3 3. Isidia present. Stictic and norsticitic acids present. . . . . . 4 3. Isidia absent. . . . . . . . . . . . . . . . . . . . . . . . . . .5 4. Lower surface of thallus black almost to edge. . .9, conspersa 4. Lower surface of thallus buff to brown throughout..9, plittii 5. Lower surface pale brown to buff; thallus more or less ascending. Salscinic acid present. . . . . . . . . . . . . . . . .9, stenophylla 5. Lower surface black except very close to margins. . . . . . . . . 6 6. Salscinic acid present. . . . . . . . . . . . . 9, tssmanica 6. Stictic and norstictic acids present. . . . . . 9, arseneana 7. Thallus olive-green (wet) or brown (dry), never grey. Irregular laminal soralia present. Medulla C + red . . . . . . . . . . . . . . . . . . . . . . . . . . 9, subaurifera 7. Thallus grey or grey-green. . . . . . . . . . . . . . . . . 8 8. Pseudocyphellae (white dots) scattered over upper surface. . . . .9 8. Pseudocyphellae absent. . . . . . . . . . . . . . . . . . . . . .12 9. Medulla C -; protolichesterinic acid present. Soredia or isidia absent, under surface black, becoming pale at margins. Very rare. . . . . . . . . . . . . 9, appalachensis 9. Medulla C + red, protolichesterinic acid absent. . . . . . 10 10. Isidia absent, soredia present. . . . . . . . . . . . . . . . . .11 10. Isidia present, soredia absent. Very common. .. . . . 9, rudecta ll. Soredia in punctiform soralia; lower surface pale brown. Frequent. . . . . . . . . . . . . . . . . . . . . 9, borreri ll. Soredia marginal; lower surface black. Very rare . . . . . . O O O O 0 O O O O O O O O O O O O O O O O O 0 £0 Olivetorum (302) (309) (312) (313) (301) (312) (30m (310 (300 (30» 12. Marginal cilia present. . . . . . . . . . . . . . . . . . . . . 13 12. Marginal cilia absent. . . . . . . . . . . . . . . . . . . . . . 17 13. Soredia absent. . . . . . . . . . . . . . . . . . . . . . 14 13. Soredia present. . . . . . . . . . . . . . . . . . . . . . 15 14. Medulla KC -, KOH + red, PD + yellow (norstictic acid present, stictic & protocetraric acids absent); cilia usually abundant; lower surface with a * broad irregular white margin; apothecia distinctly perforate. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9, perforata l4. medulla KC + red, KOH -, PD + orange (protocetraric acid present, norstictic & stictic acids absent); cilia very sparse; lower surface of thallus uniformly black, lightening to brown at margin; apothecia not perforate. . . . . . . . . . . . . . . . . . . . . 9, michauxiana 15. Rhizinaa present to the thallus edge (hypotrachynoid); upper surface of thallus t covered with reticulate.cracks and/or tiny, irregular white areas (maculae); soredia marginal or laminal. Medulla PD'+ orange, KOH + red (salacinic acid present) . . . . . . . . . . . . . . . . . . . . 9, reticulata 15. Rhizines absent from edge of thallus (amphigymnioid); reticulate cracks & maculae absent. . . . . . . . . . . . . . . . . . 16 16. Lower surface of thallus smooth, not rugulose, with a t broad, irregular white margin; soredia apical or marginal. Medulla KOH + red, PD + orange (stictic + norstictic acids). Common. .9, hypotropa(304) 16. Lower surface of thallus rugulose, uniformly black or lightening slightly to brown at margin; soredia submarginal. Medulla KOH + yellow, PD + orange (stictic acid present, norstictic acid absent). Rare. . . . . . . . . . . . . . . . . . . . . . . . . . . 9, perlata (308) 171 17. Thallus with conspicuous reticulate ridges and depressions, especially on younger portions of the thallus. Medulla KOH + blood red (salacinic acid). . . . . . . . . . . . . .18 17. Thallus t smooth, rugose or cracked, but without reticulate ridges and depressions. medulla KOH + or KOH -. . . . . . 19 18. Isidia present. . . . . . . . . . . . . . . . . . . 9, saxatilis (311) 18. Soredia present on ridges. . . . . . . . . . . . . . . 9, sulcata (313) 19. Soredia present. . . . . . . . . . . . . . . . . . . . . . 20 19. Soredia absent. . . . . . . . . . . . . . . . . . . . . . .22 20. Mbdulla pale yellow; soredia laminal. medulla PD - or PD + pale yellow, KOH + faintly yellow. . . . . . . . . . 9, aurulenta (301) 20. Medulla white; soredia marginal or laminal. . . . . . . . . . .21 21. Medulla PD + orange, KOH -, KC + red (protocetraric acid). Surface of thallus smooth with no maculae; lobes mostly 3-4 mm broad, crenate. . . . . . . . . . . . . . . . 9, robusta (310) 21. Medulla PD + orange, KOH + red (salacinic acid). Surface of thallus with reticulate cracks and maculae (see couplet #15). . . . . . . . . . . . . . . . . . . . . . . 9, reticulata (309) 22. Hedulla yellow, especially near the algal layer; chains of 2-4 moniliform cells scattered throughout the medulla. Medulla PD + orange, KOH + orange (unidentified substance). . . . . . . 9, galbina(303) 22. Medulla white thrdughout; moniliform cells absent. . . . . . . .23 23. Medulla PD + orange-red, KOH -, KC + red. Rhizines simple, unbranched; medulla thick, cottony (see couplet #14) . . . . . ..........................9.michauxisna (300 23. Medulla PD -, KOH + red-brown. Rhizines branched; medulla 1 thin, not cottony. . . . . . . . . . . . . . . . . . 9, livida ( 309 172 CETRARIA Terricolous. Thallus fruticose, dark brown, with broad or linear lobes ascending vertically producing a caespitose growth form; pseudocyphellae mostly marginal, linear. medulla PD - . . . . . . . . . . . . . . . . . . . . . . . . . 9, islandica subsp. 999929_ (318) Corticolous. Thallus foliose, brown, grey, or yellowish; lobes often ascending but never linear and never caespitose. . . . . . 2 2. Thallus grey, pitted; lower surface mostly white, sometimes mottled. Medulla I + blue. . . . . . . . . . . . 9, tuckermanii (320) 2. Thallus yellowish-green or brownish-green, never grey. . . .3 Lower surface yellow; upper surface greenish-yellow. . . .9, viridis (320) Lower surface brown; upper surface brown or greenish-brown. . . . 4 4. Apothecia originating on upper surface; thallus small, appressed; lobes narrow, finely divided, 0.5 - 0.75 mm broad, never ciliate. Very rare. . . . . . . . . . 9, fendleri (313) 4. Apothecia originating on lower surface; thallus larger, t’ascending; lobes 1.5 - 4 mm broad, often conspicuously ciliate. Canon in bogs. Medulla KC + red, UV + (in L. I. material) . . O O O O O O O O O O O O O 0 O O O 0 O O O O O O O O Q. Ciliaris (317) ALECTORIA Thallus caespitose, wiry; soralia with isidia. Common. .9, nidulifera (323) Thallus pendent, long; soralia without isidia. Very rare.. 9, glabra (322) RAMALINA Lacinae subterete or angular, t'papillste. Medulla KOH + red & PD + yellow (salacinic acid). Spores straight, ellipsoid, 11-13 x (4-) 5-6“000000000000000000eseooeo°°£‘Lfl.—LEL1(326) Lacinae strongly flattened. medulla KOH -, PD -. . . . . . . . . 2 173 2. Lacinae 3-8 mm broad, coarsely tuberculate-papillate. Very rare. . . . . . . . . . . . . . . . . . 9, cfr. complanata (324) 2. Lacinae 1-3 mm broad, not tuberculate. . . . . . . . . . . . 3 3. Spores fusiform, straight or slightly curved, 18-24 (-31) x 3-5 p; lacinae strap-shaped, with white striations (pseudocyphellae?) usually evident. . . . . . . . . . . . . . . . . . . . ..9, stenospora (325) 3. Spores ellipsoid, straight, 8-13 x 4-6 p; lacinae strap-shaped to broadened, often with numerous short proliferations along the margins; smooth, often with white punctiform pseudocyphellae, often sub- canaliculate. . . . . . . . . . . . . . . . . . . . . . . 9, fastigiata (324) USNEA l. medulla rusty-red. . . . . . . . . . . . . . . . . . . . . . . . .2 1. Medulla white. . . . . . . . . . . . . . . . . . . . . . . . . . .3 2. Thallus subpendent to pendent; branching irregular, often dichotomous, never strigose; isidiate-soralia present; apothecia rare. Norstictic, salacinic, etc., absent..9, mutabilis(327) 2. Thallus erect, shrubby, strigose; branchlets short; isidia and soredia absent; apothecia common. Norstictic acid present in about 501 of the specimens seen. . . . . . . . . . . . 9, strigosa(328) 3. Thallus pendent, filaments exceedingly slender, never tuberculate or papillate; stramineous or yellow-green. Medulla PD - . . . . . . 4 3. Thallus erect or subpendent; filaments generally coarse, papillae and/or tuberculae present; dark ashy-green, at least in older portions. Medulla PD + yellow or orange. . . . . . . . . . . . . 5 4. Branching by frequent dichotomies; perpendicular side branches infrequent; axis reddish-brown; articulations with swollen joints conspicuous; cortex intact. Common in bogs.. 9, trichodes (330) 174 4. Branching infrequently dichotomous; perpendicular side branches common and regularly spaced; axis white; articulations with swollen joints absent; cortex becoming farinose. Very rare. . . . . . . . . . . . . . . . . . . . . 9, longissima (327) 5. Medulla KOH - (or KOH i very faint yellow). Filaments papillate and tuberculate; branches coarse; erect or subpendent; cortex very thick and chondroid; isidiate-soralia usually present; base rarely blackened. Protocetraric acid, or rarely, barbatic or fumar- protocetraric5 acid present. . . . . . . . 9, subfusca sensu Motyka (330) S. Medulla KOH + deep yellow or red (often distinct only in the apothecial medulla). . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 l 6. Isidiate-soralia present; medulla very lax; apothecia not seen. KOH + red or yellow (salacinic or stictic acids present). Cape Cod region, fairly common. Not on L. I. . . . . . . . . . . . . . . . . . . . . . 9, cfr. gggggg_ 6. Isidiate-soralia absent; medulla compact; apothecia common. KOH + red (norstictic acid). Filaments strigose, scrobiculate on young branches. . . . . . . . 9, strigosa (328) CALOPLACA l. Corticolous. . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 l. Saxicolous. . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 2. Apothecial margin containing few or no algae; thallus yellow (KOH + red-purple), thin, sorediate. Spores 13-17 X 8-10 p; 181211“ 5-7 " longs s s e o e e e e e 90 (1186010! (335) 5 The two specimens containing fumarprotocetraric acid were from Cape Cod (Brodo 4161, 4338). 3. 5. 7. 175 2. Apothecial margin containing a distinct algal layer; thallus esorediate. . . . . . . . . . . . . . . . . . . . . . . . 3 Apothecial disk brown, KOH - (or vaguely pale violet), pruinose. Spores l3-l9 x 7-10 u; isthmi 3-6 u long. . . . . . . 9, camptidia (332) . Apothecial disks orange or yellow, KOH + dark purple or red-purple, not pruinose. . . . . . . . . . . . . . . . . . . . . . . . . . 4 4. Thallus pale yellow or cream-colored, KOH + red-violet (often weakly), thin, smooth. Apothecial disks yolk-yellow to yellow-orange; margins yellow; spores ll-l3 x 4-6 p; isthmi 3-4 u long. . . . . . . . . . . . . . . 9, aurantiaca (332) 4. Thallus grey-green, ashy, or dark bluish-grey, KOH - . . . 5 Amphithecium thick, ashy to blue-grey, entirely persistent; apothecial disks sordid yellow to yellow-orange; spores 12-16 x 7-8 a; isthmi (4-) 5-6 u long. . . . . . . . . . . . . . . . . . . . . 9, 999999 (333) Amphithecium very thin, pale-grey to ashy, soon disappearing and revealing an orange margin; apothecial disks dark orange to red- orange; spores 10-14 x 4-7 a; isthmi 2-4 (-5) u long. . .9, pygacea (336) 6. Spore isthmi less than 3.5 u long; thallus minutely areolate, yellow, becoming black or ashy, disappearing. Apothecia 0.25-0.40 mm in diameter; disks dark orange to orange-brown; margin yellow to orange, often becoming leprose or granular; spores 12-17 x 7-9 p. . . . . . . . . . . . . . 9, feracissima (335) 6. Spore isthmi more than 3.5 a long; thallus yellow, rarely darkening. . . . . . . . . . . . . . . . . . . . . . . . . 7 Thallus effuse granular or sorediate to subsquamulose or areolate. Apothecial disks orange; margin yellow, often sorediate; spores 9-13 x 5-7(-9) u; isthmi 3.5 - 5 u long; sometimes sterile. . . . .9, citrina (334) 176 7. Thallus smooth, rimose, areolate, squamulose, or disappearing. . . . . . . . . . . . . . . . . . . . . . . . . . 8 8. Thallus conspicuous, squamulose; apothecial disks dark red- orange to orange-brown; margins dark orange; spores 11-17 x 5-7 p. . . . . . . . . . . . . . . . . . . 9, flavovirescens (336) 8. Thallus essentially absent, or with rare yellow squamules; apothecial disks orange with yellow-orange margins; spores 11-15 x 4-6 p. . . . . . . . . . . . . . . . 9, aurantiaca (332) XANTHORIA l. Thallus with granular soredia in labriform soralia; lobes very small and narrow, 0.2 - 1.0 mm broad; apothecia rare. . . . . . 9, 999955 (337) l. Thallus esorediate; lobes broad, (2-)3-4 mm broad, flat; apothecia com“. 0 O O O 0 0 O O O 0 O O O 0 O O O O O O 0 £0 Parietina ( 337) TELOSCHISTES 1. Thallus very short, caespitose, yellowish to tan; lacinae flattened, t‘striate, giving rise to short irregularly shaped side branches ending in pointed cilia; soredia absent. . . . . . . 9, chrysophthalmus(339> l. Thallus longer, dark yellow-orange; lacinae terete or ridged and angular; cilia absent; patches of soredia frequent throughout leng ch 0 O O O O O O O O O 0 O O O O O O O O 0 O 0 O O I 1. flav1cana ( 340 BUELLIA 1. Saxicolous. . . . . . . . . . . . . . . . . . . . . . . . . . . .2 l. Corticolous or lignicolous. . . . . . . . . . . . . . . . . . . .3 3. 3. 5. S. l. l. 177 2. Medulla KOH + red (norstictic acid). Thallus whitish-grey, areolate; prothallus black, well developed. Apothecia mostly sessile or immersed between areoles. . . . . . . 9, stigmaea (342) 2. Medulla KOH - . Thallus dark ashy-brown; verrucose; prothallus inconspicuous. . . . . . . . . . . 9, turgescens (344) Apothecial sections KOH + red (norstictic acid). Apothecia 0.5 - 1.5 mm in diameter. . . . . . . . . . . . . . . . . . . . . . . . 4 Apothecial sections KOH - . Apothecia less than 0.5 mm in diameter. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 4. Exciple pallid within; grey stipe absent; spores 17-24 x 6-8 p; hymenium 80-130 p high, hyaline. . . . .9, curtisii (340) 4. Exciple uniformly dark; grey T-shaped stipe present; spores 11-17 x 6-8 a; hymenium 55-75 u, yellowish . . . . . . . . . . . . . . . . . . . . . . . . . . . . .‘9. stillingiana (343) Spores 8 per ascus. . . . . . . . . . . . . . . . . . . . . . . . 6 Spores 12-16 per ascus. Exciple pale within. . . . . . 9, polyspora (341) 6. Thallus PD + red (fumarprotocetraric acid). Spores 19-23 x 8-9 u; apothecial margin usually absent; disk hemispherical. . . . . . . . . . . . . . . . . . 9, dialyta (341) 6. Thallus PD -. Spores 9-11 x 6-7 p; apothecial margin distinct, disappearing with age; disk flat to slightly convex......................_B_.punctata (342) RINODINA sax1c°1°ua O O O O O O O O O O O O O O O O O O O O O O 0 O O O O 0 2 Corticolous. . . . . . . . . . . . . . . . . . . . . . . . . . . 4 2. Thallus pale grey or brownish-grey. . . . . . . . . . . 3 3. 178 2. Thallus yellowish-green. Thallus lobed at margins; spores 10-12 x 6-7 p. Medulla C + red, PD -, KOH - . . . 9, oreina 0n siliceous rock. Thallus verruculose to almost squamulose; spores 17-23 x 9-13 u. . . . . . . . . . . . . . . . . . . . .9, confragosa On concrete. Thallus areolate to minute verrucose; spores 10-16 x 6-8 p. . . . . . . . . . . . . . . . . . . . . . . . . . 9, 999999 4. Spores S-7(-8) x (8-)10-12(-15) u; hypothecium dark brown. Apothecia less than 0.5 mm in diameter. . . . 9, milliaria 4. Spores over 15 u long; hypothecium hyaline or yellowish. .5 Thallus brownish-green to olive, verrucose or granulose to smooth and t squamulose; spores pachysporous (examined in water), 16-23 x 6-10 u. . . . . . . . . . . . . . . . . . . . . . . 9, pachysperma Thallus thin, smooth, light grey-green; spores mostly pachysporous (examined in water), 17-24 x 9-11 p. . . . . . . . . 9, spplanata PHYSCIA Thallus deep green, olive-green, or brownish-grey; upper cortex KOH - (atranorine absent). . . . . . . . . . . . . . . . . . . . . . . . 2 Thallus grey or grey-green; upper cortex KOH + yellow (atranorine present). . . . . . . . . . . . . . . . . . . . . . . . . . . . . .3 2. Thallus esorediate; lobes finely divided, becoming covered with small lobules; lower surface white to buff; rhizines tan to brown; medulla white. Very rare. . . . . . . . . . eeseeeseeeeeeeeeeAMECIChiaealmlata 2. Thallus with greenish marginal or laminal soralia; lobes crenate to entire, never subsquamulose; lower surface black; rhizines black with white tips; medulla white (KOH -) or red- orange (KOH + purple). Common. . . . . Physcia orbicularis (346) (345) (347) (345) (346) (344) (354) (350) 179 Medulla mustard-yellow. Thallus with marginal granular soredia; lobes pruinose, 1-2 mm broad. . . . . . . . . . . . . . . Pyxine sorediata (348) O kdUIIa White. 0 O O O O O O O O O O O O O O O O O O O O O O O O 4 4. Thallus sorediate or with granules resembling soredia. . . . 5 4. Thallus esorediate. Apothecial disks very dark brown to black, somewhat pruinose. . . . . . . . . . . . . . . . . . 10 Soredia in laminal soralia. . . . . . . . . . . . Physcia tribacoides (353) Soredia (or granules) marginal or terminal. . . . . . . . . . . . .6 6. Lobes helmet-shaped, bursting into soredia. Lobes with long, white, marginal cilia. . . . . . . . . . . . . 99, adscendens (348) 6. Lobes t flat, not helmet-shaped. . . . . . . . . . . . . . . 7 Lobes broad, (2-)3-4 mm, rounded; cortical hyphae parallel to surface. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 Lobes narrow, 0.3-2 mm broad; cortical hyphae at right angles to surface. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 8. Lower surface white (KOH -), t corticate . . . . . . . . . . . . . . . . . . . . . . . . . . . Anaptychia pseudospeciosa (355) 8. Lower surface yellow (KGB + purple), ecorticate . . . . . . . . . . . . . . . . . . . . . . . . . . Anaptychia obscurata (354) Lobes 0.3 - l.0(-l.5) mm broad; spores 16-19 x 6-9 p; soredia (or granules) large, marginal, sometimes reducing thallus to a granular crust. Corticolous, or very rarely, saxicolous. .Physcia millegrana (349) Lobes very narrow, 0.1 - 0.5 mm broad; spores 12-16 x 6-8 p; soredia (or granules), marginal and apical, occasionally laminal. Saxicolous. . . . . . . . . . . . . . . . . . . . . . . 99, subtilis (353) 10. Medulla KOH + yellow. White spots (maculae) present . . . . . I O O O O O C O O O O O O O O O O O O O O 0 O O P—ll-O 8120118 (349) 10. Medulla KOH -. White spots absent. . . . . . 99, stellaris (352) i 180 ANAPTYCHIA l. Thallus esorediate, brownish; upper cortex KOH - (atranorine absent); lobes finely divided, becoming covered with small lobules . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9, palmulata (354) 1. Thallus sorediate, grayish; upper cortex KOH + yellow (atranorine present); lobes not finely divided. . . . . . . . . . . . . . . 2 2. Lower surface light to deep yellow, KOH + red-violet, not corticate. . . . . . . . . . . . . . . . . . . . 9, obscurata (354) 2. Lower surface white, KOH -, t corticate. . . 9, pseudospeciosa (355) LEPRARIA l. Saxicolous. Thallus grey to dark ashy-green; granules large, often forming a t lobed, zonata thallus. Thallus PD + red, & KOH - (fumarprotocetraric acid), or rarely, PD + yellow (barbatolic acid) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9, Eggggg_(356) 1. Corticolous or lignicolous. . . . . . . . . . . . . . . . . . . .2 2. Thallus with a distinct bluish-grey cast, a 1 thin layer of dispersed granules with little or no prothallus. Thallus KOH + yellow & PD - (atranorine) or rarely, KOH + yellow & PD + red (fumarprotocetraric acid + atranorine) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 99 992999, (355) 2. Thallus pale green or sometimes yellowish-green, thick masses of granules subtended by a thick, white, prothalline mat. Thallus KOH + yellow & PD + orange (atranorine + stictic neid)e s e e e s e s e e e s e e o e e e s e e s e e 0 CL. 81). (357) ARTHOPYRENIACEAE ARTHOPYRENIA Arthqpyrenia cerasi (Schrad.) Mass. Ricerch. Auton. Lich. 167. 1852. Verrucaria cerasi Schrad. Ann. d. Bot. 22:86. 1797. Material seen - SUFFOLK COUNTY: 9£ggg_9919 (123). Pink (1935) reports 9, cerasi from young oaks, but the Long Island material was on Myrica pennsylvanica. Distribution - Maryland, Iowa, California (Fink, 1935); Europe. Arthopvrenig_pinicolg_(Hepp) Mass. Symm. Lich. 118. 1855. Pyrenula punctiformis var. cineropruinose f. pinicola Hepp, Flecht, Europ. 106. 1853. Material seen - SUFFOLK COUNTY: Brodo 3176 (65). Degelius (1941) discusses the synonomy and gives a detailed description of his specimens. The Long Island material fits his description very closely. The specimen cited above was collected on the base of a white Oak (Quercus alba). Distribution - Tennessee; Europe. [9rthopyreni9 sublitoralis (Leight.) Arn. was not collected, but may well be found on shells and barnacles if looked forJ LEPTORHAPHIS leptorhaphis epidermidis (Ach.) T. Fr. Nova Acta Reg. Soc. Sci. Upsal. III. 3: 373. 1861. (: Lich. Arct. 273. 1860.) ‘999999_epidergigis Act. Lich. Suec. Prodr. 16. 1798. Material seen - SUFFOLK COUNTY: 99292_999§ (27), 9999 (78), 9999 (65), 1225. (91). 2m (22). 2.5.9.; (97). 21.7.; (31). 3.19.9. (122). 2:317. (66). All specimens of this species were found on the bark of 999999_populifolia. It is superficially similar to Polyblgstiopsis quercicolg but their spores 181 IIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIIiIIIIIIIHIIIIIIIIIIIIIIIIIIII' 182 are substrates and quite different. Distribution - Eastern United States (Fink, 1935); Connecticut, Michigan, Wisconsin, Arizona, Black Hills: Temperate element, East Temperate subelement; Europe. POLYBLASTIOPSIS Polyblastiopsis quercicola sp. nov. Material seen - SUFFOLK COUNTY: Brodo 2651 (61), 2674 (108), 2788 (31). Thallus subtilissimus, hypophloedalis, albus; algae non visa. Pseudothecia nigra, diam. 0.15-0.25 mm, hemisphaerica, dispersa, superficialia, sed saepe ex parte thallo vel epidermide corticis tecta; ostioles conspicues saepe; parietes carbonisati, virides-nigres ut oblinentur, praecipuus in KOH. Filamentae paraphysoideae et asci immersi in substrate gelatinoso, dissoluto in KOH. Filamentae paraphysoideae persistentes, distinctae in KOH, ramosa et anastomosa copiose, 1.5-2.5 u diam. Asci parietes t subtiles in H20, sed perspicue crassi in KOH, praecipuus spice. Sporae octonae, irregulariter seriatae, murales vel submurales, septis transversis 3-6, septis longitudinslibus 1-2, hyalinibus; vagina episporis lucida, conspicua, levis in H20, T irregularis in KOH; 16-26 X 7-10 p. Pycnidia nigra, minutissima, disperse. Pycnoconidia hyalina, non septate, elongata-cylindrica ad fusiforma, 7-9 X l u. Ad corticem Quercus alba. Holotype: New York. Suffolk County: Shoreham. Saint Joseph's Villa, N. Country Road, black oak woods, Brodo 2651, 7 July 1961, on Quercus alba, 0 ft. and higher (MSG) (see figures 102, 104, 105d). Thallus very thin, hypophloedal, appearing white; no algae evident. Pseudothecia black, 0.15 - 0. 25 mm in diameter, hemispherical, scattered, superficial, but often partially covered by thallus and/or upper layers of bark; ostioles often conspicuous; walls carbonaceous, greenish-black when smeared, especially in KOH. Paraphysoid threads and asci embedded in gelatinous material which dissolves in KOH; paraphysoid threads persistent, distinct in KOH, abundantly branched and anastomosing, 1.5 - 2.5 u in diameter. Asci appearing 1 thin walled when mounted in water, but clearly thick walled, especially at apex, when mounted in KOH. Spores eight per — 183 ascus, irregularly arranged, muriform or submuriform, 3~6 transverse septa, 1~2 longitudinal septa, hyaline; hyaline epispore sheath conspicuous, smooth in a water mount, f irregular in a KOH mount; 16~Z7 x 7~10 u. Pycnidia black, extremely minute, scattered. Pycnoconidia hyaline, nonseptate, elongate-cylindrical to fusiform, 7~9 x l u. The specimens from Long Island as well as those from New Jersey (99999 3728, 3755) were rather uniform in morphology. The thallus often covers several square centimeters or even decimaters on or near the bases of oaks. Pseudothecia vary little in size. Spores are l7~27 x 7~10 p and always show the gelatinous epispore sheath. Pycnoconidia are 6~9 x l.0~l.5 p, hyaline and nonseptate. Algae (apparently Trentepohlia) were in very small amounts just below the pseudothecia of a few specimens. The species differs from similar 9, fallaciosa (Stizenb.) Zahlbr. in having larger spores and hyaline, nonseptate rather than brown, septate pycnoconidia, and from 9, lactea in having somewhat smaller spores and eight rather than four spores per ascus. Polyblastiopsis fallax (Nyl.) Fink which is close to 9, quercicola from the description in Fink (1935) appears to be a synonym of Arthopyrenia fallax. The species was found on the bark of Quercus alba and Q, stellata. Similar species are almost always found on 999999 or some other tree, and so the unusual substrate served as the source of the specific epithet. ARTHONIACEAE ARTHONIA Arthonia caesia (Flot.) Kdrb. Parerg. Lich. 269. 1861. Coniangium caesium Flot. in K5rb. Syst. Lich. Germ. 295. 1855. Material seen - SUFFOLK COUNTY: 19 specimens collected by Imshaug and/or Brodo. Fink (1935) probably included this species in his concept of 9, impolite (Ehrh.) Borr. (Syn. 9, pruinose Ach.). The two species are similar in their 184 leprose thalli and small, dark, heavily bluemgrey pruinose ascocarps, but may be separated as follows: A. impolita6 A. caesia, Phycobiont Trentepohlia Trebouxia Fruit base hyaline yellow to redwbrown Thallus reactions KOH + yellow; KC + red KOH a ; KC - The Long Island material was mostly fertile and agreed in all respects with European descriptions of the species. The species is found on bark of all kinds in shaded or exposed woods, or on exposed downs. Under the right conditions, it apparently has a very rapid growth rate and was seen almost covering young twigs and branches in an oak forest in Laurel. Distribution - Tennessee, Wisconsin, but probably common in eastern United States; Europe. Arthonia mediella Nyl. Not. Soc. Faun. Fl. Fenn. Forhandl. l: 238. 1858 - 1859. Material seen - SUFFOLK COUNTY: §59g9_1225,(9os). The Long Island material agreed well with the description by Redinger (1937 - 1938), and with a specimen from Finland (£533.;g1, hb. MICK). Redinger states that the species is corticolous, but the Long Island specimen was an old wood on an exposed bluff overlooking Long Island sound. Distribution - First report from North America; Europe. Arthonia punctiformis Ach. Kgl. Vet. Akad. Nya Handl. 130. 1808. Material seen - SUFFOLK COUNTY: Imshaug 92139 (132). Orient, 955992 199, 6 Based on Michigan material (MSG). % 185 5 April 1914, (Latham); Montauk, Latham 3953, 6 April 1927 (Latham); Greenport, Latham 8608, 30 April 1939 (Latham); Orient, Latham 24178, 21 March 1915 (Latham). This species is similar to_A, radiata (Pers.) Ach. in many respects, but the latter has larger ascocarps and an epiphloedal thallus whereas the ascocarps of A, punctiformis are 0.1 - 0.2 mm across and its thallus is hypophloedal. Arthonia punctiformis is found on bark of various kinds, and is rare on Long Island. Distribution - Maine, Connecticut, Tennessee, Minnesota, Alaska; throughout the United States (Fink, 1935); Europe; Asia (Vainio, 1928). Arthonia sexloculares Zahlbr. Ann. Myc. 12: 336. 1914. Material seen - SUFFOLK COUNTY: EEQQQ'9§9§.(115). This single specimen was found growing on the deeply shaded base of Celtis sp. The distinctive spores agree perfectly with the description of Redinger (1937-1938) based on European material. Distribution - First North American record; Europe. Arthonia siderea Degel. Ark. Bot. 30A(1): 14. 1940. Material seen - SUFFOLK COUNTY: 99g§g_11§9 (101), 1999 (101), 1919 (1003), 1%; (117). 18.7.9. (117). 229$ (93). 2312 (113). gags (84). M (71). _2_§.§_6. (110), 9191 (111), 9931 (119); Orient, Latham 8598A,10 April 1939 (Latham); Orient, LEEEQEL§QQQ, 30 April 1939 (Latham). Although Degelius described this species are having "black apothecia," the hysterothecia actually range in color from red-brown to black. When moistened, the hysterothecia always appear a deep mahogany and never are black. The holotype, which Dr. Degelius kindly sent me, had hysterothecia :_ 186 which, although almost black when dry, also show the red~brown tint when wet. The Long Island material agrees in every respect with the holotype. This species is probably more common than would be thought judging from the number of times it has been reported. In Fink (1935), 9, siderea easily keys out to A, gregaria (Weig.) Kfirb. which is now recognized as a synonym of A, cinnabarina (DC.) Wallr. (see discussion in Redinger 1937-1938). Arthonia cinnabarina, however, has rusty red, often powdery hysterothecia, which are KOH + red~violet. The hysterothecia of 9, siderea are KOH -, smooth and often shiny. The spore type and ascocarp color clearly distinguish it from A, radiata (Pers.) Adl. Arthonia siderea is found on the bark of black oaks well above the base, and is probably photophilous. Distribution - Maine; endemic. Arthonia sp. Material seen - SUFFOLK COUNTY: 9gggg_1g1g (98). This species is similar to A, siderea in the color and stellate arrange- ment of the hysterothecia. It is, however, readily distinguished by its pruinose hysterothecia and pale brown spores with equal sized cells. It was found on the bark of Carya glabra. ARTHOTHELIUM Arthothelingtaediosum (Nyl.) Mflll. Arg. Flora 63: 287. 1880. Arthonia taediosa Nyl. Ann. Sci. Nat. Bot. IV. 3: 171. 1855. Material seen - SUFFOLK COUNTY: 9£gdg_9§99 (66); Greenport, ngggm;§§193 16 April 1938 (Latham); Greenport, 9g£9gg_19gg, 26 January 1933 (MICE). Two of the specimens from Long Island differ somewhat in spore size (32-37 x 10-17 u in Brodo 3832; 34-46(-55) x 14-23(-27) u in Latham 8610) —' 187 but the thallus and ascocarp characters agree perfectly. Except for the large spore sizes in the Latham specimen, the specimens fit the description of A, taediosum given by Redinger (l937~l938) very well. Redinger's spore measurements are 24-30(-33) x 8-13 u. The only other possible Arthothelium which could be considered for the Latham specimen would be A. distendens (Nyl.) M311. Arg. and Nearing annotated the specimen with that name. After having examined a number of specimens of both species in the Fink herbarium (MICH), I feel there is little doubt that both Long Island specimens belong to A, taediosum. I could find no spores in Latham 7206 but the specimen was identified by Josiah Lowe as this species. The specimens of A, distendens had thicker thalli and much broader hysterothecia (up to 1 mm across). In the Long Island material, the ascocarps were punctiform to irregular, and 0.1 - 0.2 mm across. Arthothelium taediosum was found on the smooth bark of red maple and oak on Long Island. Distribution - Eastern United States, California (Fink, 1935); Connecticut; Europe (Redinger, 1937-1938). MICAREA Micarea melaena (Nyl.) Hedl. Bih. Kgl. Svensk. Vet. Akad. Handl. afd. III, no. 3, 18: 82. 1892. Lecideg_melaena Nyl. Bot. Not. 182. 1853. Material seen - NASSAU COUNTY: 99992.1219_(14). This species is considered in the genus Bilimbia in Fink (1935). The Long Island specimen was found on a rotting stump. Distribution - Eastern United States (Fink, 1935); Connecticut, Michigan: Temperate element, East Temperate subelement; Europe; Asia (Vainio, 1928). Micarea prasina (Fr.) KBrb. Syst. Lich. Germ. 399. 1855. Biatora prasina Fr. Stirp. Agri Fems. 38. 1826. , . 188 var. sordidescens (Nyl.) comb. nov. Lecidea sordidescens Nyl. Flora 57: 312.'1874. Material seen - NASSAU COUNTY: Brodo 1497 (9), 3497 (4); SUFFOLK COUNTY: Brodo 1989 (51), 2578 (73). The Long Island material would fall into the var. sordidescens (Nyl.) Lettau emend. Erichs. of Catillaria prasina (Erichsen, 1957). This variety has a KOH + violet "epithecial" reaction. The spores are either mostly nonseptate or mostly uniseptate, but both types can always be found in a smear of the ascocarp. The species is treated in the genus Catillaria in Fink (1935). Micarea prasina is found on rotten wood but is rare on Long Island. Distribution - Northern and eastern United States (Fink, 1935); Connecticut, Minnesota, Black Hills: Temperate element, North Temperate subelement; Europe; Asia (Vainio, 1928). OPEGRAPHACEAE _OPEGRAPHA Qpegrapha cinerea Chev. Journ. Phys. Chim. Hist. Nat. 94: 41. 1822. Material seen - SUFFOLK COUNTY: Imshaug 25674 (72), Brodo 59-221 (72), 19; (90A), 121 (908), 1919 (98), 1199 (127), 9999 (61); Greenport, (collector unknown), April 1903 (FR). This species is somewhat similar to Q, vulgata which is distinguished by having 1) unbranched, shiny hysterothecia, 2) a greenish-brown thallus, 3) spores 15-20 u long, and 4) a specificity for coniferous bark (Redinger, 1937-1938). Opegrgpha cinerea is found on the bark of smooth-barked broadleaf trees such as 93515 sp. and Quercus sp. (which agrees with the habitat notes of Redinger, 1937~1938). All the Long Island specimens were found within a half mile of the north shore. —_ 189 Distribution ~ Florida (Fink, 1935), the Smoky Mountains of Tennessee; Europe. Opegrapha rufescens Pers. Neue Ann. Bot. 1:29. 1794. Material seen - SUFFOLK COUNTY: Orient Point, Latham, 5 June 1911 (NYS). Distribution - Florida (Fink, 1935); Europe. CALICIACEAE CHAENOTHECA Chaenotheca phaeocephala (Turn.) T. Fr. Nova Acta Reg. Soc. Sci. Upsal. III, 3: 351. 1861. (s Lich. Arct. 251. 1860). Lichen phaeocephalus Turn. Trans. Linn. Soc. Lond. 8: 260. 1807. Material seen a SUFFOLK COUNTY: Imshaug 25810 (86), 99292,919A (102). The spores of this species, normally brown, often become colorless in KOH. Chaenotheca phaeocephala is rare on Long Island and is restricted to rotting stumps of Chamaecyparis thyoides in shaded bogs. It was collected once in southern New Jersey (99g99_9119) and once on Cape Cod (99292_A991) in similar habitats and on the same substrate. Distribution - New England and Minnesota (Fink, 1935); Michigan: Temperate element, North Temperate subelement (?); Europe. VERRUCARIACEAE VERRUCARIA Verrucaria microspora Nyl. Ann. Sc. Nat. Bot. IV. 3: 175. 1855. Verruggria subsuperficialis Fink in Hedr. Mycologia 25: 304. 1933. Material seen - SUFFOLK COUNTY: Orient, Latham, 1925 (Holotype of X, subsuperficialis) (MICE); Orient, Latham, 1927, (HIGH, FH). 190 The holotype of 9, subsuperficialis was compared with a specimen of 1, microspora from Denmark kindly sent to me by Dr. Degelius. The two specimens were identical in morphology and ecology, both having thin, membranous, dark brown thalli, and small spores, and both having been found in the hydrohaline stratum (see page 74 ) on quartz pebbles. Distribution o Maine: Temperate element, Maritime subelement; maritime Europe (Santesson, 1939; des Abbayes, 1934). Verrucaria muralis Ach. Math. Lich. 115. 1803. Material seen - SUFFOLK COUNTY: 9gg§gi9§99 (115). This species was found growing alongside 9, nigrescens on mortar and brick in the aerohaline stratum at Orient Point. It differs from the latter species in having a white or ashy thallus and larger spores as well as an entirely different type of perithecium. Its involucrellum is black, hemispherical, and is almost entirely external to the thallus; the exciple appears hyaline. The species fits the description in Zschacke (1933) fairly well. Verrucaria muralis is considered a synonym of 9, ruoestris Schrad. in Pink (1935), but the latter is considered quite different by Zschacke, with an endolithic, sometimes disappearing thallus. Zschacke states that 1, muralis is found on sandstone and bricks. Distribution - Arctic-boreal element (?); circumboreal. Verrucaria nigrescens Pers. Ann. d. Bot. 15: 36. 1795. Material seen - SUFFOLK COUNTY: 99ggg_9§91 (115). The Long Island specimen together with one collected on Nantucket Island (Massachusetts) (Brodo 3964 B) agrees in most respects with descriptions by Zschacke (1933) and Fink (1935). However, the black medullary layer mentioned by Zschacke and others was not seen in all parts of the thalli although it was conspicuous in the Nantucket specimen. Servit (1954) describes the — 191 spores of this species as 20~28 x 11 u, but these measurements disagree with those of all previous authors. Both the Long Island and Nantucket material were found on concrete, and both were either in or close to the aerohaline stratum (see page 71). It is a common lichen on calcareous rocks in Europe. Distribution - Connecticut, Indiana, Black Hills, Washington, Manitoba: Temperate element, North Temperate subelement (see Fink, 1935); Europe; Asia (Zahlbruckner, 1930). Verrucaria silicicola Fink in Hedr. Mycologia 25: 305. 1933. Material seen - SUFFOLK COUNTY: Brodo 2710 (111), 2826 (115); Three Mile Harbor, Latham 32177, 16 April 1951 (Latham); Orient, Latham 36780, 14 April 1950 (Latham); Three Mile Harbor, East Hampton, Latham 36781, 19 April 1949 (Latham); Shelter Island, Latham 36785, 1 June 1944 (Latham); Sag Harbor, Latham 36786, 2 June 1946 (Latham); Orient, 99£9§m_(Holotype) (MICH); East Hampton, Latham 2647, 20 April 1926 (MICH); East Hampton, Latham 3995, 10 April 1927 (MICH); East Hampton, Latham 32177 (?? see above), 11 April 1953 (M0). This species is similar in general external appearance to 9, microspora but the latter has much smaller perithecia and spores. Both species are found on pebbles and small stones in the hydrohaline stratum in the maritime region. Distribution - Long Island (Fink, 1935): Temperate element, Maritime subelement; endemic. DERMATOCARPON Dermatocarpon miniatum (L.) Mann, Lich. Bohm. Obs. Dispos. 66. 1825. Lichen miniatus L. Sp. P1. 1149. 1753. Material seen - SUFFOLK COUNTY: Montauk, Latham 22242, 6 May 1926 (Latham). 192 The specimen was found on a rock along railroad tracks. Distribution - Massachusetts, Connecticut, Tennessee, Alabama, Oklahoma, Michigan, Ontario, Minnesota, Black Hills, Washington, British Columbia; arctic to temperate (Ahti, 1964): Temperate element, North Temperate subelement, but arctic in Asia (Lynge, 1928), Europe (Lynge, 1938b), and Iceland (Lynge, 1940 c). PYRENULACEAE PYRENULA Eyrenula nitida (Weig.) Ach. Syn. Lich. 125. 1814. Sphaeria nitida Weig. Obs. Bot. 45. 1772. Material seen - SUFFOLK COUNTY: 9mshaug 25552 (52), Brodo 59~250 (67), 953 (47), 21s (63), 1221 (100a), 1657 (88), 1787 (127), 2210 (61), 133:3 (a3), 2539 (73), 2610 (84), 3232 (35), 3320 (129); Napeague, Latham 2835, 1 March 1927 (Latham); Greenport, Latham 3989, 1 April 1927 (Latham). Pyrenula nitida is the indicator species of a well known and well studied 99399 community in Europe (the Pyrenuletum nitidae Hill.). In both Europe and Long Island, the species is characteristic of smoothwbarked trees, chiefly 99399 (and Quercus on Long Island), in moderately shaded woods (Barkman, 1958; Almborn, 1948). Its position in a 9Aggg community continuum seems to be governed by light availability (Almborn, 1948). Distribution - Nova Scotia, Maine, Connecticut, Wisconsin, Minnesota; throughout the United States (Fink, 1935); Europe; Asia (Zahlbruckner, 1930). 193 MELANOTHECA ; . 11'. q . ,1 . Melanotheca cruenta (Mont.) Mail. Arg. Bot. Jahrb. 6: 397. 1885. Trypetheligm cruentum Mont. Ann. Sci. Nat. II. 8: 53?. 183?. Material seen w SUFFOLK COUNTY: Gardiner's Island, gagggm, 23 Jay 1923 (Latham). A description and discussion of this spec: s can be found in Johnson (1959). The Long Island specimen extends the known range of g. gguenta slightly northward. This range extension is known for several other coastal plain species, among them Cladonia santensis and Cladonia evansii. The specimen was found on a tree trunk in rich woods. Distribution w Along the coastal plain, New Jersey to Texas (Fink,l935): Temperate element, Coastal Plain subelement; endemic. TRYPETHELIUM Trypethelium virens Tuck. in W. Darl. Fl. Cestr. ed. 3, 453. 1853. Material seen - SUFFOLK COUNTY; Imshaug 25735 (132), 25743 (132), 25746 (132), Brodo 59‘194 (33), 2702 (111), 3070 (128), 3291 (33), 3211 (33), 3254 (119); Mbntauk, Point Woods, Latham 3992, 7 April 192? (Iatham); Orient, Latham Bros. woods, Latham 8598 C, 10 April 1939 (Latham); Napeague, Lathég 28356 A, 9 February 1949 (M0). The unusual specificity of this species for Ilex spp. and Fages grandifo11§_ was discussed on page 34 . One specimen (Latham 85983) was collected from a black oak. Johnson (1959) lists a number of other substrates as well. Trypethelium virens is apparently skiophilous (or hygrophilous?); in holly groves, thickets, and beech forests, it is never found on well illuminated trunks. 194 Distribution - Trypethelium virens shows an unusual North American distri~ bution due to its dual substrate specificity. It has a typical coastal plain distribution from Louisiana through Florida to New England (Fink, 1935) following the range of llex opaca, as well as an Appalachianvcreat Lakes distribution following the range of Fagus grandifolia. Temperate element, Eastern Temperate subelement; endemic. FORINACEAE PORINA Porina cestrensis (Tuck. in W. Dari.) Mull. Arg. Flora 66: 338. 1883. Verrucaria cestrensis Tuck. in W. Darl. Fl. Cestr. ed. 3. 452. 1853. Material seen - $"FFOLK COUNTY: Orient Point, lathamii, 22 March 1910 (NYS). The type material of this species (Michener 204, sub Verrucaria cestrica) was examined in the Farlow herbarium. A diagnosis of the holotype is presented in tabular form in the discussion of E3 hibernica which follows. Distribution ~ New England to Georgia, Alabama, and Tennessee (Fink, l935): Temperate element, Coastal Plain subelement; endemic. Porina hibernica James & Swine. in Swine. Lichenol. 2: 35. 1962. Material seen - SUFFOLK COUNTY: Brodo 1783 B (127), 2598 (84), gggg (33). This species bears certain resemblances to two other Poriggg from the New England area, g. cestrensis and g. rhaphidosperma M511. Arg. The table presented below points out some of the differences between them. The diagnoses of g, cestrensis and g, rhaphidosperma are based on the type specimens. The values given in parentheses under g. cestrensis are measurements of other specimens which were studied. P. cestrensis 195 P° rhaphidesperma P° hibernica Thallus well developed; ashywwhite to dirty greenish to greenishwblack greenagrey; smoeth olivaceous; very t9 cracked, well thin, smooth or developed scurfy tn almost absent. Perithecium 0.15 - 0. 25 mm 0°20 0 0°35 mm 0020 - 0.35 mm Exciple hyaline carbonaceous carbonacenus Spores: 34‘46 x 3~5 u size (30w42 x 5~6 n) (63~§l®3a120 x 2&5 p 58~65 x 5&7 p shape clavate, straight acicular, .lexuous t elongatesclavate t0 f acicular, straight septa 5~8 (3~7 [~9j) (941445 (swig-13 (~i6) cell size irregular equal frequently irregular Porina hibernica was always associated with E, nucula on oaks in well shaded moist woods. Distribution w Ireland (Kilarney: type locality); oceanic (?). Porina nucula Ach. Syn. Meth. Lich. 1129 1814. Material seen - SUFFOLK COUNTY: Brodo 1783 A (127)p g§2§,sterile (84), 3:11 (33). This species is one of the few Porinae having a pale, noncarbonaceous involucrellum. The Long Island material was somewhat aberrant in that the perithecia were small and did not have the "plaques" 0r lamellae described by Swinscow (1962). Herbarium specimens of Porina nucula which I examined generally had a smooth to verrucose thallus, but the Long island specimens had diffuse coralleid thalline outgrowths. These outgrowtha appear like Iggntepchlia filaments which have partially escaped lichenization° Swineccw (in a letter) said that these specimens were poorly developed but otherwise normal, and so, perhaps l?6 the condition is not as unusual as it first appears. Its ecology is the same as that of E. hibernica. Distribution ~ Gulf coastal plain (Fink, 1935}: Tropical element (see Swinscow, 1962)‘ Coastal Plain subelement; Europe (Swinscow, 1962). GEAFHZDACEAE XYLOGRAIHA Xylographa gpggrapgella_will. in Rathr. Pros. U. S. Nat. Mae. 7: 8. 1884. Material seen - SUFFCLK COUNTY: Crient, Latham 1;, 1 May 1914 (Latham); Orient, Latham logo, 1 April, 1915 (Latham‘. Norstictic acid was demonstrated in KOH from the specimen with a well developed thallus (latham lg) and this substance nndoubtedly is the basis of the KOH + red, PD + yellow reactions in the medulla of material examined by Lamb (1954). The other long Island specimen (EEEEEE 1080) had almost no thallus (but was identical in other respects to gatham lg) and norstictic acid could not be demonstrated in the very minute thalline particles which were present. fixlggggg 3 splat; g (Pers.) Zahlbr. differs from g. ifithifitli in having broader spores and longer lirellae. An exsiccat collection of K. abietina (California Fungi no. 853) had spores liwlh x 6~7 n with lirellae up to a millimeter or more long. No norstictic acid could be found in the specimen, which had virtually no thallus. Xylographa opegraphella is confined to old wood. Distribution - New England coast (Fink, lgfifi), Nova Scotia; Alaska (Rothrock, 1884; Cummings, 1910): Temperate element, Oceanic subelement; endemic. 197 GRAPHlfi Graphis scrigta (L.) Ach. Kgl. Vet. Akad. Nya Randi. l45. 180?. Lichen scriptus L. Sp. P1. 1140. 1753. Material seen a SUFFQLK CQUNTE: 52 specimens collected by Tmshaug andfor Brodo; 12 specimens collected by Latham (Latham). The lirellee of this species are extremely variable in length, breadth, and degree of branching. Gross lirelline characters are therefore of little use in defining the species. anphi§_§gggggi is common on the bark of various deciduous trees, usually in partial shade and is mainly associated with the red oak forest on the north Shore (figure 43). Distribution - Nova Scotia, Maine, .onnecticut, hassachusetts, North Carolina, Tennessee, thhigan, Wisconsin, lndiana, Minnesota, Washington, Alaska: Tem erate element North Temperate subelement (?); Eur)“&; Asia 9 l' . (Vainio, 1928; Zahlbruckner, 1933b). EHAEDGRAPHIS 4 . . 3'?“ q ... , Phaeagraohis dendritica (Ach.) MLLio Arg. rlora 63: 382. ...;— n F" m ’1) h.) o Qpegrapha dendritica Ach. Math. Lich. 31. pl. l, f. l0. 1803. material seen a NASSAU COUNTY: giggg l509 (lb), 546 (12), £54 (12). SUFFGLK COUNTY; 72 specimens collected by Imshaug and/or Brodo; 12 specimens collected by Latham (Latham); Greenport, batham glglfi, 12 April 1953 (33). (I) p. :3, m H The species is found on the bark of various deciduous tree. lighted or partially shaded woods. Distribution w Eastern United States (Fink, 1935}; Temperate element, East Temperate subelement; Europe; Asia (Zahlbruckner, 1930b). *— Diploschistes scru osus (Schreb.) Norm. Nyt. Mag. Naturv. 7: 232. 1853. Lichen scruposus Schreb. Spic. Fl. Lips. 133. 1111. var. scrngosus material seen w SUFFOLK COUNTY? ggggo 3§§l (76); Orient. Latham, 4 October ‘1: 1917, (Latham); Sag Harbor, 3th m, 10 May 1924 (Letham). it“ N a N ’55 ‘9 [’9 .L‘ 0 var. parasiticus (Semmerf.§ Zshl: . Cst. Lieh. Univ. 2: 6' '1’ . .¢,-~ A 1n. \ .- . . .... .~ . .n‘.‘ ..., f. r" p ’ I "r. ' .F‘ ' § — .‘ .7 . . ttgattia acru egg tat. parasitica acanerf. surpl. rl. laps. ids. t826. r1 5 Material seen ~ SUFFQLY COUNTY: 41g_§ ;§;;Zg {1008). Diploschistes scruposus was treated by Fisk {1935) in the genus Erceoiar‘a. Distribution e Maine, Connecticut, Michigan, Oklahoma, Arizona, black Hills, Washington, Manitoba, Baffin Island: Arctic~boreal element; circumboreal. GYALECTACEAE EKMERELLA Bimerella dilute (Pers.) Trev. Read. Beale Ist. Lomb. Sci.l3: 65. 1880. Peziza diluta Pers. Syn. Meth. Fang. 668. 18?”. Material seen a SUFFOLK COUNTY: Brodo 3200 (33). Both this species aad the oae following were treated under the genus Microphiale by Fink (1935). The Long Island specimen was found on the bark of an old oak in the dense shade of an gle§ opaga grove on Fire Island. Distribution w Eastern United States (Fink, 1935); Maine, North Carolina, Black Hills, Saskatchewan: Temperate element, East Temperate subelement (?); \ Europe; Asia (Vainio, 1928). *- 199 Dimerella lutea (Dicks.) Trev. Rend. Reale Ist. Lomb. Sci. 13: 66. 1880. Lichen luteus Dicks. Fasc. Pl. Crypt. Brit. 1: 11, pl. 2, f. 6. 1785. Material seen u SUFFOLK COUNTY: Orient, Latham 1087, 3 May 1914 (Latham). A specimen of this species was also found on Cape Cod (Massachusetts) in a bog (Brodo 4323 B). Distribution w Eastern United States, and Canada (Fink, 1910); Maine, North Carolina, Washington: Temperate element, East Temperate subelement (?); Europe; Asia (Vainio, 1928). COLLEMATACEAE COLLEMA Collema subfurvum (M311. Arg.) Degel. Bot. Not. 139. 1948. Synechoblastus flacgidus v. subfurvus Mflll. Arg. Proc. Roy. Soc. Edinb. 11: 457. 1882. Material seen w QUEENS COUNTY: Jamaica, G. B. Brainerd, 1866 (BKL); Jamaica, G. B. Brainerd, 1866, (BKL 031870). SUFFOLK COUNTY: Orient, Latham Zfilg, 3 May 1914 (Latham); Napeague, Latham 2845, 1 March 1927 (Latham); Montauk, Latham 28309, 9 February (Latham); Shelter Island, Latham 36949, 4 May 1943 (Latham). Collema subfurvum differs from closely related Q. laccidum (Ach.) Ach. (Syn. Synechoblastus rupestris Trev.) in having globular rather than squamiform isidia, and in its corticolous rather than saxicolous substrate preference (see Degelius, 1954). It is usually found on oak bark. Distribution - New England, Smoky Mountains (Tennessee), Iowa, Illinois (Degelius, 1954): Temperate element, Appalachian subelement, Appalachian:- Great Lakes unit; Europe (oceanic localities) and Asia (Degelius, 1954). 200 LEPTOGIUM Leptogigg corticola (Tay1.) Tuck. in Lea, Cat. P1. Cinc. 47. 1849. Collema corticola Tayl. Jour. Bot. 5: 195. 1847. Material seen - SUFFOIK COUNTY: Montauk, Latham 3993 (p.p.), 6 April 1927 (Latham). Degelius (1940) discusses the nomenclatural problems pertaining to this species. The Long Island material agrees well with the original description as well as Degelius9 additions to it. Sierk (1964) presents a detailed discussion of the species. Distribution a Temperate element, East Temperate subelement, Adriatic coast in Europe (map: Sierk, 1964). Leptogium cyanescens (Ach.) KSrb. Syst. Lich. Germ. 420. 1855. Collema tremelloides v. cyanescens Ach. Syn. Meth. Lich. 326. 1814. non Lichen cyanescens Pers. or Parmelia cyanescens Ach. (Degelius, 1935). Material seen a QUEENS COUNTY: Jamaica, G. B. Brainerd, 1866? (BKL). SUFFOLK COUNTY: ggggg ;;;§ (102); Orient, Latham 181, 3 May 1914, (Latham); Orient, Latham 8199, 16 April 1928 (Latham); Greenport, Latham 8618, 1 June 1931 (Latham); North Sea, Latham 23333, 26 March 1954 (Latham); Montauk, Latham £8399A, 9 February 1949 (Latham); Three Mile Harbor, di33 gggg, 11 May 1954 (NYS). This species was apparently included in L, tremelloides (L.) S. F. Gray by Fink (1935). Leptqgium tremelloides, however, is strictly an Old World species (Sierk, 1964). The confusing nomenclature of L. cyanescens has been clarified by Degelius (1935). The species' oceanic affinities are noted by Degelius (1935 and 1941). The distribution of L. cyanescens on Long Island (figure 35), showing a restriction to the foggy eastern tip, reflects these oceanic requirements. Sierk (1964) discusses its morphology, ecology, and 201 distribution in detail. On Long Island, the species is usually found on mossy tree bases. Distribution - Temperate element, basically East Temperate subelement with scattered occurrences in the Black Hills, western Canada and coastal Alaska, Europe, Asia (map: Sierk, 1964). PANNARIACEAE PLACYNTHIUM In her recent North American monograph of the genus, Henssen (1963) placed Placynthium into the Peltigeraceae based on ascocarp development. Since Hensson's revision of the cyanophycean lichens and their families is still not complete, the older family concepts will be retained for the time being. glgcynthium_nigrug_(Huds.) S. Gray Nat. Arr. Brit. P1. 395. 1821. Lichen gigg£_fluds. F1. Angl. ed. 2, 2: 524. 1778. Material seen - SUFFOLK COUNTY: §52g2_3221 (54). Although g, nigrum is considered squamulose or even subfoliose by some authors, the Long Island material was all crustose, occasionally forming small subsquamulose areoles. Henssen (1963) presents a detailed account of the species' morphology and development. This inconspicuous species is probably more abundant than the collection records show. It was found in a shaded woods on old concrete foundations. Distribution - Arctic-boreal element (map: Henssen, 1963); Europe; Asia (Lynge, 1928). PANNARIA Pannaria lurida (Mont.) Nyl. Mem. Soc. Sci. Nat. Cherb. 5: 109. 1857. Collema luridum Mont. Ann. Sci. Nat. II. 18: 236. 1842. 202 Material seen - SUFFOLK COUNTY: Montauk Woods north of Fresh Pond, Latham 2832L, 9 February 1949 (Latham); Orient Point, Latham 2, 4 April 1910 (NYS). COUNTY UNKNOWN: Long Island (?), AggLLg (BKL()31953). The species was found on oak and red cedar bark. Distribution - Eastern United States (Fink, 1935): Tropical Element (Zahlbruckner, 1925), Appalachian~Temperate subelement. STICTACEAE LOBARIA Lobaria pulmonaria (L.) Hoffm. Deutschl. F1. 146. 1796. Lighgg_ pulmonarius L. Sp. P1. 1145. 1753. Material seen - QUEENS COUNTY: Ridgewood, G. B. Brainerd, 1867 (BKL 031881). SUFFOLK COUNTY: Brodo 887 (56), LQLL (112), 1945 (112), LL§4_(102); 12 specimens collected by Latham (Latham). Fink (1935) treated L, pulmonaria under the genus SLLng, This species shows some variation in isidia and soredia production in various parts of its range. The Long Island specimens all have isidiate- soralia on the thallus margins and ridges but they may be more common in some individuals than in others. The granular soredia are sometimes hard to see until most of the isidia have fallen away. The isidia vary from being short, almost like papillae, to elongate cylindrical, and finally coralloid. The species is rare on Long Island. It is confined to tree bases in the oceanic areas of the eastern tip of the island and bog trees (especially Acer rubrum) outside this area. Degelius (1935, p. 223) stated that L, pulmonaria favors an oceanic climate but is not restricted to an oceanic distribution. Distribution - Nova Scotia, Maine, Connecticut, Tennessee, Michigan, Ontario 9 Indiana, washington, British Columbia, Alaska: Temperate element, North 203 Temperate subelement (Appalachian-Great Lakes: Hale, 1961a); Europe; Asia (Zahlbruckner, 1930; Magnusson, 1940). Lobaria quercizans (Ach.) Michx. Fl. Bor.-Amer. ed-Z. 324. 1820. Parmelia quercizans Ach. Lich. Univ. 464. 1810. Material seen - KINGS COUNTY: New Lots, (Brainerd ?), 1867 (BKL 031874). SUFFOLK COUNTY: ngdg_LQ§Q (112), L142 (102), L§QL (102); Napeague, nghgm L838, 1 March 1927 (Latham); Napeague, Latham 2837, 1 March 1927 (Latham); Montauk, Latham 28307, 9 February 1949 (1945?) (Latham, MO); Riverhead, LEEDEE:§§§§2: 16 May 1960 (Latham); Montauk, Latham 36884, 4 April 1949 (Latham); Jamesport, Latham 36948, 19 April 1951 (Latham); (no locality), Leggygg, 6 May 1920 (Latham); Eastport, Schrenk, 28 June 1894 (MO). I Lobaria quercizans is the North American vicariad of L, amplissima (Degelius, 1940; Hale, 1957a), a well-known European oceanic species (see Degelius 1935). Lobaria quercizans was considered under the latter name in Fink (1935). The North American species also appears to have an oceanic distribution (Degelius 1941), and on Long Island is restricted to the fog belt and bogs. It often is found associated with Lobaria pulmonaria. Distribution - Temperate element, Appalachian subelement, Appalachian- Great Lakes unit (map: Hale, 1957a); endemic. NEPHROMACEAE NEPHROMA Nephroma laevigatum Ach. Syn. Lich. 242. 1814. non auct. Material seen - SUFFOLK COUNTY: Montauk, La_gt_1_a_m_ §_6_7_8_4, 6 May 1929 (Latham). A full discussion of the taxonomy and distribution of this species was 204 presented by Wetmore (1960). Long Island is the southernmost locality for the species on the east coast. The specimen was found on rock. Distribution - East and west coasts of North America: Temperate element, Oceanic subelement (Wetmore, 1960); oceanic regions of Europe (Degelius, 1935); Asia (Vainio, 1928). PELTIGERACEAE SOLORINA Solorina saccata (L.) Ach. Kgl. Vet. Akad. Nya Handl. 228. 1808. Lichen saccatus L. F1. Suec. ed. 2, 419. 1755. Material seen - SUFFOLK COUNTY: Montauk, Latham 36883, 7 October 1926 (Latham). The specimen was collected on a rocky bank. Distribution - Michigan, Ontario, Minnesota, Black Hills, Washington, Alaska, Manitoba, Quebec, Baffin Island: Arctic-boreal element; circumboreal. PELTIGERA Peltigera aphthosa (L.) Willd. F1. Berol. Prodr. 347. 1787. Lichen aphtosus L. Sp. P1. 1148. 1753. var. variolosa (Mass.) Thoma. Trans. Wisc. Acad. Sci. 38: 253. 1947. Peltigera aphthosa f. variolosa Mass. Sched. Crit. III: 64. 1856. Material seen - KINGS COUNTY: New Lots, G. B. Brginerd (BKL,031888). SUFFOLK COUNTY: Gardiner's Island, Lagggg, 22 September 1922 (Latham); Fisher's Island, Latham, 24 June 1929 (Latham); Montauk, Latham, 17 May 1942 (Latham). The dark veins on the lower surface of this variety distinguish it from var. aphthosa (var. typica in Thomson, 1950a). 205 The material is from the ground in dry woods, and from a rock. Distribution - Arctic-boreal element, circumboreal (map: Thomson, 1950a). Peltigera canina (L.) Willd. F1. Berol. Prodr. 347. 1787. 2853531 caninus L. Sp. P1. 1149. 1753. var. rufescens (Weiss) Mudd, Man. Brit. Lich. 82. 1861. Lichen caninus var. rufescens Weiss,P1. Crypt. F1. Goet. 79. 1770. Material seen - NASSAU COUNTY: Massapequa, S. Cain L88, 7 July 1935 (NY). SUFFOLK COUNTY: Devon, nghgm, 2 May 1955_(Latham); Three Mile Harbor, Latham 27207, 17 April 1947 (Latham); Napeague, north of Fresh Pond, Latham 8118, 6 April 1938 (Latham); Three Mile Harbor, Hands Creek, LQEEEE. 8848, 20 April 1926 (Latham); Napeague, Latham 36978, 3 May 1947 (Latham). Following Thomson (1950a), three varieties of this species can be recognized as occurring on Long Island. The questionable taxonomic rank of var. rufescens is discussed on pages 119- 120. On Long Island it is relatively rare occurring mainly in dry woods on tree bases. var. £22£ g (Ach.) Schaer. Lich. Helvet. Spicil. 6: 265. 1833. Lichen spurius Ach. Lich. Suec. Prodr. 159. 1798. Material seen - SUFFOLK COUNTY: ggggg_§2;;§g (53), 882; (87); Northwest, LEEEEE;£§l§§: 10 April 1947 (Latham). variety gpggig has only been collected in its sorediate stage on Long Island. It has been clearly established that the sorediate form is a juvenile stage of var. gpgg;g_(Dahl, 1950). Lagggg 88188 has both apothecia and soredia and appears similar to var. rufescens which in turn is said to intergrade with var. gaging (var. albescens) (Thomson, 1950a). It was found on dry sandy soil. 206 var. ulorrhiza (F13rke) Schaer. Enum. Crit. Lich. Europ. 20. 1850. Peltidea ulorrhigg.F13rke, Deutsch. Lich. no. 154. 1821. Material seen - SUFFOLK COUNTY: Riverhead, Latham, 1 November 1913 (Latham). The Latham specimen was found on the ground in a dry woods. Distribution (of all varieties) - Arctic-boreal element; circumboreal (maps: Thomson, 1950a). Peltigera polydactyla (Neck.) Hoffm. Desc. Adumbr. P1. Lich. 1: 19, pl. 4, f. 1. 1790. Lichen polydactylon Neck. Math, Phsc. 85. 1771. Material seen - QUEENS COUNTY: Jamaica, G. B. Brainerd, May 1866 (BKL 031889). SUFFOLK COUNTY: Montauk, ngggg, 12 May 1920 (Latham). Peltiger§_poly8§cty1a is most closely related to g, horigontglis from which it is distinguished by its vertically oriented apothecia and its longer narrower spores. Thomson (1950a) reports the spores of the latter species to be 24-45 x 3.5 - 6 n. The Long Island material, having broad conspicuous veins on the lower thallus surface, represents var. polydactyla (var. 212122 of Thomson, 1950a). Latham's specimen was found at the base of a tree in an oak woods. Distribution - Baffin Island, Manitoba: Arctic-boreal element (map: Thomson, 1950a), circumboreal. Peltigera praetextata (Flgrke in Sommerf.) Vain. Termeszetr. Fuzetek 22: 306. 1899. Peltidea ulorrhiza var. praetextata Flsrke in Sommerf. Suppl. Fl. Lappon. 123. 1826. Material seen - SUFFOLK COUNTY: §£2§2.l252 (112), L888 (48), 888; (45), 8888 (102), 8882 (23); Greenport, LEEEEEMEQ: 10 May 1914 (Latham); Three Mile Harbor, Latham, 21 November 1926 (Latham); Riverhead, Latham, 2 February 207 1923 (Latham); 16 specimens collected by Latham (Latham). This species is very similar to 8, canina var. rufescens and seems to differ only in its ability to produce regeneration squamules on the thallus surface and margins. Experiments on the production of isidia (regeneration squamules) were performed by Thomson (1948) and Lindahl (1953) and resulted in two entirely opposite points of view regarding the taxonomic value of the structures. Thomson found that wounded thalli of 8. canina var. rufescens regenerate on some lobes and not on others whereas Lindahl found that no thalli of 8, canina sens. str. regenerated and only 8, praetextata showed regeneration. It is possible that Thomson was working with true ‘8. praetextata "hidden" by its original lack of regeneration squamules (which Lindahl concedes may happen) and that true 8, canina sens. strawwould not produce regeneration even in the United States. It is also possible that only under certain conditions will species other than praetextata regenerate and these conditions were met in Wisconsin and not in Sweden, or that the America populations of‘L. $22222 differ in regeneration properties from the European populations, a difference which might be of taxonomic importance. This entire problem as it appears to me, is far from settled and should be investigated further. until more work is done, however, the European concept of 8, praetextata will be accepted. The species is most frequently found growing on mossy tree bases in oak woods. Distribution - Arctic-boreal element (map: Thomson, 1950a); Europe; Asia (Magnusson, 1940). 208 LECIDEACEAE LECIDEA Lecidea aeruginosa (see page 212). Lecidea albocaerulescens (Wulf. in Jacq.) Ach. Meth. Lich. 52. 1803. Lichen albocaerulescens Wulf. in Jacq. Collect. Bot. 2: 184, f. l. 1788. Material seen - NASSAU COUNTY: Brodo 549 (12). SUFFOLK COUNTY: Brodo 8888 (99), 8888 (20), 8888 (96), 8888_(110), 8888 (111), 8888 (17), 8888 (50), 8888 (50), 8888 (34), 8888 (119), 8888 (119), 8888_(l34), 8888 (62); Gardiner's Island, Lagggg, 30 March 1921 (Latham); Greenport, Latham 3966, 1 April 1927 (Latham); Greenport, L88888_88, 10 May 1914 (Latham); Three Mile Harbor, Latham 88888, 25 May 1954 (Latham). This striking saxicolous species is easily identified in the field by its pruinose apothecia, dark apothecial margins, and smooth grey thallus. All but one specimen on Long Island was shown to contain stictic acid (both by paper chromatography and recrystallization in GAoT solution). The exception (88888_888) contained norstictic acid (red acicular crystals in KOH). This stictic-norstictic shift is a common phenomenon in lichen chemistry and can be seen in Lecanorg_cinerea, and several species of Parmelia. Norstictic acid has never been reported for this species before. Lgcidea albocaerulescens is narrowly restricted to shaded granitic rocks,.and is only found in the poorly lighted red oak forests of the north shore. Distribution - Connecticut, Tennessee, Michigan, Indiana, Oklahoma, Minnesota, Washington, Alaska; Eastern United States and Washington (Fink, 1935): Temperate element, North Temperate subelement; Europe; Asia (Lynge, 1928). Lgeidea anthracophila Nyl. Flora 48: 603. 1865. Material seen - SUFFOLK COUNTY: 23 specimens collected by Imshaug and/or Brodo. 209 Fink (1935) lists this species with the genus 88858, The PD + red constituent of L, anthracophila is apparently fumar- protocetraric acid but does not show exactly the same Rf value as known fumarprotocetraric acid (as in Cladonia subtenuis) in paper chromatography (solvent: pyridine, ethyl acetate and water). Fumarprotocetraric acid usually has an Rf of approximately 0.30 to 0.45 and the Lecidea anthracophila material has an Rf of approximately 0.40 to 0.55. In all other characters (color reaction with PD, fluorescence in UV before and after reaction with PD, etc.) it is identical to fumarprotocetraric acid. The species is found only on fresh or charred bark of Pinus rigida (see page 56). Fink (1935) reported it from old wood. Distribution - Vermont, Massachusetts, New Jersey, and North Carolina (Fink, 1935): Temperate element, East Temperate subelement; Europe. Lecidea botryosa (Fr.) T. Fr. Lich. Scand. 1: 454. 1874. Biatora botryosa Fr. Kgl. Vet. Akad. Nya Handl. 268. 1822. Material seen - NASSAU COUNTY: 85888_8888_(4). SUFFOLK COUNTY: Imshaug 8§_6_38 (Nu of 29), gm (m of 29), Mia); (33). When sterile, this species closely resembles L, aeruginosa which, however, is C + red. If apothecia are present, the hypothecial color (hyaline in L, aeruginosa and brown in L, botryosa) distinguishes the two. The species is almost entirely restricted to old wood. It was found once (Brodo 3494) growing on the base of an old black oak in a shaded woods. Distribution - Michigan, Arizona, Manitoba; Adirondack Mountains of New York, New Hampshire with doubtful occurrences in the west coast (Lowe, 1939): Temperate element, North Temperate subelement; northern Europe, Asia (Lowe, 1939). 210 Lgcidea coarctata (Turn. in Sm. & Sowerby) Nyl. Act. Soc. Linn. Bord. 21: 358. 1856. Lichen coarctatus Turn. in Sm. & Sowerby Engl. Bot. 8: pl. 534. 1799. Material seen - QUEENS COUNTY: 88888.88; (3). SUFFOLK COUNTY: 85888 .s_9-_30§ (54). s_9_-;_3_1_q (54). 12; (90A). zeta (44), m (110). use (127). 8888_(49), 8888 (111), 8888 (112); Orient, nghgg, 18 March 1914 (Latham); Shelter Island, Latham 22177, 26 October 1944 (Latham); Montauk, ngggg 88888, 8 October 1954 (Latham). This is the only species on pebbles having small brown apothecia. The white, areolate, C + red thallus add to its distinctiveness. Lecidea coarctata is often associated with L. erratica and 888gocarpon obscuratum on pebbles and small stones. Distribution - Nova Scotia, Maine, Connecticut, Indiana, Minnesota, British Columbia; northern United States (Fink, 1935): Temperate element, North Temperate subelement; Europe. Legidea cyrtidia Tuck. Proc. Amer. Acad. Arts Sci. 12: 181. 1877. Material seen - SUFFOLK COUNTY: 85888_8888 (88), 8888 (44), 8888 (110), 88888,(128), 8888 (34), 8888 (34), 8888 (119), 8888 (112); Greenport, L88888 8888, 1 April 1927 (Latham); Shelter Island, Latham 22177, 26 October 1944 (Latham); Shelter Island, LEEEEE.§Z§125; 26 October 1944 (Latham); Shelter Island, LEEEEE.ZZ§§Q; 26 October 1944 (Latham); Shelter Island, Latham 22882, 26 October 1944 (Latham); Latham 31015,2 February 1940 (Latham); Mbntauk Point, 888888, 12 April 1956 (Latham). Lecidea cystidia is superficially very similar to L, erratica. However, the epithecium and the outer portions of the exciple are greenish-black in the latter and brown in the former species. Magnusson (1952) described L. neariggii which, from its description, appears very similar to L, cygtidia. # 211 Lecidea nearingii has a brown-black thallus whereas L, cyrtidia has a pale to dark brownish-green thallus. The thalli of both species are thin and continuous. The distinctions are therefore very questionable from the published descriptions, but since the type of L, nearingii has not been examined no further conclusions can be made concerning its validity as a species. One specimen had much larger apothecia than any of the others, but agreed in other respects with the descriptions of L, cyrtidia. The species is common on pebbles and small stones in dry woods or fields. Distribution - Eastern United States (Lowe, 1939): Temperate element, East Temperate subelement; endemic. Lecidea erratica ngb. Parerg. Lich. 223. 1861. var. erratica Material seen - QUEENS comma Brodo _s_2_4_ (3). NASSAU comm: Brodo 45 (12), 8888 (10). SUFFOLK COUNTY: 32 specimens collected by Imshaug and/or Brodo; Shelter Island, LSEEEE.Z§§12§9 26 October 1944 (Latham); Shelter Island, Latthgg 22888, 26 October 1944 (Latham); Riverhead, E8888 88888, 16 lurch 1946 (Latham); Quogue, Lech—ea 88888 (Latham); Montauk Point, _I£t__tm_n_1 88888, 6 May 1949 (Latham); Orient, Brown Brothers Site, LEEEEE:Z§2Z§3 4 November 1951 (Latham); Riverhead, North River, LEEDEE:§$§Z£: 16 March 1946 (Latham); East of Sag Harbor, Legggg, 19 October 1945 (Latham). var. planetica (Tuck.) Lowe, Lloydia 2: 279. 1939. Lecidea glanetica Tuck. Syn. N. Am. Lich. 2: 131. 1888. Material seen - SUFFOLK COUNTY: 8:888 8818 (17). Magnusson (1936) recognized several species as being closely related to L. erratica: L. sylvicola Flot., L. czrtidea Tuck., L. micytho Tuck., and L, plantetica Tuck. Lowe (1939) whose work is being followed here, treats L, planetica as a variety of L, erratica having a more well-developed thallus 212 than the variety erratica. Lowe regards L, micytho as a yellowish form of var. planetica. The separations of L. erratica and L. cyrtidea have already been discussed under the latter species. L, sylvicola differs from L, erratica in having a greenish or greenish black hypothecium with a doubtfully dis- tinguished pale bluish-black exciple (Lowe, 1939) as opposed to a reddish- brown to almost black hypothecium and an exciple greenish-black externally and hyaline within. This common species is found on pebbles in exposed fields and downs, and is particularly abundant in well-lighted areas on the Ronkonkoma moraine. Some observations on its development have been presented on pages 47-48. Distribution - Eastern United States west to Minnesota (Lowe, 1939): Temperate element, East Temperate subelement: Europe (ibid). Lecidea aeruginosa Borr. in Hook. and Sowerb. Suppl. Engl. Bot. 1: tab. 2682. 1831. Lecidea flexuosa (Fr.) Nyl. Act. Soc. Linn. Bord. 21: 356. 1856. uterial seen - SUFFOLK COUNTY: $3318.23 88888 (86), B_rg§_o_ 888 (79), 8888 (70). _1_§_I_2_ (69). 22;; (61). '_2_;3_3 <44). _2_s_4a (73). 21;; (111). £295; (95). 8888 (129), 8888 (18), 8888 (49). The separation of sterile material of L, aergg8nosa from L, botrzosa was discussed under the latter species. Laundon (1962) regarded L, aeruginosa (sub L, flexuosa) as synonomous with L, guadricolor (syn. L, granulose). On Long Island, however, except for spore size, the two are not at all similar either morphologically or ecologically. L, aeruginosa has black or lead- colored, plane apothecia each with a thin hyaline hypothecium whereas L, guadricolor has large brown, irregularly convex to almost hemispherical apothecia each with a thick opaque hypothecium. In addition, the former 213 species is restricted to lignum and the latter is found only on sandy soil. I have examined material from the Black Hills of South Dakota where both species occur on old wood and still the fertile material of L, aeruginosa is easily distinguished from L, quadricolor. Distribution - Connecticut, Minnesota, Black Hills; throughout the United States (Fink, 1935): Temperate element, North Temperate subelement (?); Europe; Asia (Vainio, 1928). Lecidea macrocarga (DC. in Lam. & DC.) Steud. Nomencl. Bot. 245. 1824. Patellaria gacrocarpa DC in Lam. & DC. F1. Franc. ed. 3. 2: 347. 1805. Material seen - SUFFOLK COUNTY: Imshaug 28888 (52). There has been much disagreement concerning the name of this species. Fink (1935) discussed it under the name L, platycarga Ach. and Lowe (1939) following Vainio (1909, 1934) used L, steriza (Ach.) vain. Clauzade and and Rondon (1959) recently considered the species under the name Lecidea contigua (Hoffm.) T. Fr. Most other workers have used L, macrocarpa. The epithet Wmacrocarga" was first used at the species level in the genus Patellaria by DeCandolle in 1805 which makes it the oldest available name. "Steriza" was only considered at the infraspecific level (L, confluens 5,L, steriza Ach.) until Vainio raised it to a species in 1909. Lecidea platycarpa was not described until 1810 by Acharius. vainio (1934) states that Theodor Fries used the name contigua incorrectly in referring Hoffmann's Verrucaria contigua to the genus Lecidea. Fries' lichen was L, macrocarpa but Hoffmann's name referred to a different species. The Long Island specimen has a rather well-developed continuous to cracked and areolate thallus. It was found on a siliceous roadside pebble. Distribution - Nova Scotia, Maine, Connecticut, Tennessee, Michigan, Minnesota, Idaho, Alaska, Saskatchewan, Baffin Island: Arctic-boreal element; circumboreal. 214 Lecidea myriocarpoides Nyl. Flora 48: 355. 1865. Material seen - SUFFOLK COUNTY: §£2§2.Z§§§.(49): 8888 (42), 8888_(62), 8888_(112). This species was found only on well illuminated hard lignum. Distribution - Eastern United States and California (Fink, 1935, Lowe, 1939); Europe. Lecidea nylanderi (Anzi) T. Fr. Lich. Scand. 1: 462. 1874. Biatora nylanderi Anzi, Cat. Lich. Sondr. 75. 1860. Material seen - SUFFOLK COUNTY: Brodo 1400 (65), 8888 (85),8888.(51), 88888,(73). The very small reddish-brown apothecia and the subglobose to globose spores of this species easily distinguish it from other pine bark lichens. On Long Island, it is limited to the bark of Pinus rigida. Culberson (1958a) studying the pine-inhabiting lichen vegetation of North Carolina found Lgcidea nylanderi only in the mountains of North Carolina. 88888_58g888_ is also found only in the mountains. This correlation may indicate a very high degree of substrate specificity, but since the specificity of the species was not indicated in that paper, and since other pines occur in the mountains, no such conclusion can be made. The species is found on Pinus ponderosa in the Black Hills of South Dakota and was also collected twice (§£2§2.&122: 8888) on Pinus rigida in the Cape Cod region of Massachusetts. Distribution - Adirondack Mountains of New York, Massachusetts, California, (Lowe, 1939); North Carolina, Wisconsin, Black Hills, Manitoba: Temperate element, North Temperate subelement; Europe; Asia (Vainio, 1928). Lecidea quadricolor (Dicks.) Borr. ex Hook. in Sm. Eng. Fl. 5: 182. 1833. Lichen qggdr8color Dicks. Fasc. P1. Crypt. Brit. 3: 15, tab. 9, fig. 3. 1793. 215 Material seen - SUFFOLK COUNTY: Imshaug 25644 (64), 25656 (64), 25788 (86); Brodo 888 (79), 1900 (114), 1939 (85), 3372 (94), 3401 (75), 1404 (83); Shinnecock Hills, Latham 7873, 14 February 1938 (Latham); Southold, L§£E§!;Z§§§a 11 February 1938 (Latham); North Sea, Latham 28128, 16 May 1955 (Latham); Riverhead, 8888, September (NYS). This species has generally been treated under the name L. granulose. Laundon (1962) discusses the nomenclature. Lecidea qgadricolor is similar in some respects to L, aeruginosa but the two are quite distinct on Long Island (see discussion under L, aeruginosa). It is known to grow on old wood as well as soil but is restricted to sandy soil on the island. Distribution - Nova Scotia, Maine, Connecticut, North Carolina, Michigan, Minnesota, Arizona, Black Hills, Washington, Alaska, Saskatchewan; Northern United States (Fink, 1935): Temperate element, North Temperate subelement; Europe; Asia (Vainio, 1928). Lecidea scalaris (Ach.) Ach. Math. Lich. 78. 1803. L88888 scalaris Ach. Kgl. Vet. Akad. Nya Handl. 127, tab. 5, f. 2. 1795. Material seen - NASSAU COUNTY: §E2§2.§29§,(10)- SUFFOLK COUNTY: 19 specimens collected by Brodo and/or Imshaug. As with L, anthracophila, this species is treated under 88858 by Fink (1935). Lecidea scalaris has a high specificity for the bark of 88888_58g888 but is not restricted to it. Barkman (1958, p. 38) and Lowe (1939) state that the species is commonly found on burned wood and this is certainly true on Long Island where it is often found on charred pine bark (see page 50). éEEE saccharinum and Acer rubrum were the preferred substrates in an area in central New York (Brodo, 1959). The reasons for these substrate preferences are not clear, although all these substrates are highly acid. Ochsner 216 (1928 in Barkman, 1958 p. 102) stated that L, scalaris is nitrophobous, but this is yet to be proven. Distribution - Central New York, North Carolina, Arizona, Black Hills, Washington, Saskatchewan: Temperate element, North Temperate subelement (?); Europe; Asia (Lowe, 1939). Lecidea uliginosa (Schrad.) Ach. Math. Lich. 43. 1803. 8888881 uliginosus Schrad. Spic. F1. Germ. l: 88. 1794. Material seen - SUFFOLK COUNTY: 22 specimens collected by Imshaug and/or Brodo. Laundon (1961) discusses in detail the similarities of this species with Lecidea oligotropha Laund. The latter is mainly characterized by its coarsely granulose to verruculose, pale brown to yellowish thallus. In contrast, L, uliginosa has a finely granular to almost leprose dark brown to black thallus. Only one North American specimen of L, oligotropha (from Minnesota) is cited by Laundon. Lecidea uliginosa often forms conspicuous tar-like patches on partially stabilized sand. Closer examination will reveal tiny black apothecia scattered among the dark brown thalline granules. Alvin (1960) reported the species as occurring in dune communities in southern England, especially in the heath, ecologically very similar to some of the Long Island habitats. Distribution - Nova Scotia, Connecticut, Michigan, Indiana, Minnesota, Black Hills; throughout United States (Fink, 1935): Temperate element, North Temperate subelement (?); Europe; Asia (Vainio, 1928). Lecidea ygrians Ach. Syn. Math. Lich. 38. 1814. Material seen - SUFFOLK COUNTY: 37 specimens collected by Imshaug and/or Brodo; East Marion, Latham 88 (22249), 3 May 1914 (Latham); Greenport, Latham 3984, 1 April 1927 (Latham); Orient, Latham.88, 30 May 1914 (Latham). 217 This species occurs on the bark of various trees from completely exposed dune areas to protected oak forests. Distribution - Nova Scotia, Connecticut, Michigan, Minnesota, Washington, Manitoba; throughout the United States (Fink, 1935); France (Acharius, 1814). Lecidea vernalis (L.) Ach. Meth. Lich. 68. 1803. Lichen vernalis L. Syst. Nat. 3: 234. 1768. Material seen - SUFFOLK COUNTY: Imshaug 25752 (132), Brodo 888 (55), 888 (47), 8888 (61); Greenport, Latham 1998 (22247), 27 February 1927 (Latham); Greenport, Latham 22254, 14 May 1914 (Latham); Greenport, ngggg, 1 March 1923 (Latham). This species is distinguished by its strongly convex, pale apothecia and its fusiform, occasionally one-septate spores. It is not common on Long Island, but where it occurs, it often covers large portions of the tree trunk. It is found on the bark of various trees particularly in rain tracks or in other equally moist or humid microhabitats. Outside of Long Island, the species is known to occur over moss. Distribution - Nova Scotia, Connecticut, Michigan, Wisconsin, Minnesota, Arizona, Alaska, Manitoba, Baffin Island: Arctic-boreal element; circumboreal. ngidea viridescens (Schrad. in Gmel.) Ach. Meth. Lich. 62. 1803. 888888_viridescens Schrad. in Gmel. Syst. Nat. 2(2): 1361. 1791. Material seen - SUFFOLK COUNTY: §£222.§Ql§ (17). This rare species was found growing over rotting wood. It somewhat resembles a Lepraria in its granulose, effuse thallus. Distribution - Michigan, Minnesota, Arizona, Alaska; Eastern United States and California (Fink, 1935): Temperate element, North Temperate subelement; Europe; Asia (Vainio, 1928). 218 CATILLARIA Catillaria g8auconigrans (Tuck.) Hasse, Bryol. 12zl02. 1909. Biatora glauconigggns Tuck. Proc. Amer. Acad. Arts. Sci. 12: 179. 1877. Material seen - SUFFOLK COUNTY: Brodo 59-272 (53), 88888,(73). This species, rare on Long Island, was found only on pine bark. Nearing (1947) stated that it is an oak-and pine-dwelling lichen. It is therefore surprising and noteworthy that Thomson (1951) reported the species as growing on the bark of aspens (ngulus tremuloides). This substrate usually bears a typically neutrophytic community of Caloglaca spp., Physcia spp., and Xanthoria spp. quite opposite from the communities on highly acid conifer bark. Distribution - Massachusetts and California (Fink, 1935), New York (Nearing, 1947), Michigan, Arizona, Manitoba: Temperate element, North Temperate subelement; endemic. BACIDIA Bacidia atrogrisea (Del. in Hepp) K3rb. Parerg. Lich. 133. 1860. Biatora atrogrisea Del. in Hepp, Flecht. Europ. 26. 1853. Material seen - SUFFOLK COUNTY: Orient, L2£E§E.§i§» 10 May 1914 (Latham); Orient, Lathag 791, 5 April 1914 (Latham); Orient, ngggg, 6 May 1915 (Latham); Orient, 888888, 10 April 1921 (Latham). Bacidia luteola (Schrad.) Mudd (syn.,8, rubella [hoffmj Mass.) and 8, fuscorubella (Hoffm.) Banach. can sometimes be confused with 8, atrogrisea. Bacidia luteola is distinguished by its uniformly brown or reddish apothecia often with a conspicuous white pruina, and 8, fuscorubella differs in having a dark brown hypothecium. Both these species have thicker thalli than 8, atrogrisea. The Long Island material agrees very well with the Migula exsiccat, Krypt. Germ. no. 52. 219 The species is usually found on the bark of various coniferous trees. Thomson (1951) reported it from Michigan on 88818_bark. Distribution - Connecticut, Michigan; Eastern United States (Fink, 1935): Temperate element, East Temperate subelement; Europe; Asia (Ikoma, 1957). Bacidia chlorantha (Tuck.) Fink, Cont. U. S. Nat. Herb. 14: 91. 1910. Biatora chlorantha Tuck. Syn. Lich. New Engl. 60. 1848. Material seen - NASSAU COUNTY: 88888_888 (11). SUFFOLK COUNTY: §£222 8888.(51), 8888_(ll3), 8888 (61), 8888 (sterile) (134), 8888 (134), 8888 (sterile) (66). Lamb (1954) presented a description and a discussion of this species and Thomson (1951) compared it with 8, chlorococca with which it is sometimes confused. The Long Island material agrees well with Lamb's description of the specimens from Nova Scotia. This species is often found sterile, but with many clusters of minute brown pycnidia containing pycnoconidia measuring 1.2 x 0.5 u. It is found on the bark of various species of deciduous and coniferous trees. Distribution - Nova Scotia, Connecticut, Smoky Mountains of North Carolina and Tennessee, Michigan; New England, New York, Ohio, Illinois, Minnesota (Fink, 1935): Temperate element, Appalachian subelement, Appalachian - Great Lakes unit; endemic. Bacid88_chlorococca (Graewe in Stizenb.) Lett. Hedw. 52: 131. 1912. Lecideg_chlorococca Graewe in Stizenb. Nova Acta Acad. Leop. Carol. 34 (2): 24. 1867. mterial seen - NASSAU COUNTY: my; 15.3.9. (16), .525. (12), 5_§_§ (11), 8898 (15). SUFFOLK COUNTY: 75 specimens collected by Imshaug and/or Brodo; Riverhead, Latham, 1 May 1960 (Latham). 220 Degelius (1940) described the spores of his material from Maine as slightly smaller than those of typical European specimens although the specimens from Long Island fit the spore size of the European material well. The species is found on a variety of substrates including twigs and bark of coniferous and deciduous trees as well as old wood. It is found in exposed and shaded localities. Distribution - Maine, central New York, North Carolina, Michigan, Wisconsin: Temperate element, Appalachian subelement, Appalachian-Great Lakes unit; Europe. Bacidia chlorostica (Tuck.) Schneid. Guide Study Lich. 109. 1898. Lecidea chlorostic8_Tuck. Proc. Amer. Acad. Arts Sci. 5: 419. 1862. Material seen - SUFFOLK COUNTY: 88888_8921|(78); Riverhead, L88888_8888, 24 June 1924 (Latham). This rare species is distinctive in many ways. Its paraphyses appear to be branched giving the apothecium an sscolocular appearance, but the olivaceous, minutely verruculose to subgranulose thallus, the large-celled Trebouxioid phycobiont, and the lack of thick-walled asci all are characteristic of Bacidia and not Micarea. The apothecia are small, lead-black, and convex, with the margin disappearing. They are sessile or buried in the granular crust, or sometimes they become more or less stipitate. The hypothecium is dark brown becoming sordid blackish-violet below and olivaceous above in KOH. The margins are reddish-violet in KOH. The species is apparently restricted to Chamaecygaris thyoides, at least in the coastal plain region. Three specimens (Brodo 3676, 3765, 3768) were collected in southern New Jersey on white cedar in cedar bogs just as they were on Long Island. 221 Distribution - Connecticut; Massachusetts, South Carolina, Illinois (Fink, 1935); New Jersey (see above): Temperate element, Coastal Plain subelement; endemic. 8§gidia integgedia (Hepp in Stizenb.) Arn. Flora 54: 54. 1871. non Hampe in Mass. Biatora anomala var. intermedia Hepp in Stizenb. Nova Acta Acad. Leopold. - Carolin. 30 (3): 42. 1863. Material seen - SUFFOLK COUNTY: 85888_8888 (33); Orient, L££2é2,§fl» 10 May 1914 (Latham). It seems clear from published descriptions that the Long Island material belongs to what Tuckerman (1888, sub Biatora), Fink (1935), and Erichsen (1957) have called Bacidia effuse (Sm. in Smo& Sowerby) Trev. However, there are a number of problems involved in the use of the name 8, 888888, To begin with the epithet 888888 cannot be used for any Bacidia since its basionym, Lichen effusus Sm. in Sm. & Sowerby (1808), is a later homonym of £12222 effusus Ach. (1798), a synonym of Lecanora saligna (Schrad.) Zahlbr. Secondly L88888 effusus 8m. in Sm. & Sowerby is listed as a synonym of 8898888 arceutina (Ach.) HArn." by vainio (1922). The latter species as described by Vainio differ in many respects from the Long Island specimens. These Long Island specimens do fit Vainio's description of Bacidia intermedia ("Hepp’0» Arn. and they agree almost perfectly with the Rabenhorst exsiccat no. 509 (distributed as 8gcidia effuse) cited by Vainio as typical 8, intermedia. Still another name which must be considered is 8gcidia albescens (Hepp) Zwack. which some authors (e.g., Arnold, 1884) considered as including intermedia as only a form with flatter larger apothecia. Erichsen (1957) used all four names (8, effuse, 8, arceutina, 8, intermedia, and 8, albescens) as separate species distinguishing them as follows: 222 1. Apothecia at first light, darker in age, never black; spores mostly 20 - 50 u long . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 1. Apothecia soon or from the beginning dark to black; spores mostly 4O - 60 u long (but f. brevispora is 25 - 39 u long). . . 8, arceutina 2. Apothecia whitish, flesh-colored, or rose. . . . . . . . . . . . 3 2. Apothecia light brown, brick red, darker in age. . . . . 8. effuse 3. Hymenium 50 - 60 u high; spores 34 - 48 u long7; apothecia remaining flat, 0.3 - 0.4 (0.6) mm in diameter. . . . . . . . . . . 8, intermedia 3. Hymenium 35 - 50 u high; spores 20 - 33 u long; apothecia first flat, then soon convex and marginless, 0.2 - 0.4 mm in diameter.8, albescens Fink (1935) did not use 8, intermedia. He separated 8, arceutina, 8. §££2££.8nd 8, albescens as follows: 1. Spores rarely more than 40 u in length. . . . . . . . . . . . . . . 2 l. Spores rarely less than 40 u in length (35 - 50 u). Hypothecium yellowish; disks pale light brown to blackish. . . . . . . . . . . . . 8, arceutina 2. Hypothecium hyaline; apothecia not more than 0.5 mm across; disk light pink to pale reddish. . . . . . . . . . . . . . . .8. albescens 2. Hypothecium pale yellowish; apothecia small, 0.4 - 0.8 mm across; disk pale flesh-colored to reddish brown. . . . . . . . . . 8, effuse It appears that 8gcidig_effusa sensu Fink and Erichsen is probably synonmous with Vainio's 8, intermedia and intermedia is in all likelihood merely a form of 8, albescens. Since I have not yet seen any authentic material of 8, albescens, Vsinio's interpretation is followed at this time. Unfortunately, the epithet intermedia on the species level (Arnold, 1871) 7 Based on vainio (1922) which in turn is based on a single specimen. 223 is preempted by Bacidia intermedia Hampe in Mass. (1861) and therefore is invalid. Until further studies are done to determine the correct name for this taxon, however, 8, intermedia (Hepp in Stizenb.) Arn. will be used. Distribution - Massachusetts, Iowa, California (Fink, 1935, sub 8, effusa); Europe; Asia (Vainio, 1928, sub 8, intermedia). Bacidia inundata (Fr.) Korb. Syst. Lich. Germ. 187. 1855. Biatora inundata Fr. Kgl. Vet. Akad. Nya Handl. 270. 1822. Material seen - SUFFOLK COUNTY: Brodo 761 (67), 8281 (54). This species is usually found on siliceous rocks in or near streams or brooks (Hale, 1950; Thomson, 1951), but the Long Island specimens were collected in comparatively dry habitats on concrete. One (no. 188) was growing on a concrete foundation within a few feet of a swampy brook, and the other was collected in a shaded oak woods, on an old concrete foundation. However, Sandstede (1913) reported 8, inundata from brick walls and Fink (1902) listed the species from limestone bluffs in Minnesota. Tuckerman (1888) stated that 8, inundata is found "on various rocks, especially such as contain lime; as also on brick;..." Distribution - Connecticut, Michigan, Minnesota, Indiana, Black Hills; East of Rocky Mountains (Fink, 1935): Temperate element, East Temperate subelement; Europe. Bacidia schweinitgii (Tuck. in W. Darl.) Schneid. Guide Study Lich. 110. 1898. Biatora schweinitzii Tuck. in W. Darl. Fl. Cestr. ed. 3. 447. 1853. Material seen - SUFFOLK COUNTY: 8£ggg_8888_(102), 2157 (102); 2802 (102). This species is unique among the Bacidiae in having Trentepohlig_as a phycobiont rather than Trebouxia. Lamb (1954) discusses this fact and some other aspects of the history of the species. 224 Bacidia schweinitzii was found in only one locality, as a member of the 522$ rubrum - bog community. It was also found in southern New Jersey (Burlington County, Atsion, 85282,8888) on a roadside oak close to a hog. Distribution - Nova Scotia, Maine, Connecticut, Tennessee, North Carolina, Oklahoma, Michigan, Indiana, Minnesota; eastern United States (Fink, 1935): Temperate element, East Temperate subelement; endemic. Bacidia cfr. trisepta (Naeg. in M511. Arg.) Zahlbr. Lecidea trisepta Naeg. in Mull. Arg. Mem. Soc. Phys. Hist. Nat. GenEve 16: 403. 1862. Material seen - SUFFOLK COUNTY: 85282 888Z_(44). 8gcidia trisepta has usually been reported to have ascocarps darker than that seen on the Long Island specimen, and, except for f. saxicola (Kgrb.) Lettau is mainly known from lignum and bark. The ascocarps appeared to be ascolocular in the Long Island specimen. Since I have not examined the type and the Long Island material was questionable, I will not transfer the species into Micarea where it might very well belong. The specimen cited was collected on a partially shaded granitic rock. Distribution - Massachusetts (Fink, 1935); Black Hills; Europe. Bacidia umbrina (Ach.) Bausch, Verh. Nat. Ver. Carls. 4: 103. 1869. Lecidea umbrina Ach. Lich. Univ. 183. 1810. Material seen - SUFFOLK COUNTY: 85282.8188 (111). The distinctive twisted and curved spores of this species easily separate it from all other Bacidiae on Long Island. Accurate measurements of the spore length were difficult due to the strong curvature of the spores, and the values appear to be somewhat lower than those reported by Hillman & Grummann (1957) or Erichsen (1957) (15 - 20 u rather than 17 - 40 u in length). The ecology of the specimen found on Long Island was extremely unusual for the species. It was found in the hygrohaline stratum on a granite boulder — 225 above the littoral zone but well within the storm tide level and certainly exposed to salt spray in windy weather. Growing alongside the specimen was Acarospora fuscata (see page 74). Distribution - Maine, Connecticut, Tennessee, Minnesota; northern United States (Fink, 1935): Temperate element, North Temperate subelement; Europe. RHIZOCARPON Rhizocarpon cinereovirens (Mull. Arg.) Vain. Acta Soc. Faun. F1. Fenn. 53. (1): 280. 1922. Patellaria cinereovirens M511. Arg. Flora 51: 49. 1868. Material seen - SUFFOLK COUNTY: 23222.2l12 (99), 8888 (119), 8822 (112). The very lightly tinted or hyaline one-septate spores of this species give it the appearance of a saxicolous Catillaria or a light spored Buellia (especially 8, stigmaea). However, gelatinous episporic sheaths are usually conspicuous indicating its true position. Runemark (1956) identified both norstictic and stictic acids from 8, cinereovirens by chromatographic analysis. The presence of norstictic acid in the medulla, an unusual feature among the Catocarpons, was detected in two of the Long Island specimens. The third specimen (Brodo 3899) was KOH + yellow; chromatography showed the presence of stictic acid, but not norstictic acid. Unfortunately, the Long Island material was too scanty to enable a more thorough chemical analysis. However, the presence of stictic and norstictic acids together is by no means uncommon (cfr. Parmelia conspersa, 8, hypotropa, etc.). Distribution - Black Hills; Minnesota (Fink, 1935); Europe. 226 Rhizocarpon gggggg.(F13rke in Flat.) Arn. Flora 54: 149. 1871. Lecideg,petr§gg_var. fuscoatra f. grandis Flgrke in Flat. Flora 11: 690. 1828. Material seen - SUFFOLK COUNTY: 88282 8882 (76). 88289carpon grande was discussed at length by Degelius (1940, 1941). Degelius (1940) mentioned the KOH + yellow to testaceous reaction of the medulla as well as the C + red reaction. The substances responsible for these reactions were identified by Runemark (1956) as stictic and gyrophoric acids. Stictic acid was found in the Long Island specimen (paper chromatography) and the C + red reaction indicates that gyrophoric acid is probably present as well. The specimen was found on an exposed granite boulder. Distribution - Maine, Tennessee, Michigan, Minnesota, Idaho, Black Hills, Hashington, Saskatchewan, Manitoba, Baffin Island: Arctic-boreaJ.element; circumboreal. Rhizocaggon intermedium.Degel. Ark. Bot. 30A (3): 43. 1941. Material seen - SUFFOLK COUNTY: 8£ggg_8298,(ll4), 28825’(108), 8218 (119). The Long Island specimen agreed perfectly with the type material (US). The type specimen contained stictic acid (by chromatography) as did the Long Island material (except one poorly developed specimen). I also collected the species on Cape Cod (Massachusetts) (85282_82§1, 828;, 52218). Distribution - Tennessee, Massachusetts (see above); endemic. Rhisocarpon obgcurgtum (Ach.) Mass. Ricerch. Anton. Lich. 103. 1852. Lecidea petraea var. abscurata Ach. Lich. Univ. 156. 1810. ' Material seen - SUFFOLK COUNTY: Imshaug 28822,(52), 85282.288,(53), 8128 (126). 12.6.7. (91). 3.3.9. (44). £1.12; (In). 2.7119. (111). 9218; (128). as; (119), 22g (112); Orient, 88582 1428, l by 1933 (Latham); Quogue, _L£_t_l_1_a_g 282548 (Latham); Shinnecock, Latham 27288, 8 May 1945 (Latham). — 227 This species is apparently extremely variable with many forms having been described for it (Erichsen [1951] included ten). 0f the many forms, f. reductum (T. Fr.) Eitn. seems to be most common on Long Island. This form is distinguished by a “more granulose thallus, smaller apothecia with thinner and disappearing margin, indistinctly papillated disk and submurale (not murale) spores” (Degelius, 1940). All Long Island specimens lacked any clearly positive chemical tests, although Runemark (1956) reported both stictic and gyrophoric acids from a specimen which he tested. Rhizocarpon orphninum (Vain.) Zahlbr. is very similar to 8, obscuratum but differs in having a KOH + violet or magenta reaction in the exciple and epithecium (Laundon, 1960). 88188garpog_obscurgggg_is common on pebbles and small stones and is often associated with Lecidea erratica. Distribution - Maine, Minnesota (Fink, 1935); Nova Scotia, Connecticut, Tennessee, Saskatchewan; Greenland (Lynge, 1940); Europe; Asia (Lynge, 1918). Rhizocaggon plicatile (Leight.) A. L. Sm. Monogr. Brit. Lich. 2: 197. 1911. Lecidea plicatilis Leight. Ann. Mag. Nat. Hist. IV. 4: 201. 1869. Material seen - SUFFOLK COUNTY: §£2§2.£§§§ (44), 8818 (128). Rhizocagpon plicatile was found on well illuminated or partially shaded boulders. I have collected specimens from.the Adirondack Mountains of New York. Distribution - Nova Scotia, Maine, northern New York (see above), North Carolina; Europe. STEREOCAULACEAE PYCNOTHELIA Pycnothelia papillaria (Ehrh.) Duf. Ann. Gen. Sci. Phys. Brux. 8: 5. 1817. Lichen pgpillaria Ehrh.Phytophyl. no. 100. 1780. 228 Material seen - NASSAU COUNTY: Brodo 3345 (8); Plain Edge, 8. Cain 371, 818, 1936, Andropoggggtum Hemsteadi (NY). SUFFOLK COUNTY: Brodo 59-177 (1008), 1111 (101), 1882 (103), 1888 (88), 1188 (126), 1188 (126), 1288 (91), 8818 (51), 8888 (49), 8888_(l7), 881|(55); 16 specimens collected by Latham (Latham); Orient, 88888, August 1877 (PH); Orient, 888889 ll: V. 1914 (EH); Orient Point, 888888, 1927 (NY); Montauk Point, R. H. Torrey, 1933 (NY); Selden, S. Cain §£§:.§§2: 888, 1936 (NY); Coram, R. H. Torrgy, 1936 (NY); Calverton, R. H. Torrgy, 1936 (NY); East of Calverton, R. H. Torrey, 1936 (NY); Route 112, north of Coram, R. H. Torrey, 1936 (NY). The important characters which separate Pycnothelia from Cladonia, e.g. pseudopodetia rather than podetia (see Lamb, 1951), and septate spores rather than nonseptate spores, have for some reason have ignored by most recent workers in the recent past with the notable exception of Watson (1953) and Mattick (1938). However, even Mattick (1940) later chose to regard chnothelia as part of Cladonia "for practical reasons." Pycnothelia pagillaria seems to be narrowly restricted to well illuminated localities on eroding sandy loam (see page 66). Distribution - Temperate element, East Temperate subelement, EurOpe (map: Sandstede, 1932). STEREOCAULON Stereocaulon saxatile Magn. Cgteb. Kgl. Vet. Samh. Handl. IV. 30: 41. 1926. Material seen - SUFFOLK COUNTY: 85888_8888 (76). Lobaric acid and atranorine were demonstrated by recrystallization in CAM and GAoT, respectively, in the Long Island and Cape Cod specimens. These chemical constituents were reported for this species by Lamb (1951) and Ramaut (1962). 229 The epithet evolutoides was published as a variety of 8. paschale by Magnusson in 1926 and was first used on the species level by Frey in 1932. It is necessary, therefore, to refer to this species as 8, saxatile, although most recent authors treat saxatile as a variety of evolutoides. A specimen of this species in much better condition than the Long Island material was found on Cape Cod (East Dennis, Brodo 4467). Both specimens were growing on granite boulders, the former in partial shade and the latter in full sun. Distribution - Nova Scotia, Massachusetts, Ontario, Saskatchewan: Temperate element, North Temperate subelement (?) (see Ahti, 1964); Europe. Listed as an “amphiatlantic, boreal” species by Lamb (1951). BAEOMYCETACEAE BAEOMYCES Baeomyces EgggggiPers. Neue Annal. Bot. 1: 19. 1794. Material seen - NASSAU COUNTY: Brodo 59:118_(12), 8888 (5). SUFFOLK COUNTY: Imshaug 8888Q.(52), 88888 (52), 88888 (72); §E222.§2:112.(54): 888_ (55), 888 (55), 218 (63), 1818.(31), 1888 (100A), 1888 (88), 1288 (91), 8QQ8_(51), 8281 (26), 8881_(128), 8888_(76); 13 specimens collected by Latham (Latham); Wildwood State Park, S. Smith 18882, 17 October 1952 (NYS). This species is usually found on eroding sandy loam especially on the moraines (figure 57), and is often associated with Pycnothel18DEapillaria. Distribution - Nova Scotia, Maine, Massachusetts, Connecticut, Tennessee; Appalachian-Great Lakes distribution (Hale, 1961a): Temperate element, Appalachian subelement, Appalachian-Great Lakes subelement (?); Europe, Asia (circumboreal: Sandstede, 1932). 230 CLADONIACEAE CLADONIA Section Clausae Kgrb. Subsection Cocciferae Del. Series Subglaucescentes Vain. Cladonia floerkeana (Fr.) FlSrke, Clad. Comm. 99. 1828. Cenomyce floerkeana Fr. Lich. Suec. Exs. 82. 1824. Material seen - SUFFOLK COUNTY: EEQQQHEQLLQL (83), 8218 (38), 8228 (17), 3426 (134); Southold, Latham 7573 (+ 7581, + 7588, + 7590), 3 January 1934 (Latham). This species is found on the ground in open sandy or grassy fields. Distribution - Vermont8, Massachusetts, Connecticut, New Jersey, Tennessee, Michigan: Temperate element, East Temperate subelement (?); Europe; Asia. Cladonia bacillaris (Ach.) Nyl. Bot. Sallsk. Faun. Fl. Fenn. F3rh. 8: 179. 1866. Baeomyces bacillaris Ach. Meth. Lich. 329. 1803. Material seen - KINGS COUNTY: New Lots, G. B. Brainerd, 1860's? (BKL 031984). NASSAU COUNTY: Brodo 550A (12),1888.(9), 1828 (14). SUFFOLK COUNTY: 112 specimens collected by Imshaug and/or Brodo; 39 specimens collected by Latham (Latham); Orient, 222222.392: 10 May 1914 (PH); Barling Hollow (: Baiting Hollow7), R1181_Torrey, 1934 (NY); Holtsville, R. H. Torrey, 1936 (NY); Southold, R. H. Torrey, 1937 (NY); Hyendanch Club Game Reserve south of Smithtown, R. H. Torrey, 1937 (NY); E. of Greenport, S. Smith 11888, 13 March 1955 (NYS). 8 In addition to those locality references given on page 121, the following references pertain to Cladonia: Vermont (Evans, 1947), Connecticut (Evans, 1930, 1944),N¢w Jersey(Evans 193,5)Tennessee (Mozingo, 1961), Michigan (Evans, un- published key to the Cladoniae of Michigan). References to presence in Asia are based on Asahina (1950) unless otherwise stated. 231 Caldonia bacillaris is one of the commonest lichens on Long Island. The species is very variable having numerous sterile and fertile forms. Red apothecia are present on approximately 50% of the specimens and appear either as conspicuous hemispherical terminal caps or mere dots of red at the podetial summits. The podetia either taper very gradually to a point, are almost entirely uniform in diameter, or are distinctly clavate. The species is found on a variety of substrates including soil, tree bases, and rotten wood, but it is found most frequently on wood. Distribution - Vermont, Massachusetts, Connecticut, New Jersey, Michigan, Indiana, Minnesota, Oklahoma, Arizona, Black Hills, washington, Alaska, Saskatchewan, Manitoba, Ontario: Temperate element, North Temperate subelement; Europe, Asia. Cladonia mag11enta Hoffm. Deutschl. Fl. 2: 126. 1796. Material seen - SUFFOLK COUNTY: 16 specimens collected by Imshaug and/or Brodo; Montauk, Hither beach, 88888!_88881, 88888, 28 October 1945 (Latham); Amagansett, Eghajm 25991, 11 birch 1947 (Latham); Greenport, m 81812, 30 April 1950 (Latham); Flanders, £££E22;3£ZZ§2 8 April 1946 (Latham); Riverhead, 1.1.5.9192 88881, 6 February 1953 (Latham); East Marion, 1312888, 1 September 1947 (Latham); Riverhead, 188888, 16 May 1960 (Latham); Orient, 1.88118! 818, 23 my 1914 (PH); Orient, 2.9921! 882, 10 May 1914 (PH). C1adonia macilenta closely resembles 8, bacillaris and they are best separated by their chemistry: 8, macilenta has thamnolic acid and 8, bacillaris does not. Although 8, macilenta, like 8, bacillaris, is found on many different substrates, it is usually found on sandy soil. Distribution - Vemmont, Massachusetts, Connecticut, New Jersey, Tennessee, North Carolina, Michigan, Ontario, Minnesota, Black Hills, washington, coastal Alaska: Temperate element, North Temperate subelement (?); Europe; Asia. 232 Cladonia vulcanica Zoll. Natur - et Geneeskundig Arch. Nearl. Indie 1: 396. 1847. Material seen - SUFFOLK COUNTY: Imshaug 88881_(86), 88888 (86); 88888 8188_(102), 8188_(102); Northwest, Latham 27458, 12 April 1948 (Latham); Flanders, 141919.39. 88188, 8 April 1946 (Latham); Riverhead, Latham 32871, 18 April 1955 (Latham); North Sea, LEEEEE.323172 35349, 26 March 1954 (Latham); Riverhead, 888885, 2 May 1957 (Latham); Riverhead, EEEEEEp 16 May 1960 (Latham). The presence of thamnolic acid in 8, vulcanica distinguishes this species from.the very similar 8, 818y88, Both species are found on rotting logs in bogs. It is interesting that 8, vulcanica was found to be abundant in the white cedar bogs of Cape God but was never collected in my studies of similar bogs in southern New Jersey. Exactly the reverse was true of 8, 818y88, Distribution - South America, and from New Jersey to Florida (Evans, 1952): Tropical element, Coastal Plain subelement; Asia. Cladonia didygg (F‘s) Vain. Acts Soc. Faun. F1. Fenn. 4: 137. 1887. Sgyphgpgorug didyggs Fee, Essai Crypt. Ecorc. Off. 98 & 101, pl. 3, f. 13. 1824. Material seen - SUFFOLK COUNTY: §£2§2.ZLQ§§.(85): 8188.(102); Montauk, Hither Beach, EE£§§!:§591§J 28 October 1945 (Latham); Riverhead, LEEEE! 29580, 7 August 1950 (Latham); Riverhead, 888882_(88888)?, 16 May 1960 (Latham); Northwest, 188888 88488, 10 April 1947 (Latham). A discussion of some aspects of the ecology and taxonomy of this species can be found with the comments on 8, vulcanica. Distribution - Connecticut to Florida along the coast: Tropical element, Coastal Plain subelement; much of South America, Africa, Hawaii, Ceylon, Japan (map:Sandstede, 1932), but not listed from Japan by Asahina (1950). 233 888§onia incrassata Flarke, Clad. Comm. 21. 1828. Material seen - NASSAU COUNTY: 88888 59-113 (12), 888,(l3),_888 (11), 8818 (10). SUFFOLK COUNTY: 29 specimens collected by Imshaug and/or Brodo; 39 specimens collected by Latham (Latham). This species is narrowly restricted to rotting wood and to pine bases. As in Cladonia cristatella, if podetia are produced, they are always capped by large red apothecia. Distribution - Along the coast from Nova Scotia to Florida (Evans, 1952): Temperate element, Coastal Plain subelement; Europe; Asia. Cladonia cristatella Tuck. Am. Jour. Sci. 25: 428. 1858. Material seen - KINGS COUNTY: Gowanus, G. B. Brainerd, (1866?), on ground (BKL). NASSAU COUNTY: §£2§2.§2§.(15)» 888 (12), 1888_(15), 1888 (14), 8128 (6), 8881 (8), 8828,(4). SUFFOLK COUNTY: 93 specimens collected by Imshaug and/or Brodo; 61 specimens collected by Latham (Latham); Orient Point, £2£E§E: 6 November 1911 (NYS); near Orient, 888888_11, 1914 (FH); near Orient, 153318231: 1914 (FH); ?, M18; 1914 (FH); Orient, 13.9.1312! 1_91, 20 my 1914 (FH); East of Calverton, R. H. Torrgy, 1936 (NY); Holtsville, 81_8, Iggggy, 1937 (NY); Pikes Beach, Westhampton, R. H. Torrey, 1936 (NY); Selden, R. H. Torrey, 1936 (NY); Selden, S. Cain 882, 1936 (NY); 2.3 miles SW of Riverhead, S. Smith 11888, 11881, 11882, 14 August 1952 (NYS). Cladonia cristatella is common and widespread on Long Island, occurring on a variety of substrates in a multitude of forms (see page 119). Distribution - Eastern United States (Sandstede, 1939): Temperate element, East Temperate subelement; endemic. Cladonia deformis (L.) Hoffm. Deutschl. F1. 2: 120. 1796. Lichen deformis L. Sp. P1. 1152. 1753. Material seen - SUFFOLK COUNTY: Montauk Point, R. H. Torrey, 1933 (NY). 234 This species, found only once on Long Island, is very similar to 8, pleurota differing in having farinose soredia, and podetial cups which are often lacerate and with involute margins. Distribution - Vermont, Massachusetts, Connecticut, Michigan, Ontario, Minnesota, Black Hills, Washington, Alaska, Saskatchewan, Manitoba, Canadian East Arctic: Arctic-boreal element; circumboreal. Cladonia Eleurota (Florke) Schaer. Enum. Crit. Lich. Eur. 186. 1850.. Capitalaria pieurota-Florke,~Mag. Ges. naturf. Freunde, Berlin 2: 218. 1808. ’ Material seen - NASSAU COUNTY: Brodo 3344 (8); Massapequa, S. Gain 88, 20 June 1935 (NY). SUFFOLK COUNTY: 20 specimens collected by Imshaug and/or Brodo; Riverhead, 52.112! 1181, 1 May 1937 (Latham). Sterile podetia of 8, glggyg;§_bear many resemblances to sterile 8, chloroghaea and the two are often found together on various types of soil. The yellow color of 8, Elggyggg_(due to usnic acid) distinguishes the two in the field. In addition, Long Island material of 8, chloroghaea almost always can be shown to contain grayanic acid which is absent in 8, Bleurota. Its similarity to 8, deformis has already been mentioned (see above).~ Cladonia Eleurota grows well on eroded sandy loam as well as on mossy soil and so has a broad distribution over both moraines. It is also occasion- ally found on tree bases. Distribution - Nova Scotia, Vermont, Massachusetts, Connecticut, New Jersey, Mflchigan, Ontario, Black Hills, Alaska, Manitoba, Canadian East Arctic: Arctic-boreal element; Europe; Asia; "hemiarctic" (Ahti, 1964). Subsection Ochroleucae Fr. Cladonia carneola (Fr.) Fr. Lich. Eur. 233. 1831. Cenomyce carneola Fr. Sched. Crit. 4: 23. 1825. Material seen - SUFFOLK COUNTY: Brodo 2693 (110); Montauk, R. H. Torrey, 1933 (NY). 235 The presence of barbatic acid and farinose rather than granulose soredia are usually sufficient to separate sterile specimens of this rare species from the more common 8, Eleurota. Fertile material is easily distinguished since the apothecia are brown rather than red. Cladonia carneola is found on well illuminated eroding soil. Distribution - Black Hills, Washington, coastal Alaska, British Columbia, Saskatchewan, Manitoba: Arctic-boreal element; circumboreal (Sandstede, 1939; Ahti, 1964). Cladonia piedmontensis Merr. Bryologist 27: 22. 1924. Material seen - NASSAU COUNTY: §£242.§§§l (8). SUFFOLK COUNTY: Imehaug 88811 (116), Brodo 2821 (115); Montauk Point, R. H. Torrey, 1933, (NY). Cladonia suhggraggnea Nyl. (p.p.) is listed as a synonym of 8, piedmontensis by Fink (1935). It is also, in part, a synonym of 8, cristatella f. ochrocarpia Tuck. (Evans, 1930; Fink, 1935). Until the type is examined and the true identity of.§, substraminea is determined, the name 8, piedmontensis will be used. Distribution - Massachusetts and Connecticut southward to Alabama and Mexico (Evans, 1930): Temperate element, Appalachian subelement (?), Appalachian unit (?); endemic. Subsection Foliosae Cladonia robbinsii Evans, Trans. Conn. Acad. Arts Sci. 35: 611. 1944. Material seen - SUFFOLK COUNTY: Southold, LE£E§!;Z§§Q.( +-1881) 20 January 1934 (Latham); Orient, 191888 8_4_6_7_, 5 May 1939 (Latham); Orient, west Long Beach, 8581182 88888, 88382, 7 December 1944 (Latham); Shinnecock Hills, m 88288, 8 my 1946 (Latham). . This species bears many similarities to closely related 8, stregsilis but differs from the latter in color (dark yellowish-green as opposed to olive 236 green) and in chemistry (usnic and barbatic acids present rather than baeomycic acid and strepsilin). Distribution - Connecticut (Evans, 1944), Tennessee, Black Hills: endemic. Cladonia stregsilis (Ach.) Vain. Act. Soc. Faun. Fl. Fenn. 10: 403. 1894. Bagggyces strepg1118_Ach. Math. Lich. Suppl. 52. 1803. Material seen - NASSAU COUNTY: 88888_882 (16), 8881_(5), 8888 (8), 8818 (10). SUFFOLK COUNTY: 23 specimens collected by Imshaug and/or Brodo; 17 specimens collected by Latham (Latham); Orient Point, 1881198, 1927 (NY); Shinnecock Hills, R. H. Torrey, 1933 (NY); Rt. 112 north of Coram, 81_8. Iggggy, 1936 (NY). No other Cladonia on Long Island has strepsilin and the accompanying C‘+ green medullary reaction. This species is fairly common on waste soil and sandy roadbanks; it is occasionally found on mossy soil. Distribution - Eastern United States southward to Mexico (Sandstede, 1939): Temperate element, East Temperate subelement; Europe; Asia. Subsection Podostelides (wallr.) vain. Series Helogodium.(Ach.) vain. Cladonia cagitata (Michx.) Spreng. Syst. Veg., ed. 16, 4: 271. 1827. Helogodium cagitatum Michx. Fl. Bor. Am. 2: 329. 1803. Material seen - NASSAU COUNTY: Valley Stream, E. A. Warner, 17 November 1900 (BKL). SUFFOLK COUNTY: Igggaug 88888 (52), EEQSQ.22:£§Z.(54)a 82:188 (54). 221m (68). .91_s, (39). up; <65). 19¢; (53). 2Q (59). _1_:_ss_9 (65). 1811 (65), 3121 (99). 8181,(111). 21$; (111). 8888_(23), 8882 (128); 25 specimens collected by Latham (Latham); Orient, 112.2113! 128, 4 July 1914 (FH); Greenport, EEEEEEHE: 1914 (FH); Coram, R. H. Torrey, 1936 (NY). Fink (1935) listed this species under the name Cladonia mitrula Tuck. in W. Darl. 237 Cladonia cagitata is most commonly found on tree bases in well lighted oak forests but sometimes is found on sandy soil. Distribution - Eastern United States and Cuba (map: Sandstede, 1938): Temperate element, East Temperate subelement; Europe (Poelt, 1963); Asia. 81889nia car19§a (Ach.) Spreng. Syst. Veg. ed. 16, 4: 272. 1827. Lichen cariosus Ach. Lich. Suec. Prodr. 198. 1798. Material seen - SUFFOLK COUNTY: Montauk Point, R. H. Torrey, 1933 (NY). Distribution - Vermont, Connecticut, Tennessee, Michigan, Ontario, Indiana, Minnesota, Black Hills, Arizona, washington, Alaska, Saskatchewan, Manitoba, Baffin Island: Arctic-boreal element; circumboreal. Cladonia subcariosa Nyl. Flora 59: 560. 1876. Material seen - QUEENS COUNTY: §£2§2.§ZQ.(3)- NASSAU COUNTY: 88888_8128 (6), 8888 (8). SUFFOLK COUNTY: Imshaug 25619 (116), 88888 (116), 88888 (116), 229;; (116), 2.5.632 (116); 232.42 222.5. (53). 2.2.1.28. (82). .1191 (127). .227; (38), 8881 (20); 11 specimens collected by Latham.(Latham); Southold, 81_88 Torre , 1933 (NY); Montauk Point, R. H. Torrey, 1933 (NY); Montauk Point, 1:. n. Torrey, 1933 (NY); 2, Latham (81?), 1914 (PH). One can consider 8, subcariosa the central element of a group of closely related taxa called the Cladonia subcgg1gs4a group. Members of this group are morphologically almost indistinguishable, but show some differences in distribution and chemistry. Of this group, 8, subcariosa contains norstictic acid, 8, clavulifera contains fumarprotocetraric acid, 8, 888y18_contains psoromic acid, and 8, golycargia contains atranorine. Cladonia golycaggia, which is not found on Long Island, is considered synonymous with 8, clavulifera by Mattick (1940). In this paper, the first three species will be recognized although there is considerable question as to whether they are distinct (see Mozingo, 1961). In view of the fact that these species differ little in 238 their morphology, and their chemical components are closely "related" (all being Porsellic acid depsides or depsidones with a substantial history of chemical shifting between closely related taxa), it might be better to consider them in an apprOpriate infraspecific rank. Pending further study of the morphology, chemistry, and phytogeography of members of the 8, subcariosa complex, the various "microspecies" will be recognized. Cladonia subcariosa is found in dry sandy or grassy fields. Distribution - Eastern United States (map: Sandstede, 1938): Temperate element, East Temperate subelement; Europe; Asia. Cladonia clavulifera vain. in Robb. Rhodora 26: 145. 1924. Material seen - NASSAU COUNTY: Brodo 2529 (5), 8828.(4), 8888_(10). SUFFOLK COUNTY: 18 specimens collected by Imshaug and/or Brodo; Southold, 1._a_t_l'1a_n_|_ 129_5 ( + 8888), 11 February 1938 (Latham); Orient, Long Beach, 8am 88888, 88821, 88822, 7 December 1944 (Latham); Napeague, Latham 22988_(= 88288), 20 February 1941 (Latham); Amagansett, LEE!§E.Z§221: 11 March 1947 (Latham); Noyack, LEEEE!.Z§§$§: 9 March 1947 (Latham); Bridgehampton, Latham 27050, 14 September 1947 (Latham); Bridgehampton, 888888 81888, 14 September 1947 (Latham); North Sea, L2£E£E:Z§l§£p 16 May 1955 (Latham); Orient Point, 1.3.9.3.! 8, 9 January 1911 (NYS); near Orient, m 18 (FH); (locality unknown) 888888, 1914 (FH); Montauk Point, R. H. Torrey, 1933 (NY); Southold, R. H. Torrey, 1936 (NY); Pike's Beach, westhampton, R. H. Torrey, 1936 (NY); Selden, S. Cain 888, 881, 30 June 1936 (NY). This species is usually found on exposed sandy ground. Distribution - Maine, Massachusetts, Connecticut, New Jersey, Maryland, washington, D. C., Virginia (Sandstede, 1939); Vermont, Tennessee, Oklahoma: Temperate element, East Temperate subelement (?); Asia. 239 Cladonia brevis Sanst. Abhandl. Naturv. Ver. Bremen. 25: 192. 1922. Material seen - SUFFOLK COUNTY: Imshaug 82§Q§.(64); Brodo 8852 (69); Southold, ngggg_1882, 11 February 1938 (Latham); Riverhead, ESSE.(NYS)3 Montauk Point, R. H. Torrey, 1933 (NY); East of Calverton, R. H. Torrey, 1936 (NY); Airport near Westhampton, R. H. Torrey, 1936 (NY). Cladonia brevis, like the other members of the g, subcariosa group (see page 237 is found on dry sandy soil. Distribution - Maine, Massachusetts, Connecticut (Sandstede, 1938); Vermont, New Jersey, Tennessee, Manitoba: Temperate element, East Temperate subelement (?); Europe. Subsection Thallostelides Vain. Cladonia verticillata (Hoffm.) Schaer. Lich. Helv. Spic. 31. 1823. Cladonia pyxidata *'Q, vergioillata Hoffm. Deutschl. Fl. 2: 122. 1796. Material seen - KINGS COUNTY: New Lots, G. B. Brainerd, (1866?) (BKL 031990); New Lots, G. B. Brainerd, (1866?) (BKL). SUFFOLK COUNTY: 85282 22:222.(53); Greenport, Eggggg‘82322, 12 April 1945 (Latham); Northwest, Third station, Latham 27447, 27 April 1948 (Latham); Orient, EEEEEE;2§§§Z: 17 April 1950 (Latham); Sag Harbor, 825329, 15 September 1941 (Latham); ? near Orient, 9.9.3133 82 (PR). The separation of this species from closely related 8, calycantha is often very difficult. Such characters as smooth cup margins and gradually expanding cups usually attributed to Q, verticillata are not always evident. The ecology of the two species, however, seems to be different with Q, verticillata being found in open sandy or grassy fields, especially on neutral soils, and Q, calycantha being found mainly in boggy or acid sand localities usually under pines. The geographical distribution of the two species is basically different as well. 240 Distribution - Nova Scotia, Vermont, Massachusetts, Connecticut, New Jersey, Tennessee, Michigan, Ontario, Minnesota, Black Hills, Hashington, Alaska, Saskatchewan, Manitoba, Canadian East Arctic: Arctic-boreal element; circum~ boreal. Cladonia calycantha Nyl. Syn. Meth. Lich. 192. 1858. Material seen - SUFFOLK COUNTY: 8gphaug 25856 (60); Brodo 59-23 (83), _5_9_-_25_ (83). 22102 (68), 2229216». 1132 (78). .2_<_>_9_2_ (83), 1222, (87). £223 (49), 8228 (75), 8888 (66); 29 specimens collected by Latham (Latham); Napeague, Latham 26024, 11 March 1947 (US: Evans); Springs, Latham 26432, 17 April 1947 (US: Evans); Northwest, Latham 26391, 17 April 1947 (FH); Northwest Section 2, Latham 27480, 21 April 1948 (US: Evans); Airport near Hesthampton, R. H. Torrey, 1936 (NY); Pike's Beach, westhampton, R. H. Torrey, 1936 (NY); Sweezy Pond, 2.3 miles SW of Riverhead, S. Smith_8882§, 14 August 1952 (NYS). The relationship between this species and Q, verticillata has been discussed with the latter. Distribution - Newfoundland to Florida, South America, Australia (Sandstede, 1938): Tropical element, Coastal Plain subelement; Europe (Poelt, 1963); Asia. Cladonia mateocygtha Robb. Rhodora 27: 50. 1925. Material seen - NASSAU COUNTY: 85282 252 (16). SUFFOLK COUNTY: 7 LEEEEE. 85 1914 (FH); Montauk Point, R. H. Torrey, 1933 (NY); between Commack and Kings Park, R. H. Torrey, 1937 (NY). This species, very rare on Long Island, shows considerable morphological variability. Its smooth or cracked, completely corticate podetial surface together with its irregularly proliferating cups, giving rise to contorted branches from both central and marginal areas, distinguish Q, meteocyatha from other nonsorediate species. 241 This species is found on exposed soil. Distribution - Massachusetts, Connecticut, Washington D. C., West Virginia, New Mexico (Sandstede, 1939); Vermont, New Jersey, Tennessee, Michigan: Temperate element, Appalachian subelement, Appalachian - Great Lakes - Rocky Mountain unit (?); endemic. Cladonia pyxidata (L.) Hoffm. Deutschl. F1. 2: 121. 1796. .EiSEEE pyxidatus L. Sp. Pl. 2: 1151. 1753. Material seen - KINGS COUNTY: Gowanus, G. B. Brainerd (BKL 031992). SUFFOLK COUNTY: Imshaug 25628 (116), 88888 (60); Brodo 2838 (115); 10 specimens collected by Latham.(Latham); Montauk Point, R. H. Torrey, 1933 (NY). gladonia pyxidata is usually found on the ground in dry, sandy localities. Distribution - Vermont, Massachusetts, Connecticut, New Jersey, Tennessee, Michigan, Ontario, Minnesota, Wisconsin, Black Hills, Arizona, Idaho, Washington, Alaska, British Columbia, Saskatchewan, Manitoba, Canadian East Arctic, Baffin Island: Arctic-boreal element; circumboreal. Cladonia chlorgphaea (Flsrke in Sommerf.) Spreng. Syst. Veg. ed. 16. 4: 273. 1827. Cenomyce chlorophaea Flgrke in Sommerf. Suppl. F1. Lapp. 130. 1826. Material seen - COUNTY UNKNOWN: Fresh Pond, 88888, 1890 (BKL 031986). QUEENS COUNTY: §£2§2.§l2 (3). NASSAU COUNTY: 85882_881 (16), 888 (12), 888 (13), 8888 (15), 8888 (8), 8888 (10); Oyster Bay, L. P. 1e 7, September 1889 (NY); Valley Stream, 285885, 17 November 1900 (BKL). SUFFOLK COUNTY: 109 specimens collected by Imshaug and/or Brodo; 70 specimens collected by Latham (Latham); (locality unknown) 8.58892, 1914 (FH); Orient, LEM 1.88, 1914 (FH); (locality unknown) 2.9.1193» 1914 (FH); Orient, 2.91%! 888, 23 May 1914 (FH); Shinnecock Hills, R. H. Torrey, 1933 (NY); Holtsville, R. H. Torrey, 1936 (NY); Pikes Beach, Westhampton, R. H. Torrey, 1936 (NY) (PD + red); 242 Pikes Beach, Westhampton, R. H. Torggy, 1936 (NY) (PD -); Suffolk County Airport near Westhampton, R. H. Torrey, 1936 (NY); Selden, S. Cain 345, 30 June 1936 (NY); Wyandanch Club Game Preserve south, R. H. Torrey, 1937 (NY); Horton's Beach, Southold, S. Smith 11896, 11897, 14 August 1952 (NYS); Wildwood State Park near Riverhead, S. Smith 12744, 17 October 1952. This species, one of the most abundant on Long Island, is extremely variable in morphology and ecology. Soredia range from almost farinose to granular and even appear corticate in some specimens; cups are either simple, goblet-shaped structures with smooth margins, or have many often large, marginal proliferations bearing large brown apothecia. Fumarprotocetraric acid, as determined by a PD + red reaction on the podetia, was demonstrated in about 60% of the specimens. The presence of grayanic acid was determined by the microscopic examination of acetone extracts of the podetia with supplementary recrystallization in GAW solution if necessary. It was found in all the Long Island specimens except one (85889 8889 [112]) in which cryptochlorophaeic acid was demonstrated. In addition, nine specimens of 8, chlorophaea from southern New Jersey and 12 from.Cape Cod were chemically examined. All of these also contained only grayanic acid except for one specimen (§£2§2.$§§Z from.Cape Cod) which showed neither grayanic nor cryptochlorophaeic nor merochlorophaeic acids. Fumarprotocetraric acid is regarded as an accessory substance in 8, chlorophaea by most modern workers (see Evans, 1944). However, the presence or absence of the other substances mentioned above have been used by Evans, Asahina, and others as a basis for recognizing four species: 9, gggyi'Merr. in Sandst. with grayanic acid; 8, cryptochlorophaea Asah. with cryptochlorophaeic acid; 8, merochlorophaea Asah. with merochlorophaeic acid; 8, chlorophaea sens. str. with none of these chemicals. Several of these chemical segregates seem to have some geographic restrictions (grayanic acid strain : eastern, inactive 243 strain 3 northern). The other strains are rather rare, and are geographically poorly defined. Until the full chemical story is known, at least in North America, it seems best to regard these segregates as chemical strains within 8, chlorophaea sens. lat. although with further study at least a few may prove to be more logically considered as subspecies, or perhaps even species. Cladonia chlorophaea is found on soil, lignum, or tree bases. Distribution - Nova Scotia, Maine, Vermont, Massachusetts, Connecticut, New Jersey, Tennessee, Alabama, Florida, Michigan, Ontario, Indiana, Wisconsin, Minnesota, Oklahoma, Black Hills, Arizona, Washington, Alaska, British Columbia, Manitoba, Canadian East Arctic, Baffin Island: Arctic-boreal element; circum- boreal. Cladonia fimbriata (L.) Fr. Lich. Eur. Ref. 222. 1831. 888888 fimbriatus L. Sp. P1. 1152. 1753. Material seen - SUFFOLK COUNTY: Riverhead, LEEEEE;Z§§§§: 1 May 1937 (Latham); Bridgehampton, Latham 27043, 14 September 1947 (Latham); North Sea, EEEE§!;§Z2§§’ 26 April 1954 (Latham); Riverhead, Latham 33318, 1 June 1923 (Latham); East Marion, Latham, 3 May 1914 (Latham). Cladonia fimbriata varies from a very narrow cupped condition very similar to 8, coniocraea, to a broader trumpet shaped condition resembling g, conista. However, its podetia always show a distinct, deep, though often very narrow cup and rarely are as subulate as those of 8, coniocraea in which the podetial cups are flat or very shallow. Cladonia fimbriata also has much smaller podetial and basal squamules than the latter. In addition, the podetia of 8, fimbriata never arise from the center of a primary squamule as do the podetia of 8, coniocraea. gladonia cgnista, with broad goblet-shaped cups has its soredia confined to the upper third of the podetium and the inner. surface of the cup and contains substance "H" whereas 8, fimbriata has narrower podetia covered with soredia and does not contain substance H. 244 Cladonia fimbriata is found on the ground and on tree bases. Distribution - Arctic-boreal element (Hale, 1954b; Thomson, 1953, 1955); circumboreal. Cladonia conista (Ach.) Robb. in Allen, Rhodora 32: 92. 1930. Cenomyce fimbriatap—8. conista Ach. Syn. Meth. Lich. 257. 1814. Material seen - KINGS COUNTY: Gowanus, G. B. Brainerd, 1866 (BKL). QUEENS COUNTY: Cypress Hills, flsleg. 1890 (BKL (B1991). SUFFOLK COUNTY: Imshaug 88888 (72), 88888 (132); Brodo 811 (908), 8888 (1008), 8188_(111), 8888 (128); 10 specimens collected by Latham (Latham). The presence of substance H in 8, conista easily separates it from similar species which lack it such as 8, fimbriata (see above) and 8, chlorophaea In addition, the latter usually has distinctly granular soredia covering the entire podetium. Cladonia conista grows on soil or tree bases. Distribution - Vermont, Massachusetts, Connecticut, New Jersey, Tennessee, Michigan, Black Hills: Temperate element, North Temperate subelement (?); Europe, Asia. Cladonia coniocraea (Flgrke) Spreng. em. Sandst. Syst. Veg. ed. 16. 4: 272. 1827. Sandstede, Abh. Naturw. ver. Bremen 21: 373. 1912. Cenomyce coniocraea Flgrke, Deutschl. Lich. 138. 1821. Material seen - QUEENS COUNTY: Brodo 526 (3). NASSAU COUNTY: 17 specimens collected by Brodo. SUFFOLK COUNTY: 83 specimens collected by Imshaug and/or Brodo; 47 specimens collected by Latham (Latham); Mbntauk Point, R. H. Torrey, 1933 (NY); Coram, R. H. Torrey, 1936 (NY); Greenport, Latham 30938a, 30 May 1952 (NYS). This species is one of the most common and variable of the Cladoniae. Cladonia coniocraea is usually said to have abruptly tapering podetia entirely 245 covered with farinose soredia except for a narrow basal zone. Long Island specimens, however, show every gradation from this "typical" form to a condition having almost entirely corticate podetia with patches of farinose or granular soredia scattered along their length. This latter form has generally been considered under the name 88§80nia ochrochlora Flc'irke. Evans (1935) discussed the difference between these two species. In general, five characters are fairly constant in 8, coniocraea: 1) the podetia are sorediate, 2) the podetia are usually short, stout, and abruptly tapering to a sharp point, 3) the podetia arise from the center or near center of a primary squamule, 4) the primary squamules are broad, and 5) the podetial cortex and the soredia usually have a yellowish or yellow-olive caste (not due to usnic acid). A great deal of variability can be seen in l) the extent of podetial cortex, 2) the type of soredia (although the granular sorediate condition is very rare), 3) the presence of soredia on the primary squamules (from abundant to essentially absent), 4) the presence of cupped podetia (see discussion under 8, fimbriata, page 243), 5) the lobing of the primary squamules (entire to crenate), and 6) the degree of branching (podetia are almost always simple, but rarely have one, or at most, two simple branches). Apothecia, which are rare in this species, are brown and irregular, occurring at the edges of poorly developed cups or trays. Cladonia coniocraea is usually found on mossy soil or tree bases and almost always in shaded situations. Its city-tolerance is discussed in Section IV. Distribution - Nova Scotia, Maine, Vermont, Massachusetts, Connecticut, New Jersey, Tennessee, Alabama, Ontario, Michigan, Wisconsin, Minnesota, Indiana, Oklahoma, Arizona, Washington, Alaska, British Columbia, Manitoba, Canadian East Arctic: Temperate element (?), North Temperate subelement; Europe, Asia. 246 Cladonia nemoxyna (Ach.) Arn. Lich. exs. no. 1495. 1890. Baeomyces radiatus (8,88moxygus Ach. Math. Lich. 342.1803. Material seen - SUFFOLK COUNTY: Brodo 2713 (111); Orient, Long Beach, 888888_82, 15 April 1914 (Latham); Montauk Point, R. H. Torrey, 1933 (NY); Southold, R. H. Torrey, 1936 (NY); 2.3 miles southwest of Riverhead, 88 88888_& Qagen, Smith 11853, 11854, 14 August 1952 (NYS). Cladonia nemoxyna may contain fumarprotocetraric acid; it always contains homosekikaic acid (Evans, 1944). All the Long Island specimens were PD + red and presumably contained fumarprotocetraric acid. The specimens of this species cited by Degelius (1940) from Maine also were PD + red. Specimens from Ontario (Ahti, 1964), the Great Lakes region, and the Black Hills are PD -. The presence of fumarprotocetraric acid, therefore, may prove to have geographic correlation as do many other chemical populations of lichens (Cladonia uncia888, 8, 888grophaea, Lecanora caesiorubella, etc.). Homosekikaic acid is very difficult to demonstrate apparently because. it occurs in very minute concentrations. The directions for its recrystalli~ zation from GAOT solution as given by Evans (1943) should be followed carefully. This species was found on eroding soil associated with 8, conista, 8, farinacaea, and 8, cristatella. Distribution - Vermont, Massachusetts, Connecticut, New Jersey, Tennessee, Ontario, Michigan, Black Hills, Washington: Temperate element, North Temperate subelement; Europe; Asia. Cladonia cylindrica (Evans) Evans, Rhodora 52: 116. 1950. Cladonia borbonica f. cylindrica Evans, Trans. Conn. Acad. Arts Sci. 30: 482. 1930. Material seen - NASSAU COUNTY: §£2§2.l§92.(15): 8888 (10). SUFFOLK COUNTY: 88888 8888 (65), 288(35-36), 8818 (27), 886_2_ (29), 8288 (133), 8888 (28), 8282 (112); Greenport, Gull Pond, EEEEEB.32§2§: 20 January 1951, dry soil in woods (Latham). 247 This lichen bears many similarities with 8, coniocraea. Both have more or less short, sorediate, usually sterile podetia, and both are PD + red. However, 8, cyl8ndrica has a clear gradation of coarse granules at the podetial base to farinose soredia at its tip and contains grayanic acid, whereas 8, con89craea is entirely covered with farinose soredia and lacks grayanic acid. 88ggonia cylindrica is found on tree bases, usually in shaded woods. Distribution - Vermont, Massachusetts, Connecticut, New Jersey, Michigan; West Virginia (Sandstede, 1939): Tropical element, Temperate-Appalachian subelement; Asia; circumtropic (Sandstede, 1939). 8888onia pityrea (F13rke) Fr. Nov. Sched. Crit. 21. 1826. Cap8§plaria pitygea F13rke, Ges. Naturf. Fr. Berlin mag. 2: 135. 1808. Material seen - QUEENS COUNTY: Brodo 523 (3). SUFFOLK COUNTY: Brodo 585' (92), 8188 (111), 8888 (119), 8888.(129); East Marion, EEEEEE.§; 3 May 1914 (Latham); Greenport, E££E§!.Z§&Z§: 12 April 1945 (Latham); Riverhead, Latham 888888, 1 May 1960 (Latham); Riverhead, 888888, 25 May 1960 (Latham); Greenport, 888118818888 (?) (MICH); (locality unknown), 8.8818818, 1914 (EH). Almost all of the Long Island specimens of 8, pitygea are identical with Connecticut material identified by Evans (in herb. FH) as 8, pitygea var. zwackii vain., and either form subacuta Vain. or form sggamulgfera Vain. The podetia were contorted and covered with coarse granules or granular soredia. In the type of form sguamulifera (Thaxter 88, Trinidad 1912-13, YFH], the podetia were densely squamulose and granular sorediate, not very contorted, and were not pellucid and dark in the decorticate areas. The podetial squamules were finely lobed and almost nonsorediate. In other words, the type of 8, sguamulifera does not seemto agree with Evans' identifications, and 8, sguamulifera sensu Evans is probably a kind of 8, subacuta (especially 248 since collections containing both forms in the same packet were common). The value of these infraspecific taxa is doubtful. Cladonia gitygea has been collected on various substrates including dry ground, tree bases, rocks, and wood. Distribution - Vermont, Connecticut, New Jersey, Tennessee, Florida, Michigan; South America, West Indies, East Indies (Sandstede, 1939): TrOpical element, Appalachian-Temperate subelement; Europe; Asia. 88ggonia simulata Robb. Rhodora 31: 105. 1929. Material seen - SUFFOLK COUNTY: Northwest, Latham 27200, 27 April 1947 ' (US: Evans). This species had been placed in the subsection Ochroleucae by Sandstede (1938) and Mattick (1940) on the basis of its resemblance to 8, piedmontensis. Evans (1952), following Robbins, pointed out that the similarity of the species to 8, piedmontensis is entirely superficial, and its chemistry (lack of usnic and presence of fumarprotocetraric acid) places it in the subsection Thallostelides, close to 8, pityrea. The specimen was growing on dry sand. Distribution - Massachusetts, North Carolina, Georgia, Florida (Evans, 1952): Temperate element, Coastal Plain subelement; endemdc. Section PERVIAE (Fr.) Matt. Subsection Chasmariae (Ach.) Flgrke Series Megaphyllae Cladonia agodocarpa Robb. Rhodora 27: 211. 1925. Material seen - SUFFOLK COUNTY: Riverhead, Latham 24794, 5 April 1946 (US: Evans). This species is the only non-podetiate Cladonia on Long Island containing 249 both atranorine and fumarprotocetraric acid. Latham's specimen was from a dry woods. Distribution - Northeastern states (Sandstede, 1939); Tennessee, Alabama: Temperate element, East Temperate subelement; endemic. Series Microphyl888_Vain. Cladonia caespiticia (Pers.) Flgrke, Clad. Comm. 8. 1828. Baeomyces caesp888cus Pers. Ann. d. Bot. 7: 155. 1794. Material seen - QUEENS COUNTY: m _5_g_1_ (3). NASSAU COUNTY: 13324.9. 1_312. (15), 8888 (5). SUFFOLK COUNTY: 8mshaug 25693 (72); Brodo 2426 (20), 8888 (107), 8888 (50), 8888 (57), 8288 (36); Orient, Latham.789, 3 March 1914 (Latham); Montauk, L££E§9;§l§2§: 16 May 1951 (Latham); Mattituck, LEEEQE. 88888, 7 June 1955 (Latham); East Marion, 888888, 3 May 1914 (Latham); Orient, LEEEEE.122L 15 May 1914 (FH); (locality unknown), 888888, 1914 (FH); Barling Hollow ( ' Baiting Hollow?), R. H. Torrey, 1934 (NY); Wyandanch Club Game Preserve, south of Smithtown, R. H. Torrey, 1937 (NY). This species is found on bare or mossy ground, often on charred ground, and less frequently on tree bases. It is almost always in shaded or partially shaded localities particularly in the black and red oak forests. Distribution - Eastern United States (Sandstede, 1938): Temperate element, East Temperate subelement; EurOpe; Asia. Cladonia paras888gg_(Hoffm.) Hoffm. Deutschl. Fl. 2: 127, 1795. 888888_parasiticus Hoffm. Enum. Lich. 39, tab. 8, fig. 5. 1784. Material seen - NASSAU COUNTY: §£2§2.l§92.(14): 8888 (14). SUFFOLK COUNTY: mag; (59). 1mg (21). 132.; (19). 122g (19). 21(1), (78). z_1_s_3_ (102). .2316. (22). £93; (71). 2.9.7.5. (43>. 222g. (43). 29s; (50). 212 (65). gm (62). 8288 (112); 12 specimens collected by Latham (Latham); Shinnecock Hills, R. H. Torrey, 1933 (NY). 250 Cladonia parasitica is usually described as having granular soredia. On many specimens which I have seen, however, the so-called soredia appear to be corticate and,therefore, are actually granules. The species has been called 8, delicata (Ehrh) Flgrke, by most authors. This species is almost entirely confined to decaying logs and wood of various origins, but especially coniferous trees. It is usually found in shaded situations. Distribution - Throughout eastern United States (Evans, 1930): Temperate element, East Temperate subelement5EurOpe; Asia. Cladonia santensis Tuck. Am. Jour. Sci. Arts 11. 25: 427. 1858. Material seen - SUFFOLK COUNTY: Lmshaug 88888 (86); Riverhead, Sweezy Pond, I__.£_1_t_:_h_a_8m_ 88888, 18 April 1955, (Latham); North Sea, 8.8881818 88282, 20 May 1954 (Latham); Riverhead, 888888, 1 May 1960 (Latham). This species can be recognized, even in the field, by its very pale, almost white color, and its contorted, bent, minutely squamulose podetia. Sterile material of 8, santensis sometimes closely resembles sterile 8, pargsitica which also contains thamnolic acid. The latter, however, has finely divided, "lacy,” granulate primary squamules whereas in 8, santensis the squamules are thicker and not granular or sorediate. Long Island is the northern limit of this species. It was found to be abundant in cedar bogs in southern New Jersey. Distribution - New Jersey to Florida (Evans, 1952): Temperate element, Coastal Plain subelement; endemic. Cladonia sguamosa (Scop.) Hoffm. Deutschl. Fl. 2: 125. 1796. Lichen sguamosus ScOp. Fl. Carn. ed. 2. 2: 368. 1772. Material seen - SUFFOLK COUNTY: Brodo 2164 (102), 2358 (42), 2569 (73), 2998 (17), 3001 (17), 8008 (17), 8888 (50); 10 specimens collected by 251 Latham (Latham). This species, which is uncommon on Long Island, is usually found on mossy ground, rocks, or wood in partially shaded black or red oak forests. Its relationship to 8, atlantica is discussed under the latter species. Distribution - Nova Scotia, Maine, Vermont, Massachusetts, Connecticut, New Jersey, Tennessee, Alabama, Ontario, Michigan, Minnesota, Indiana, Washington, coastal Alaska, Baffin Island; “arctic to southern temperate" (Ahti, 1964): Arctic-boreal element; circumboreal. Cladonia atlantica Evans, Trans. Conn. Acad. Arts Sci. 35: 573. 1944. Material seen - NASSAU COUNTY: Brodo 542 (12), 8888 (15), 8888.(10). SUFFOLK COUNTY: 59 specimens collected by Imshaug and/or Brodo; 66 specimens collected by Latham (Latham); West Suffolk Co. Airport near Westhampton, R. H. Torrey (NY); Baiting Hollow, R. H. Torrey, 1934 (NY); Holtsville, R. H. Torrey, 1936 (NY); Smithtown, R. H. Torrey, 1936 (NY); 2.3 miles SW of Riverhead, S. Smith 88888, 88888, 88888, 14 August 1952 (NYS); Horton's Beach, Southold, S. Smith 88828, 14 August 1952 (NYS); 1.3 miles W of Middle Island, S. Smith 88888, 12 March 1955 (NYS). This species, which is very common throughout the sandy parts of the island, is very variable in its morphology. Podetia devoid of squamules commonly are found as well as podetia entirely covered with small or large squamules. Apothecia seem to be more common on the squamulose forms. The main difference between 8, atlantica and 8, sguamosa (from which it was segregated by Evans) is in the production of baeomycic acid in the former. Evans (1944) discussed their differences and similarities in detail. The two species also differ in ecology and distribution. Cladonia atlantica grows on acid sand and lignum and is more or less photophilous. Cladonia squamosa is a species of partially shaded,mossy, rich soil habitats. 252 Distribution - Temperate element, Coastal Plain subelement (see Hale, 1961a); endemic. Cladonia beaumontii (Tuck.) Vain. Acta Soc. Faun. Fl. Fenn. 10: 455. 1894. Cladonia santensis f. beaumontii Tuck. Syn. N. Am. Lich. 1: 245. 1882. Material seen - SUFFOLK COUNTY: 8rodo 2282 (87); 21 specimens collected by Latham (Latham). This species is very closely related to 8, atlantica which, however, always shows more or less distinct cups. In addition 8, beaumontii is usually more decorticate than 8, atlantica. Cladonia beaumont88 is a lignum-inhabiting bog species. Distribution - Massachusetts, Connecticut, New York, North Carolina, Florida (Evans, 1950): Temperate element, Coastal Plain subelement; endemic. Cladonia floridana vain. in Sandst. Clad. Exsic. 1196. 1922. Material seen - SUFFOLK COUNTY: Imshaug 88888 (64), 88888 (64), 88888 (86): 9.2212 .623. (79). 15.23 (79). 11.51 (70). 1.2.2.1185). 12.4.;(85). 1:32; (75). 8888 (75); 23 specimens collected by Latham (Latham); Suffolk CO. Airport near Westhampton, R. H. Torre , 1936 (NY); Rock Hill (near) south of Calverton, 885888 8828, 28 June 1937 (NYS). Cladonia floridana is found on exposed or partially shaded sand, or rarely on wood. Although it is almost entirely limited to the coastal plain, R. H. Torrey (in Smiley, 1940) reported its occurrence in Ellenville, N. Y. (Ulster County) in the Shawangunk Mbuntains at an elevation of 2,200 feet. This unlikely distribution is repeated by the heath, 888888.conradii which is typically a coastal plain species but is also found at 1500 feet on Gertrude's Nose, also in the Shawangunk mountains. Distribution - Cape God to Florida (Evans, 1952): Temperate element, Coastal Plain subelement; endemic. 253 Cladonia multiformis Marr. Bryologist 12: 1. 1909. Material seen - SUFFOLK COUNTY: 88888_8888_(111); (locality unknown), Latham 88, 1914 (FH). This very rare species was found on dry soil. Distribution - Nova Scotia, Vermont, Connecticut, Ontario, Michigan, Black Hills, washington, Saskatchewan, Manitoba: Temperate element, North Temperate subelement; Africa (des Abbayes, 1938). Cladonia scabriuscula (Del. in Duby) Nyl. Compt. Rendu 83: 88. 1876. genogyce scabriuscula Del. in Duby, Bot. Call. 632. 1830.- Material seen - KINGS COUNTY: New Lots, G. B. Brainerd, 1866 (BKL); New Lots, G. B. Brainerd, 1866 (BKL 031988); New Lots, G. B. Brainerd, 1866 (BKL). (SUFFOLK COUNTY: Imshaug 88888 (132); Brodo 59~276 (54), 8888_(125), 8888 (84); near Orient, EEEEEELE: May 1914 (FH). This species and its relationship to 8, farinacea is discussed in detail with the latter. Cladonia scabriuscula sens. str. is very rare on Long Island. It is found on mossy ground and on tree bases usually in the shade. Distribution - Nova Scotia, Vermont, Massachusetts, Connecticut, Michigan, Ontario, Black Hills, coastal Alaska; arctic to southern temperate, with oceanic tendencies (Ahti, 1964): Arctic-boreal element-(?); Europe; Asia. Cladonia farinacea (vain.) Evans, Rhodora 52: 95. 1950. 8, furcata scabriuscula f. farinacaea vain. Acts Soc. Faun. F1. Fenn. 4: 339. 1887. Material seen - KINGS COUNTY: New Lots, G. B. Brainerd, (with Cladonia bacillaris) (BKL 031984). SUFFOLK COUNTY: 2E2§2.§2:Zl§,(54)v 828828 (54), 8888 (125), 8888 (111), 8888 (134), 8888 (72); 19 specimens collected by Latham.(Latham). After looking at many specimens of both 8. scabriuscula and 8. firinacea from several parts of the country, I am not at all convinced that the two are —r____i 254 actually different species. In 8, scabriusculg, the podetia are typically tall, branched, covered with small or large squamules (often sorediate) and become granular sorediate towards their tips. The squamules are often very inconspicuous on the upper half of the podetium and the granular soredia are often abundant over the greater part of the podetium. The Long Island material of 8, scabriusculg_usually is short (less than 20 mm tall), squamulose and irregularly sorediate with clumps of granular soredia. 88§doni8_farinacea, typically, has tall podetia which are infrequently branched, farinose sorediate for most of their length, and almost devoid of podetial squamules. Long Island 8, farinacea, however, is rather short, often has granular soredia and occasionally even shows some podetial squamules. Evans identified all of Latham's 8, gggbriuggula sens lat. as 8, farinacea including a specimen which I am calling 8, scabriuscula sens. str. (Latham.8888). In other words, Evans' concept of 8, farinacea was apparently very broad and allowed for considerable variation in the principal separating characters. Ellflsnié.£££12££2&.1i usually found in dry exposed grassy fields or on eroded ground. Cladonia scabriuscula is usually on richer soil in more shaded localities. The distributions of the two species seem to be fairly distinct in most areas. ‘ Distribution - Widely distributed in North America; in eastern part, south to North Carolina and west to Wisconsin (Evans, 1950); Fonts Arenas, southern tip of Chile (type locality), Port Famine, Straits of Magellan (Evans, 1950); Asia. 255 Cladonia furcata (Huds.) Schrad. Spic. F1. Germ. 107. 1794. Lichen furcatus Huds. F1. Angl. 458. 1762. Material seen - QUEENS COUNTY: Fresh Pond, 88888, 1890, (BKL 031989). SUFFOLK COUNTY: Imshaug 88888? (116), 88888 (116); §£9d° 59-174 (1008), 8888 (70), 8888 (125), 8888 (45), 8888 (84), 8888 (111), 8888 (111), 8828 (126), 3.18.5. (69), _3_gs_q (119), _3_3_§_s_ (94), 2229.9 (94), 11.3.92 (112); 52 specimens collected by Latham including 8888? (Shinnecock Hills) (Latham); Northport, 88888, December 1900 (BKL 031987); ?, 888888, May 1914 (FH); Southold, Latham 188, 4 October 1914 (FH). Cladonia furcata shows considerable morphological variation with various ecological situations. It is usually pale green and more or less squamulose in the shade in mossy banks, and is slender, distinctly browned, and essentially devoid of squamules in fully exposed localities (see page 119). A small percentage of the northeastern material of 8, furcata shows the presence of atranorine, including several specimens from Long Island (see above), one from Nantucket (88888 8888) and one from Cape Cod (§£2§2.&§§Q)' These specimens which appear like 8, furcata but differ chemically were called 8, subrangiformis Sandst. by Evans (1954). Ahti (1962) examined the type of the latter and stated that it seems to be distinct from 8, furcata. He believes the North American material with atranorine, however, is merely a chemical race of 8, furcata. 88889nia furcata is found on exposed or partially shaded sandy or grassy ground and, rarely, is also found on wood or mossy boulders. Distribution - From arctic regions southward into Mexico (Evans, 1930), but not reported by Hale (1954a), Thomson (1953, 1955), or Ahti (1964): Arctic-boreal element (7); circumboreal. Contains atranorine, as demonstrated with GAOT. 256 Cladonia carassensis Vain. Acta Soc. Faun. F1. Fenn. 4: 313. 1887. Material seen - SUFFOLK COUNTY: Three Mile Harbor, EEEEEE.Z§£§£: 17 April 1947 (Latham); Riverhead, LéfihaE.30565: 3 April 1952 (Latham). The Latham specimens were found on rotten wood and sandy soil in bogs and swamps. Evans (1950) discusses the species in detail. Distribution - Massachusetts, Connecticut, Oregon, Haiti, Brazil , New Zealand (Evans, 1950): Tropical element, Oceanic subelement; eastern Europe (Evans, 1950); Asia (Asahina, 1950, sub 8, jagonica Vain.). Subsection Uhciales (Del.) Vain. Cladonia boryi Tuck. Lich. Am. Sept. Exsic. 36. 1847. Material seen - NASSAU COUNTY: Meadowbrook valley, Hemstead Plains, 888888, 27 March 1918(mSUFFOLK COUNTY: 28 specimens collected by Imshaug and/or Brodo; 40 specimens collected by Latham (Latham); 16 specimens collected by R. H. Torrey, 1933-1937 (NY); Orient Point, (collector unknown), September 1870 (FH); Southampton, 88888, 3-7 September 1898 (NY); Wading River, 8888, September (NYS); Wading River,‘888§ (NYS); Orient, 88883 (BKL); Orient Point, Latham, 8 December 1909 (NYS); Orient, Latham 8, 1913 (FH); (locality unknown), LEEEEE;§Q: 1914 (FH); Three Mile Harbor, LEEEEE;Z§ELZ: 17 April 1947 (FH); Tiana Beach, 8. Smith 88888, 4 August 1959 (NYS); 3 miles south of Montauk Point, 888888_8288, 7 September 1961 (M30). The external morphology of this species is very variable and one should mainly rely on the internal anatomy described in the key. Cladonia 885y8 is strictly an exposed sand plain and sand dune species. Distribution - Nova Scotia, Massachusetts, Connecticut, New Jersey: Temperate element, Coastal Plain subelement (?); Asia and Brazil (Vainio, 1887). Cladonia caroliniana Tuck. Am. Jour. Sci. Arts 11. 25: 427. 1858. Material seen - NASSAU COUNTY: Brodo 3346 (8), 3349 (8); Plain Edge, 257 S. Cain 888, 3 August 1936, Andropogonetug_Hemgsteadii (NY). SUFFOLK COUNTY: 25 specimens collected by Imshaug and/or Brodo; 38 specimens collected by Latham (Latham); Coram, N. Taylor 8, 15 June 1922 (NY); Selden, S. Cain 358, 30 June 1936 (NY); Coram, R. H. Torrey (2 specimens), 1936 (NY); Reeves Bay near Flanders, R. H. Torrey, 1937 (NY); Rt. 112 north of Coram, R. H. Torrey, (2 specimens), 1936 (NY); Barling Hollow ( : Baiting Hollow ?), R. H. Torrey, 1937 (NY); Pikes Beach, Westhampton, R. H. Torrey, 1936 (NY); 1.3 miles W of Middle Island, S. Smith 17716, 12 March 1955 (NYS); Tiana Beach, S. Smith 88888, 4 August 1959 (NYS). Cladonia caroliniana, like 8, uncialis, is found on sandy or mossy soil in exposed or partially shaded localities. Distribution - Throughout eastern United States (Evans, 1952): Temperate element, East Temperate subelement; endemic. 888don88_uncialis (L.) Web. in Wigg. Primit. F1. Holsat. 90. 1780. 888888Duncialis L. Sp. P1. 1153. 1753. Material seen - QUEENS COUNTY: Ridgewood, G. B. Brainerd, (1866?) (BKL). NASSAU COUNTY: Meadow Brook Valley, Hempstead Plains, 888888, 27 March 1918 (NY). SUFFOLK COUNTY: 36 specimens collected by Imshaug and/or Brodo; 66 specimens collected by Latham.(Latham); Orient, Egggg (BKL); Shinnecock Hills, R. H. Torrey, 1933 (NY); Montauk Point, R. H. Torrey, 1933 (NY); Airport near Westhampton, R. H. Torrey, 1936 (NY); Coram, R. H. Torrey, 1936 (NY); Selden, S. Cain 888, 30 Jane 1936 (NY). 888donia ggcialis shows several growth forms apparently in response to different ecological situations. In exposed areas on bare sand, the podetia are slender and crowded forming tight flattened cushions; in shaded localities on mossy soil or in protected spots where moisture is usually abundant, the podetia become broad, tall, and erect without forming distinct cushions. 258 The smooth, somewhat pruinose podetial inner lining, however, is constant for the species. The chemistry of the species is somewhat variable, with squamatic acid occurring in some geographic areas and not in others (see Evans, 1944). On Long Island, all specimens have a medullary white UV fluorescence and all those extracted with acetone and tested with GE solution showed the presence of squamatic acid. A study of the material of 8, uncialis in the Michigan State University herbarium revealed that the squamatic acid strain is found in the Appalachian Mountain range and along the northeast coast as far north as New Brunswick as well as in boreal and arctic Canada and Alaska. The squamatic negative strain seems to be confined to the Great Lakes region and northern New England. In Europe the squamatic strain is found in central portions of the continent and the inactive strain is mainly found in Scandinavia and Russia (Evans, 1944). Distribution - Arctic regions south to Alabama (Evans, 1930): Arctic- boreal element; circumboreal. Subgenus CLADINA (Nyl.) Leight. em. Vain. Section Bicornutae Abb. Cladonia evansii Abb. Lond. Jour. Bot. 76: 351. 1938. Material seen - SUFFOLK COUNTY: Shinnecock, Latham 33156, 30 April 1926. (US: Evans, Latham). Latham's specimen was first identified as Cladonia 888888 f. condensate (Flgrke) Sandst. by Evans who noted the presence of usnic acid, perlatolic acid and atranorine. Ahti and Thomson later studied the same specimen and called it 8, evansii (Ahti, 1961). When I first came upon a duplicate of the specimen in the Latham.herbarium, I referred it to 8, terrae-novae Ahti having demonstrated atranorine and what appeared to be usnic acid in GAOT solution. However, after seeing the Evans herbarium material which was much 259 better developed, and after examining many specimens of both 8, terraennovae, which I collected on Cape Cod and Nantucket Island, and 8. evansii from the Michigan State University herbarium, I also came to the conclusion that the Latham specimen must indeed be 8, evansii with Long Island thus representing its northernmost locality. Ahti (in letter) emphasized the importance of the different kinds of branching in the two species: 8, evansii has isotomic branching whereas 8, terrae~novae has anisotomic branching. The podetial surface in both species is more or less decomposed and fibrous. Usnic acid is rarely present in 8, evansii and very rarely absent in 8, terrae-novae. Cladonia terrae-novae is common throughout the Cape Cod region and is also found in the Forked River bogs of southern New Jersey. It is, therefore, likely that it will be found on Long Island with additional collecting. An interesting phytogeographic parallel with the distribution of 8, terrae- 22222 involves the tiny fern, Schizaea pusilla. The two species have their southern-most locality in the very same bog and are found almost side-by-side. The fern also has not been found on Long Island but reoccurs furthér north (especially Nova Scotia and Newfoundland) in great abundance. Latham's specimen of 8, evansii was found on dry sandy soil on an open hill. Distribution - Southeastern United States and the West Indies (map: Ahti, 1961): Temperate element, Coastal Plain subelement; endemic. Section Algestres Abb. Cladonia algestris (L.) Rabenh., Clad. Europ. 11. 1860. 888888 rangiferinus (K.) algestris L. Sp. P1. 1153. 1753. Material seen - KINGS COUNTY: Forest Park, 88888, 31 November 1890, (BKL 031993). QUEENS COUNTY: Ridgewood, G. B. Brainerd, (1866?) 260 (BKL). SUFFOLK COUNTY: 88888_888 (79), 8888 (120); 12 specimens collected by Latham (Latham). Cladonia alpestris at one time was probably fairly abundant throughout the eastern part of Long Island (Latham, 1949) even having occurred in the New York City area at one time. Latham (1949) gives an extensive account of the species' distribution and ecology on the island. I have only seen 8, alpestris twice on Long Island. The first observation was in a pine barren area south of Riverhead and was represented by a tiny fragment of a thallus possibly blown there from a larger colony nearby which I could not locate. Latham took me to his "Colony seven" (Latham, 1949) at Napeague Beach which, at one time, was "in excess of 300 feet in diameter" but at the time of our visit consisted of but a few plants scattered among low shrubs, bearberry and Cladonia submitis. Distribution - (Figure 25 ) Arctic-boreal element, circumboreal (Ahti, 1961). Section 222222.Abb- Cladonia subtenuis (Abb.) Evans. Trans. Conn. Acad. Arts Sci. 35: 536. 1944. Cladonia £222l£.*.§l° subtenuis Abb. Bull. Soc. Sci: Bretagne 16: 108. 1939. f. subtenuis ‘ Material seen - KINGS COUNTY: Forest Park, 88888, 31 November 1890, (BKL 031993). NASSAU COUNTY: Valley Stream, 888888, 17 November 1900 (BKL). SUFFOLK COUNTY: 95 specimens collected by Imshaug and/or Brodo; 92 specimens collected by Latham (Latham); East Point, 221125.23; 2-3 July 1918 (BKL); Orient Point, Dillman, 1927 (NY); Calverton, EEEEEE;Z§§2: 17 September 1937 (NYS); 10 specimens collected by R. H. Torrey (NY); 1.3 miles W of Middle Island, S. Smith 17715, 12 March 1955 (NYS); Eastport (vicinity), 261 S. Smith 28512, 28511, 28510, 5 August 1959 (NYS). f. cinerea Ahti, Ann. Bot. Soc. 2001. Bot. Fenn. "Vanamo" 32: 69. 1961. Material seen -SUFFOLK COUNTY: Promised Land, Latham 27630, 2 June 1951 (Latham) (Holotype); Peconic, Latham 23445, 11 April 1945 (Latham). Ahti (1961) presents a full discussion of f. cinerea, which differs from f. subtenuis only in lacking usnic acid. If the thalli of 8, subtenuis are fertile, which is rare, the branches are shorter, stouter, and more verrucose than sterile specimens. Cladonia subtenuis and 8, arbuscula are the two species in the Long Island Cladinae most difficult to separate. They have the same chemical constituents (fumarprotocetraric acid, usnic acid, and ursolic acid) and their morphologies overlap to a large degree. Ahti (1961) separates the two largely as is shown in the following table: C. subtenuis C. arbuscula l. branching mostly dichotomous 1. branching mostly tri- and tetrachotomous 2. axils mostly closed 2. axils mostly Open 3. slender branches 3. heavy robust branches 4. main branch often sub- to 4. main branch robust and very indistinct distinct 5. branchlets mostly erect 5. branchlets mostly unilaterally falcate 6. podetial surface smooth 6. podetial surface t’warty 7. pycnidial jelly red 7. pycnidial jelly hyaline Upon examining specimens determined by Ahti as 8, subtenuis and 8, arbuscu88, and after personally examining scores of specimens from Long Island and nearby Cape Cod and southern New Jersey, it appears that only a few of these characters approach constancy. There are many specimens of 8, subtenuis which are quite robust and have heavy main stems, unlike typical subtenuis. On occasional specimens, axils may commonly be open and sometimes may even show whorls of branchlets around 262 the gaping hole, although this latter condition is very rare. The pycnidial jelly of specimens approaching 8, arbuscula so closely should be examined. The jelly will be reddish or red-brown in subtenuis and colorless in arbuscula. This was done in some of the questionable, very robust specimens of subtenuis from Long Island, and helped establish the range of variation to be expected in this very variable species. Unfortunately, pycnidial jelly can only be examined from relatively fresh specimens (not more than a few years old). Cladonia subtenuis is most characteristic of partially shaded oak or pine- oak forests but can also be found in Open sand barrens associated with 8, submitis and 8, 888y8. Distribution (f. subtenuis)- Eastern United States (map: Ahti, 1961): Temperate element, East Temperate subelement; British Guiana (map: Ahti, 1961). (f. cinerea) - New England (Ahti, 1961). Section Cladina Cladonia rangiferina (L.) Web. in Wigg. Prim. Fl. Holsaticae 90. 1780. Lichen rangiferinus L. Sp. P1. 1153. 1753. subsp. ran ferina var. ranggfgrina. Material seen - SUFFOLK COUNTY: §£2§2.lQl§ (82), 8888 (83); 25 specimens collected by Latham (Latham). Cladonia rangiferina, like 8, algestris is a rare member of the community on open sand-dunes and sand plains on Long Island but is found abundantly in the Cape Cod region in the same community. Both species were previously more comon on the island than they are now (see page 370). Distribution - Throughout arctic, boreal, east temperate, and west montane North America (map: Ahti, 1961): Arctic-boreal element; circumboreal. 263 Cladonia arbuscula (Wallr.) Rabenh. Deutschl. Kryp. Fl. 2: 110. 1845. Pa§e88aria foliacea var. m. Arbuscula ws11r. Fl. Crypt. Germ. 1: 425. 1831. Material seen - SUFFOLK COUNTY: Brodo 59-284 (82), 223;.(120)3 Calverton, Latham 7547, 10 October 1933 (Latham); Flanders, Latham 24717 (Latham); Calverton, R. H. Torrey (with 8, subtenuis), 1936 (NY). The nomenclatural problems of this species, called 8, sylvatica (L.) Hoffm. by most authors are discussed in detail by Ahti (1961). The Long Island material of this species belongs to Ahti's subsp. arbuscula, chemical strain I (with fumarprotocetraric acid). The similarities between 8, arbuscula and 8, subtenuis are discussed under the latter species (page 261) . Cladonia arbuscula was found associated with 8, submitis and Cetraria islandica on sand dunes. Distribution - (see maps: Ahti, 1961). (sens. lat.) - Arctic~boreal element; circumborea1o(subsp. arbuscula, chemical strain 1) - Eastern boreal and temperate North America; Temperate element, North Temperate subelement (?), but clearly boreal to arctic in Eurasia. Cladonia submitis Evans, Rhodora 45: 435. 1943. Material seen - KINGS COUNTY: Forest Park, 88888, 31 November 1890 (BKL 031993). SUFFOLK COUNTY: 66 specimens collected by Imshaug and/or Brodo; 36 specimens collected by Latham.(Latham); Southampton, 88888, 3-7 September 1898 (NY); East Point, 88y888_88, 2-3 July 1918 (BKL); Farmingville, 88y88, August 1916 (Staten Island); Pike's Beach, Westhampton, R. H. Torrey, 1936 (NY); Holtsville, R. H. Torrey, 1936, (NY); Selden, R. H. Torrey, 1936 (NY); Route 112 north of Coram, R. H. Torrey, 1936 (NY); Reeves Bay near Flanders, R. H. Torrey, 1937 (NY); Riverhead, S. Smith 28444, 28559, 28560, 7 August 1959 (NYS); Selden, S. Cain 888, 30 June 1936 (NY); Noyack, Latham 26423, 17 April 1947 (FH). 264 Ahti (1961) reported that an isotype specimen of 8, submitis which he examined appeared to be 8, 88888. Upon checking the holotype specimen in the Evans herbarium I discovered that it was already annotated by Ahti (in 1961) and by Thomson (in 1962). Ahti marked it as pseudonorangiformic absent, but Thomson noted that with the help of Kurakowa, he demonstrated pseudonorangiformic acid in small amounts in the greater portion of the material. I attempted to recrystallize the crucial chemical myself, but met with no success. In this connection, it should be mentioned that the holotype material is not a typical example of 8, submitis from a morphological point of view. Although some branchlets show the characteristic prongs and robust nature of the species, most of the material is rather slender. The Sandstede exsiccats nos. 1564 and 1565, both on the same sheet as the holotype and both annotated by Evans as being 8, submitis and containing “C” (pseudonorangiformic acid), are much more representative of typical 8, submitis. 88880nig submitis seems to have two basic growth forms on Long Island: one is prostrate and sprawling and the other is erect and often tufted. The former is characteristic of the isolated thalli in Open sand dune areas and exposed sand barrens; the upright form is usually seen in protected situations, between clumps of grass, in extensive colonies on the dunes, and in partially shaded localities. This latter form often appears very much like 8, arbuscula, which, however, is PD + red and lacks pseudonorangiformic acid. The prostrate form has no parallel in the Cladinae and is easily identified in the field. It is interesting although puzzling that 8, submitis is very abundant in south shore dune habitats, but is entirely absent from very similar habitats on the north shore. There are three observable factors correlating with its occurrence on the south shore: the presence of a continuous fore- dune between the community and ocean, the presence of Pinus rigida in the 9 ¥# 265 immediate areas, and the high acidity of the sand. These three factors are probably somewhat interrelated, and may affect the Cladonia distribution directly or indirectly, and act either independently or together. It is known that salt mist and salt spray causes maritime substrates to become more alkaline than normal (Barkman, 1958). It is also known that Pinus rigida is intolerant of large quantities of salt spray (Boyce, 1954). Ahti (1961) stated that Cladonia submitis is intolerant to salt spray and is never found near the ocean. All these facts seem to strongly suggest that the salt spray on the south shore, blocked to a large extent by the foredune, is not nearly as abundant as it is on the north shore where the only protection comes from low dunes and hollows (see Oostings and Billings, 1942). It is, therefore, the salt spray, rather than any directly observed factor such as sand pH or the accumulation of pine detritus, which very likely limits the distribution of 8, submitis. Distribution - Atlantic coastal plain (map: Ahti, 1961): Temperate element, Coastal Plain subelement; Japan (Ahti, 1961). 88adonia m8888_Sandst. Clad. exs. no. 55. 1918. Material seen - SUFFOLK COUNTY: Orient, West Long Beach, Latham 23437, 7 April 1945 (Latham); Promised Land, Latham 25473, 1 April 1946 (Latham). This species is extremely rare on Long Island. It is apparently a member of the community on exposed sand with other Cladinae. Distribution - Throughout the arctic and boreal northern hemisphere (map: Ahti, 1961): Arctic-boreal element; circumboreal. 266 UMBILICARIACEAE UMBILICARIA Umbilicaria mammulata (Ach.) Tuck. Proc. Am. Acad. Arts Sci. 1: 261. 1848. gyrophora mammulata Ach. Syn. Lich. 67. 1814. Material seen - SUFFOLK COUNTY: Brodo 3848 (76); Mbntauk, 888888, May 1920 (Latham); Plum Island, 888888, July 1931 (Latham). Llano (1950) considered Tuckerman's transfer of Gyrophora mammulata Ach. to umbilicaria invalid since Tuckerman, not knowing the true identity of Acharius' species, was actually working with what is now known as 8, car888niana Tuck. Llano, therefore, proposed the new combination 8, 98mmulata (Ach.) Llano and considered 8, mammulata (Ach.) Tuck. 888 Llano as a synonym of 8, caroliniana. Llano's transfer is not necessary since, although Tuckerman was mistaken about the identity of his new combination, it was still validly published and must stand. The species has often been considered under the names Umbilicaria dillenii Tuck. or Gyrophora dillenii (Tuck.) M311. Arg. Of the three Umbilicariae on Long Island, this species is the only one I saw growing in the field. It was found on exposed granitic boulders at the summit of a morainal hill south of Riverhead. Torrey (1933) also reported it from the Wading River region. Distribution - Temperate element, Appalachian subelement, Appalachian- Great Lakes unit; endemic (map: Llano, 1950). Umbilicaria muhlenbergi8 (Ach.) Tuck. Enum. N. Am. Lich. 55. 1845. Gygoghora muhlenbergii Ach. Lich. Univ. 227. 1810. Material seen - SUFFOLK COUNTY: Gardiner's Island, 888888, 28 June 1927 (Latham); Bald Hill, 3 miles S of Calverton, 888888, 1 July 1937 (Latham); Yaphank, Wm. Davis, 3 January 1929 (Staten Island). 267 This species is treated in the genus Actinogyra by Llano (1950). It is found on boulders. Distribution - Temperate element, Appalachian subelement, Appalachian» Great Lakes unit; Europe (Pbelt, 1963); north temperate regions of Asia (map: Llano, 1950). Umbilicaria papulosa (Ach.) Nyl. Mem. Soc. Sci. Nat. Cherb. 5: 107. 1857. gerphora papulosa Ach. Lich. Univ. 226. 1810. Material seen - SUFFOLK COUNTY: Wading River, Latham 2643, 20 July 1926 (Latham). Llano (1950) considered this species in the genus Lasallia. The species apparently is unknown on Long Island outside the Wading River region. I have searched the area for Umbilicariae without success but Latham (see above) and Raymond Torrey (1933) collected 8, papulosa there. On Cape Cod (Barnestable County, N of Woods Hole, Brodo 3927, 3956), I collected several specimens of this species. It was growing abundantly over almost all exposed and partially shaded boulders in the area but was found nowhere else on the Cape. Distribution - Temperate element, Appalachian subelement, Appalachian~ Great Lakes - Rocky Mountain unit, with several west coast localities; Africa (map: Llano, 1950). ACAROSPORACEAE SARCOGYNE Sarcogyge gigggg (Ram. in Lam. & DC.) Kremp. Denkschr. Kgl. Bayer. Bot. Gas. 4: 212. 1861. Lighen clavus Ram. in Lam. & DC. F1. Franc. ed. 3, 2: 348. 1805. Material seen - SUFFOLK COUNTY: Brodo 779 (90A), Z88_(90A), 1715 (133), 268 l888_(127), 8888 (123), 8198.(111), 8818 (106), 8888 (62), 8811 (94), 8888. (134), 8888 (134), 88§8_(76); Orient, Latham 22246, 3 May 1914 (Latham); probably Montauk Point, von Scheur, 22 July 1895 (M0). Fink (1935) treats this species in the genus Biatorella. Sarcogyne clavus is found on exposed granitic boulders. Distribution - Connecticut, New York, Alabama, and California (Magnusson, 1935); Minnesota. Black Hills: Temperate element, North Temperate subelement (?); Europe. Sarcogyne privigna (Ach.) Mass. Geneac. Lich. 10. 1854. Lecidea privigna Ach. Meth. Lich. 49. 1803. Material seen - SUFFOLK COUNTY: Brodo 961 (S of 50). The similarity of this species with 8, glgggg and 8, simplex (as well as 8, pruinose) is discussed in some detail by Magnusson (1935). Sarcogyne privigna is similar to 8, 213222.1n having a smooth, red-black apothecial disk but differs from the latter in having small (less than 1 mm broad), concave, irregular apothecia with prominent margins. Distribution - New Hempshire, Connecticut, and New Mexico (Magnusson, 1935); Black Hills; Europe. Sarcggyne simplex (Dav.) Nyl. Mem. Soc. Sci. Nat. 2: 337. 1854. Ligggg_sim21ex Dav. Trans Linn. Soc. Load. 2: 283. 1793. Material seen - SUFFOLK COUNTY: Brodo 1907 (114), 8888_(128), 8888 (119); Orient, LEEEEE: 25 April 1921 (Latham). This species is the only Sarcogyne with small apothecia having rough disks. It was collected on exposed or partially shaded granitic boulders. Distribution - Maine, Connecticut, Tennessee, Minnesota, Black Hills, Hashington, Manitoba; Canadian archepelago (Thomson, 1960): Arctic-boreal element (?); circumboreal. L—_____ 269 ACAROSPORA Acarospora fuscata (Schrad.) Arn. Verhandl. Zool. -Bot. Ges. Wien. 20: 528. 1870. Lichen fuscatus Schrad. Spicil. F1. Germ. 83. 1794. Material seen - NASSAU COUNTY: Brodo 3513 (10). SUFFOLK COUNTY: Imshaug 2255561 (52); Brodo 1556 (103), 8188 (126), 8818 (123), 8888 (108), 8188 (111), £23888_(94), 8888_(134), 8888_(76), 8888 (62); 16 specimens collected by lLfiitham (Latham); Mbntauk, Hither Beach, Latham 27289, 28 October 1947 (M0). Both Magnusson (1929) and Weber (1962) have commented on the extreme morphological variability of this species. The C + red reaction of the cortex 198 also somewhat variable being strongly positive in some cases and almost negative in others. Aggrospora fuscata is found on granite boulders and pebbles in exposed <>r partially shaded localities. In addition, one questionable specimen was <:ollected on calcareous rock (Lgthag 22332, with Lecanora disperse), and one sees found growing on a storm.tide-washed boulder (hygrohaline zone) (85282_ £7.19). Distribution - Northern and middle states (Fink, 1935): Temperate element, North Temperate subelement; Europe; Asia (Magnusson, 1929). PERTUSARIACEAE PERTUSARIA Pertusaria alpine Hepp in Ahles, Pertus. et Conotr. 12. 1860. Material seen - SUFFOLK COUNTY: Orient, 885889, 23 May 1914 (Latham). The Long Island specimen was compared with Cummings' exsiccats (Decades of North American Lichens no. 281 and Lichens Boreali-Americani no. 211), the former of which was cited by Erichsen (1941) as 8, 382828, (These exsiccats are both mixtures of 8, pustulata which has 2 spores per ascus, and 270 8, alpine which has 4-8 spores.) The Long Island specimens were morphologically and anatomically identical with these exsiccats, but the Cummings specimens contained stictic acid (by chromatography) and were K + yellow and PD*+ orange whereas the Long Island specimen was K - or K + yellowish and PD -. (The specimen was too scanty to extract for chromatography.) The ultraviolet fluorescence of the thallus (orange-pink) was the same in all the material however. The Latham specimen is on cedar lignum and not on bark as is the case with the Cummings material. Distribution - Nova Scotia, District of Columbia (Cummings' Decades no. 281), Michigan; Europe. Pertusaria amara (Ach.) Nyl. Bull. Soc. Linn. Norm. II. 6: 288. 1872. Variolaria amara Ach. Kgl. Vet. - Akad. Hya Handl. 163. 1809. Material seen - SUFFOLK COUNTY: 85282,888,(56), 888 (56), 8888_(83), 88818_(83), 8888 (102), 8888 (112); Orient, Long Beach, LEEEEE.§;§§§2 7 December 1944 (Latham); Orient, LEEE§!.§lp 10 May 1914 (Latham). This species is the only KC1+ violet Pertusaria on Long Island. It was always found sterile. Pertusaria amara grows on the bark of various trees usually in or near bogs. Distribution - Nova Scotia, Quebec, Maine, Connecticut, the Smoky Mountains of Tennessee, North Carolina, Michigan, Wisconsin, Black Hills; Washington (Fink, 1935): Temperate element, North Temperate subelement; Europe. Pertusaria multipuncta (Turn.) Nyl. Lich. Scand. 179. 1861. Variolaria multipuncta Turn. Trans. Linn. Soc. Load. 9: 137. 1806. Material seen - SUFFOLK COUNTY: Brodo 857 (47); Greenport, Latham 1983, 27 February 1923 (Latham); Montauk, Latham 3944 (sterile), 7 April 1927 271 (Latham); Greenport, Latham 27287, 16 April 1945 (Latham); Greenport, Latham, 27 February 1923 (Latham). The Pertusaria multipuncta group seems to be a rather heterogeneous complex of KOH -,PD-taxa having one spore per ascus, and includes at least three populations having spore size ranges which do not overlap. Representatives of two of these populations are on Long Island and seem to be morphologically distinct as well (see key). This group certainly needs further study. These specimens were found on the bark of various deciduous trees. Distribution - No comment seems warranted until the taxonomy of the group is clarified. Pertusaria propinqua M311. Arg. Flora 67: 273. 1884. Material seen - SUFFOLK COUNTY: §E£22.§21§ (119). This species, though represented by only a single collection from Long Island, was found abundantly in the locality where it occurred. The description of 8, rubescens Erich. (Erichsen, 1941) agrees very well with the Long Island material except for the lack of zoned spore walls in the former. The type specimen of 8, rubescens is from a hickory in the New Jersey coastal town of "Sea Girton" ( = Sea Girt?). This exactly parallels the Long Island collection in a coastal oak-hickory woods on 888y§.cfr. tomentosa. Without having seen authentic material of 8, rubescens, I am not listing it as a synonym. Although 8, propinqua was described from.a specimen on granite, it appears to be identical with the original collection of 8. torguata 'M'i11.Arg,whichms on bark. I saw the original material of both taxa (in herb. MICH). Since the species is poorly known, a short description of the Long Island material follows: 272 Thallus dark ashy-grey, smooth to rugose, becoming thick and cracked; fruit warts smooth or rough, becoming distinctly constricted at the base in maturity, lighter in color than the thallus (appearing as if their top surfaces were rubbed), l - 2 mm in diameter; ostioles single to many, usually large, ashy to black, usually somewhat depressed; epithecium cinereous, turning violet in KOH; spores 8, irregularly arranged in the ascus, 89-96 x 40-41 u; spore walls zoned. Medulla of fruit warts and thallus PD + yellow, KOH + deep blood red. Norstictic acid demonstrated in KOH. Distribution - Temperate element, Coastal Plain subelement (Fink, 1935 sub 8, marginate Nle; endemic. Pertusaria sghpertusa sp. nov. Material seen - SUFFOLK COUNTY: Brodo 8935 (112), 1436 (83), 1674 (88), 2163 (102), 2289 (87). Thallus virido-cinereus, continuus, rimae acquirendus, rugosus, ultimus minutissime verrucosus; verrucae fructae plerumque disperses, (0.5-) 0.65-1.10 (-1.30) mm diam., leves, hemisphericalesad.subplaniferes; colorae thallis, basibus perspicues constrictis; ostioles 3-7 per verruca fructus, obscures vel pallides, tantum depressiuscules, 0.05 - 0.15 mm diam. Paraphyses tenuisimae (t 1 u), ramiosissimae. Asci 194-236 X 35-42 a, parietis crassa. Sporae hyalinae, non septatae, 97-138 X 35-41 g, parietes 4-10 h crassae, zonates et canaliculates, 2 vel rerissime l per ascus. Epithecium obscurum, KOH + violaceum. Medulla verrucarum fructus et thalli PD +-rubro-aurantiaca, KOH + fulvescens, C -, KC -. Materia chemica: acidum fumarprotocetraricum. Corticola. Holotype: SUFFOLK COUNTY: Three Mile Harbor, on Old Northwest Road 0.7 miles from junction with Alewife Brook Road, 8ggggw8888, 12 July 1960, on bark of éEEE.£22£2! in bog (MSC) (see figure 101). Thallus grey to greenish-grey, continuous, becoming cracked, rugose, and finally minutely verrucose; fruit warts mostly scattered, (0.5-) 0.65 - 1.10 (-1.30) mm in diameter, smooth, hemispherical to flattened, the same 273 color as the thallus, distinctly constricted at the base; ostioles 3-7 per fruit wart, dark or pale, only slightly depressed, 0.05-0.15 mm in diameter. Paraphyses very slender (ca. 1 n), much branched; asci thick- walled, 194-236 X 35-42 p; spores hyaline, nonseptate, 97-138 X 35-41 p, walls 4-10 h thick, zoned and channelled, 2 or very rarely l spore per ascus; epithecium dark, KOH + violet. Medulla of fruit warts and thallus PD + red-orange, KOH * brownish, C -, KC -; fumarprotocetraric acid present. Corticolous. The Long Island material is rather uniform in morphology, but does show some variation in the color of the ostioles (becoming pale in some specimens) and in the depth and breadth of the ostiole depressions (often becoming very deep and up to 0.20 mm broad in maturity). Salscinic acid was demonstrated in all specimens except the type, and is therefore an accessory substance. Specimens containing this substance will have a KOH + yellow changing to red medullary reaction. If we were to accept Erichsen's sections of the genus based on KOH reaction, some specimens would have to be placed in one part of the genus (section Rubescentes) and some in another (section Insensibiles). Since this is untenable, Erichsen's sections cannot be used. The epithet "subpertusa" is used for this new species to emphasize its similarity in general appearance and spore type to 8. pertusa (L.) Tuck. Pertusaria pertusa has larger spores (145-229 X 40-82 g) and contains stictic acid. All other North American KOH + red Pertusariee have norstictic acid rather than salacinic acid and none, to my knowledge, contain fumar- protocetraric acid as well. Of the four Long Island specimens three were found growing on the bark of Acer rubrum in bogs or swamps and one was on black oak bark. I 274 also collected a specimen in southern New Jersey (Atsion, Brodo 3587) on a black oak just outside a bog. Distribution - New Jersey; endemic. Pertusaria trachythallina Erichs. in Degel. Ark. Bot. 30A(1): 36. 1940. Material seen - SUFFOLK COUNTY: 31 specimens collected by Imshaug end/or Brodo. This species is discussed at length by Erichsen in his original description. The species is found on the bark of various deciduous trees, usually at breast height. It can be considered a member of the black oak-breast height community although it has also been found on Quercus alba and Fagus grandifolia. Distribution - Maine; endemic. Pertusaria tuberculifera Nyl. Act. Soc. Scien. Fenn. 7: 448. 1863. Material seen - QUEENS COUNTY: Ridgewood, G. B. Brainerd, (1866?) (BKL 031906); Ridgewood, G. B. Brainerd, (1866?) (BKL). SUFFOLK COUNTY: 24 specimens collected by Imshaug and/or Brodo; Mentauk, LEE!§E:ZZ§§§2 17 April 1946 (Latham); Springs, East Hampton, Latham 28321, 9 February 1949 (Latham). Pertusaria tuberculifera belongs to Erichsen's subgenus Eupertusaria section Insensibiles.v The material treated here was probably considered under the name 8, leioplaca (Ach.) Lam, & DC. in Fink (1935) where 8, 1e10plac§_ is described as having 4-8 spores. Erichsen (1935) regards 8, leioplaca to be 4-spored alone, or rarely 2-3 or S-spored. The Long Island specimens are all dominantly 8-spored with the 4 spore condition occurring frequently in the same apothecia. Pertusaria'tuberculifera and 8, tetrathalamia (Fee) Nyl. are often considered to be conspecific, but Erichsen (1936) pointed out that the latter has only four spores per ascus. 275 Since the species apparently is fairly common and yet poorly known, a more detailed description of the Long Island material may have some value: Thallus dark ashy-grey, continuous, smooth, becoming rugose and verrucose; fruit warts large, 1-3 mm in diameter, irregular, crowded, distinctly constricted at the base in maturity; spores 4-8, hyaline, (30-)34-40 X (SS-)62-80 (-97) n, walls 6 n thick, zoned, smooth. Medulla of fruit warts KOH -, PD -, KC -, C -. Thallus UV + orange fluorescence. Distribution - South America (type locality), West Indies (Imshaug, 1957b), New Jersey (see above): Tropical element, Coastal Plain subelement(?). Pertusaria velata (Turn.) Nyl. Lich. Scand. 179. 1861. Parmelia 3888£8_Turn. Trans. Linn. Soc. Lond. 9: 143. 1808. Material seen - SUFFOLK COUNTY: §£222.lflll§.(83): 8888 (102), 8888 (102), 8888_(122), 8818 (119); Riverhead, LEEEEE: 16 May 1960 (Latham); Three Mile Harbor, East Hampton Twp., QEQEE.§&Q§J 11 March 1952 (NYS). The C + red disks and thallus and the lecanorine apothecia of this species give it a superficial similarity with a species of Ochrolechia. However, the very large spores, one per ascus, easily refer it to Pertusaria. Pertusaria EELEEE is usually found in humid forests or bogs on the bark of deciduous trees. Almborn (1948) stated that the species is typical of the Pzgenula 888888 society on 8gggg_and has an oceanic affinity. This would be borne out to some extent by its "bceanic" distribution on Long Island. Distribution - Nova Scotia, Maine, Connecticut, Tennessee, North Carolina, Oklahoma, Michigan, Indiana, Iowa, Wisconsin, Minnesota, Black Hills, washington, coastal Alaska, British Columbia: Temperate element, Oceanic subelement (?); Europe (oceanic), Asia, Africa, South America (Almborn, 1948). 276 Pertusaria xanthodes M311. Arg. Flora 67: 286. 1884. Material seen - SUFFOLK COUNTY: 80 specimens collected by Imshaug and/or Brodo; 10 specimens collected by Latham (Latham). The Long Island material agrees well with Maller's type specimen from Texas which I saw in Geneva. The thallus of the type was yellow, having amplieriate fruit warts, each with one pale, more or less depressed ostiole and containing one or two apothecia. The spores were 2 per ascus and showed clearly zoned walls often with one of the walls roughened on the inner surface. This species is easily confused with certain forms of Pertusaria pustulate (Ach.) Duby and was almost surely considered under this name in Fink (1935). Pertusaria pustulate is characterized by spores with thin, smooth, unzoned walls, and by dark ostioles. In addition, the epithecium of 8, pustulate generally turns KOH + violet. Since my material is rather variable, I will break down its description as follows. anstant characters: variable characters: 1. on bark l. ostiole color (pale or dark) 2. spores 2 per ascus 2. concentration of stictic acid 3. spores 30-45 X 70-120 u 3. thickness of thallus 4. spore walls zoned, thick, rough 4. degree of density of fruit 5. stictic acid present warts 6. UV fluorescence pink-orange Almost constant characters: Never seen: 1. yellow color of thallus 1. spore walls thin, smooth 2. ampliariate fruit warts 2. epithecium KOH + violet 3. hypophloedal thallus Pertusaria xanthodes is found on the bark of various species of deciduous trees usually in well lighted situations. Distribution - Cape Cod (Massachusetts), New Jersey, Texas (type locality), 277 West Indies (Imshaug, 1957b) Temperate element (?), Coastal Plain subelement; endemic. MELANARIA Melanaria macounii Lamb, Ann. Rep. Nat. Mus. Can. 132: 286. 1954. Material seen - SUFFOLK COUNTY: 88982.888’(56), 8888 (117), 8888 (102), 8888 (102), 8888_(1l9); Napeague, Latham 2848, 1 March 1927 (Latham). Melanaria macounii resembles Pertusaria pertusa in many respects. Both have polycarpous, smooth fruit warts of the same color as the thallus, both have two spores per ascus, and of approximately the same size range, and both contain stictic acid. In 8, macounii, however, the spores are distinctly radiately channeled and often are brownish, turning sordid violet in KOH. (The hyaline spores, which predominate in the Long Island material, do not give this KOH reaction.) The species is found on the bark of various deciduous trees in humid forests or bogs. Distribution - Nova Scotia (type locality); Great Lakes region(seen in herb. MSC); endemic. LECANORACEAE IONASPIS Ionespis 2§2££.(A0h- in Schaer.) T. Fr. Lich Scand. l: 273. 1871. §y§8ggta odora Ach. in Schaer. Lich. Helv. Spic. 2: 80. 1826. Material seen - SUFFOLK COUNTY: Shelter Island, LEEEEE;Z£§§£» 26 October 1944 (Latham). Latham's specimen agrees perfectly with the description of the species given by Magnusson (1933) in his monograph of the genus. Although Tuckerman (1882) cited a specimen of 8, odora from New Hampshire, Magnusson (1933) 278 stated that the species is "most likely not in North America." Ionaspis lavata Megn. was described from a Merrill collection from Mount Rainer, Washington. This species, however, differs from 8, 88888 in having confluent, brownish apothecia rather than scattered pale pink or yellow-brown apothecia. For some unknown reason, the type specimen of 8, lavata is absent from its packet (in herb. FH) and I therefore have not seen it. The species is listed under Lecanora by Fink (1935). In addition to the reports of this species from the White Mountains of New Hampshire (Tuckerman, 1882) I collected a specimen from Cape Cod, Massachusetts (Brodo 43998) on a granitic boulder in partially shaded oak woods, a similar habitat to that of the Long Island collection. Magnusson (1933, p. 20), however, states its ecology as "on granitic stone on the banks in brooks and lakes at least part of the year wetted by water." Distribution - New Hampshire (Tuckerman, 1882), Massachusetts (see above); Europe ("boreal-alpine species") (Magnusson, 1933). LECANORA Lecanora atra (Huds.) Ach. Lich. Univ. 344. 1810. Lichen ater Huds. Fl. Angl. l: 445. 1762. Material seen - SUFFOLK COUNTY: Montauk, LEEEEE;Z&l§Z; 4 May 1926 (Latham). The species is usually found on stone or tree bark, but is also known to occur on lignum on occasion (Hillman and Grummann, 1957; Erichsen, 1957). Distribution - Alaska, washington, Idaho, Quebec, Michigan, Minnesota, Black Hills, Arizona: Temperate element (1), North Temperate subelement, reported from European and Asian arctic by Lynge (1938, 1928). 279 Lecanora caesiocinerea Nyl. in Malbr. Bull. Soc. Amis Sci. Nat. Rouen 5: 320. 1869. Material seen - NASSAU COUNTY: 8:9do 3505A (10). SUFFOLK COUNTY: EEQQQ. 88_7_l (62) . The Long Island specimens of 8, caesiocinerea agree well with Magnusson's description of this species (Magnusson, 1939) except that they have moniliform rather than submoniliform paraphyses and spores which are slightly smaller (15-20 p long rather than over 20 a long). This species is very similar to 8, cinerea and perhaps should be included there. It differs in the slightly shorter pycnoconidia, the KOH - thalline reaction, and the fewer apothecia. (See additional notes under 8, cinerea.) Lecanora caesiocinerea grows on exposed or partially shaded granitic boulders. Distribution - Nantucket Island (Massachusetts) (Brodo 4004), Black Hills, Arizona; Europe. Lecanora caesiorubella Ach. Lich. Univ. 366. 1810. subsp. lathamii Imsh. & Brodo, in press. Material seen - NASSAU COUNTY: §£2§2.§§2.(13)' SUFFOLK COUNTY: 103 specimens collected by Imshaug and/or Brodo; 19 specimens collected by Latham (Latham); Eastport, Schrenk, 26 June 1894 (MD); Greenport, LEEEEE.l§v 2 August 1914 (FH); Greenport, Latham 195, 20 March 1914 (PH). All the Long Island material of this species belongs to subspecies lathamii. Since a complete discussion of 8, caesiorubella and other members of’the.8v pallida group will soon be published (Imshaug and Brodo, in press), it suffices to say that this subspecies of 8, caesiorubella is characterized by a C + orange-yellow disk and by the presence of protocetraric acid and norstictic acid in apothecial sections, the latter being confined to the stipe. 280 This subspecies of 8, caesiorubella is found on the bark of deciduous trees usually in exposed situations. Distribution (subspecies lathamii) - Nova Scotia to Texas: Temperate element, Coastal Plain subelement; endemic. The species as a whole has a tropical - temperate distribution (including South America, Africa) (Imshaug and Brodo, in press). Lecanora chlarotera Nyl. Bull. Soc. Linn. Norm. II. 6: 274. 1872. Material seen - NASSAU COUNTY: Cold Spring, 88888, 1 April 1900 (BKL). SUFFOLK COUNTY: 83 specimens collected by Imshaug and/or Brodo; 14 specimens collected by Latham (Latham); Montauk Point, Easthampton Twp., northeast of Prospect Hill between Great Pond and Oyster Pond, Ogden 5411, 12 May 1954 (MSG). This species is very common on Long Island, and it is extremely variable. The color of the disks, for example, varies from a pale yellow-brown to a dark chocolate brown; the margins are usually somewhat crenate, but sometimes are quite smooth and even; the epithecium is usually conspicuously granular but sometimes is almost without granules. Large irregular, colorless crystals, however, always can be found in the amphithecium, particularly in the margins, and the epithecial granules and pigment always dissolve completely in KOH. Lecanora ghlarotera is not listed in Hale and Culberson's (1960) check- list but was reported from the Chirichahue Mountains in Arizona by Weber (1963). A number of corticolous species belonging to the Lecanora subfusca group have been reported from eastern or northern United States and in an effort to show the differences between these species and 8, chlarotera, a key to their separation follows. This key is mainly based on the work of Magnusson (1932) to which I have added information which has been published since then as well as some of 281 my own observations. 1. 3. 5. 5. Apothecial margin cortex 50-100 p thick, strongly gelatinous. Exclusively on the bark of 88388 (see Lamb, 1954) (Nova Scotia, Maine) . . . . . . . . . . . . . . . . . . . . . 8, glabrata Ach. Apothecial margin cortex less developed, 8-35 (~50) n thick. . .2 2. Cortex little developed, 8-15 u thick, KOH -; crystals lacking in medulla. Epithecium inspersed with crystals, PD + red (see Degelius, 1941) (Maryland, West Virginia) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8, cinereofusca Magn. 2. Cortex 20-35 (-50) n thick, inner portion distinctly delimited, KOH + strong yellow; medulla usually with heaps or clumps of crystals. . . . . . . . . . . . . . . . . . . . . . . . . . 3 Upper part of hymenium (epithecium) coarsely or finely granular.5 Upper part of hymenium (epithecium) without granules, more or less reddish-brown. . . . . . . . . . . . . . . . . . . . . . . 4 4. Apothecia thick, urn-like, margin coarsely crenulate (Maine, Michigan, Manitoba). . . . . . . . . . . . . 8, subrugosa Nyl. 4. Apothecia thin, margin finely crenulate or smooth (Maine, New York, Connecticut, Tennessee, Muchigan, Oklahoma, Manitoba, Quebec). . . . . . . . . . . . . . . . . . . 8, subfusceta Magn. 0n the bark of conifers. . . . . . . . . . . . . . . . . . . . 6 On the bark of broadleaf trees. . . . . . . . . . . . . . . . . 7 6. Thallus leprose or finely granular; epithecium PD -; disks dark red-brown; margins with a yellowish tint (Maine, Tennessee). . . . . . . . . . . . 1v pinastri (Schaer.) Magn. 6. Thallus smooth or rugose; epithecium PD‘+ orange crystals. Spores (13-) 17-20 x 8-13 u. . 8, insignis Degel. (see below) 282 7. Apothecial margin PD + red (substance unknown)(Connecticut, New York, Nova Scotia, Saskatchewan, Quebec). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8, chlarona (Ach.) Nyl. 7. Apothecial margin PD - or PD + pale yellow . . . . . . . . . .. 8 8. Epithecium PD + red with the production of orange acicular crystals (substance unknown). Apothecial disks strongly convex; margins beaded and often discontinuous; spores 12-14 8‘ x 7-8 p. (Long Island, North Carolina, and Tennessee) . . . . . . . . . . . . . .8. cfr. insignis Degel. (see page286). 8. Epithecium PD -. Apothecial disks flat to somewhat convex; margins sometimes crenulate but always continuous; spores 10-13 x 6-7 u . . . . . . . . . . . . . 8, chlarotera Nyl. Lecanora chlarotera is found on the bark of various deciduous trees throughout the island. Distribution - North American distribution largely unknown, but un- doubtedly common, at least in the east; coastal Massachusetts, New Jersey, Arizona; Europe. Lecanora cinerea (L.) Sommerf. Suppl. Fl. Lapp. 99. 1826. LEEEEB. cinereus L. Mantissa 1:132. 1767. Material seen - NASSAU COUNTY: 88282.88988 (10). SUFFOLK COUNTY: 88288 8888 (100A), 8888_(6l), §§§3,(108): 8188.(111), Z§Q§.(106):.§2§§.(62): 3419 (134), 8888 (134), §£§§.(134): 8888 (76); Orient, LEEEEE:2§Q¢ 19 April 1924 (Latham); Shelter Island, 8888ggi88818, 1 April 1946 (Latham, M0). There are two Long Island species of Lecanora in the section Aspicilia, 8. cinerea and 8. caesioc8nerea, and they appear to be very closely related if in fact they are not conspecific. 283 In Europe, 8, cinerea and 8. caesiocinerea are separated easily by their differing reactions with KOH and by their pycnoconidia which do not even come close to overlapping in length. Lecanora cinerea has a rapid KOH + yellow to red reaction (due to norstictic acid) and pycnoconidia 16-20 u long, whereas 8, caesiocinerea has a KOH - (or dirty reddish-brown) reaction and pycnoconidia 6-12 u long (Hillman & Grummann, 1957). On Long Island, the situation is much more complicated. To begin with, there are three divisions based on KOH reactions instead of two: KOH + red (norstictic acid), KOH + yellow (stictic acid), and KOH -. Secondly, there are three size classes of pycnoconidia, each class correlated with a KOH reaction type. Thirdly, the three pycnoconidial size classes overlap, especially in the KOH positive groups. The KOH + red group on Long Island has pycnoconidia on the small side of the European scale, the KOH negative group has pycnoconidia on the large side of the scale, and the KOH + yellow group (the one for which there seems to be no European parallel), introduces an intermediate size range. Since stictic acid and norstictic acid commonly shift within species, it is reasonable to presume that the stictic acid specimens represent North American chemical variants of the well known 8, cinerea which normally produces norstictic acid. The fact that their pycnoconidial sizes overlap considerably adds to the likelihood of the two variants being conspecific. The KOH - material apparently is 8, ggesiocineree with somewhat larger pycnoconidia than seen in European specimens. Lecanora cinerea is found on exposed or partially shaded granitic rocks. Distribution - Nova Scotia, Maine, Michigan, Iowa, Minnesota, Arizona, Black Hills, Washington, Alaska, arctic Canada: Arctic-boreal element; Europe; Asia (Zahlbruckner, 1930; Lynge, 1928). 284 Lecanora conizaea (Ach.) Nyl. Flora 55: 249. 1872. Lecanora expallens pm 8. conizaea Ach. Lich. Uhiv. 374. 1810. Material seen - QUEENS COUNTY: Ridgewood, G. B. Brainerd, (1866?) (BKL 031909). NASSAU COUNTY: Cold Spring 88288, 1 April 1900 (BKL). SUFFOLK COUNTY: Imshaug 88188 (132), 88118 C (121); §£222.119§ (133), 8813 (123), 8888_(97), 8888 (115); Orient, Latham 100, 23 May 1914 (Latham); Orient, Latham 22257, 20 May 1914 (Latham); Orient Point, 888888, 11 April 1910 (NYS). This species is very similar to Lecanora symmicta (Ach.) Ach. in many respects. The thallus is whitish-green to yellow-green, verruculose to granulose, the disks are yellow to buff or brown, and the spores are of the same size and shape. In 8, syggicta, however, the apothecial margins which are at first smooth, pale, and usually transluscent, rapidly disappear with the spothecial disks becoming strongly convex. Lecanora conizaea has white or thallus-colored, smooth apothecial margins which soon become granulose- sorediate, and finally disappear leaving the disks more or less flat or slightly convex. Some apothecia always show the typical granulose lecanorine margin. Lecanora conizaea grows on the bark of various trees usually in exposed habitats, especially near the ocean. Distribution - Maine, Tennessee, North Carolina, Black Hills; Europe; Asia (Lynge, 1928). Lecanora cupressi Tuck. in Nyl. Flora 55: 251. 1872. Material seen - SUFFOLK COUNTY: Montauk, Latham 3662, 28 April 1926 (Latham). The species was found only once, and was growing on wood of what seems to be Juniperus. — 285 Distribution - Massachusetts to Florida and Louisiana (Fink, 1935): Temperate element, Coastal Plain subelement; endemic. Lecanora dispersg (Pers.) Sommerf. Suppl. Fl. Lapp. 96. 1826. Lichen dispersus Pers. Neue Ann. Bot. 1: 27. 1794. Material seen - KINGS COUNTY: §£2Q2.&§§§ (l). NASSAU COUNTY: 88983 8888 (7). SUFFOLK COUNTY: §£2§2.212§.(84): 8888 (115); Orient, Long Beach, Latham 22332, 7 December 1944 (Latham). The Long Island material of this species agreed well with both American and European specimens examined at the Farlow herbarium. The apothecial margins of 8, disperse are usually described as pruinose or even powdery, but in my observations this is not always the case. The species is similar to 8, hgggp8_in many respects and is often included in that species. Lecanora hageni, however, always shows a distinct, thick, gelatinous apothecial margin cortex whereas 8, disperse is essentially without a cortex of any kind. The latter seems to be confined to calcareous rock and mortar, and the former is most frequently found on bark. Both species are commonly found growing with species of Galaplaca. On Long Island, 8, disperse was only found on mortar and brick. It has the distinction of being the only species found in the westernmost collection locality on the island... in the heart of thickly populated Brooklyn. It is well known in Europe as being a highly city tolerant species (Erichsen, 1957). Distribution - Michigan, Indiana, Minnesota, Black Hills, Arizona, Manitoba, Canadian archipelago: Arctic-boreal element; circumboreal. Lecanora hageni (Ach.) Ach. Lich. Univ. 367. 1810. Lichen hageni Ach. Lich. Suec. Prodr. 57. 1798. Material seen - SUFFOLK COUNTY: Brodo 3361 (S of 97). 286 The similarity of this species to 8. disperse has been discussed with the latter. Lecanora hageni was collected only once, on a roadside ELEEE growing with Xanthoria fallax and 8, pgrietina. Distribution - Nova Scotia, Maine, Connecticut, Michigan, Indiana, Minnesota, Black Hills, Rocky Mountains (seen in herb. MSC), washington, Alaska: Temperate element, North Temperate subelement; Europe; Asia (Magnusson, 1940). Lecanora cfr. insignis Degel. Ark. Bot. 30A (3): 53. 1941. Material seen - SUFFOLK COUNTY: Napeague, Latham 2847, 1 March 1927, (Latham). The Latham specimen was compared with the type of 8, insignia kindly sent to me by Dr. Degelius and the two agreed in all respects except spore size and substrate type. The spores of the Long Island material, from oak bark, were smaller than those of the type from the bark of 88888. Poelt and Schauer discovered a correlation of spore size and substrate in specimens of 8, insignis collected recently by the latter in Austria. Small spored specimens were from.deciduous trees and larger spored specimens were from coniferous trees (Poelt, and Schauer, pers. comm.). Degelius' specimens from.the Smoky Mountains showed the same correlation. The small spored population will be described as a new species (Schauer and Brodo, in prep.). The beaded almost disappearing margin and the production of PD + orange needles from the epithecium were both evident in the Long Island specimen. The species is very similar to 8. chlarotera but the larger spores and PD + epithecium of 8, cfr. insigpis easily distinguish the two. Distribution - Smoky Mountains of Tennessee and North Carolina; Austria (see above). 287 Lecanora 8§gp8§_Poelt, Ber. Bayer. Bot. Ges. 29: 64. 1952. Material seen - SUFFOLK COUNTY: 88888_8888_(115); Orient, 888888_81, 23 May 1914 (Latham); Orient, Latham 817, 29 October 1916 (Latham); Orient, Latham 7421, 5 June 1933 (Latham); Orient, Latham 7424 (22245), 5 June 1933 (Latham); Orient, Long Beach, Latham 3940, 27 March 1927 (Latham); Orient, Latham, 3 March 1915 (Latham); Orient Point, Latham 8, 18 April 1910 (NYS); Flanders, S. Smith 34927, 1 August 1962 (NYS). Lecanora laevis bears an external resemblance to 8. allophana (Ad1.)Nyl.which, however, has a thick, gelatinous apothecial margin cortex and seems to be exclusively European. Both Lamb (1954) and Laundon (1958) have presented detailed descriptions of 8, laevis. On Long Island it seems to be restricted to the aerohaline stratum on the eastern Long Island coastline. Distribution - Nova Scotia; southern Europe and North Africa (Lamb, 1954); Ireland (Laundon, 1958). Lecanora muralis (Schreb.) Rabenh. Deutschl. Krypt. F1. 2: 42. 1845. 88g§gg_muralis Schreb. Spic. F1. Lips. 130. 1771. Material seen - SUFFOLK COUNTY: Gardiner's Island, Latham 36807, 23 May 1923 (Latham). This species, common on limestone outcrops in the northeast and else- where, probably was introduced into Long Island with a shipment of limestone building materials. It is found nowhere else on the island probably due to the lack of naturally occurring limestone. Distribution - Connecticut, central New York, Michigan, Iowa, Minnesota, Oklahoma, Black Hills, Arizona, Idaho, Washington, Alaska: Temperate element, North Temperate subelement; Europe; Asia (Zahlbruckner, 1930; Magnusson, 1940). 288 Lecanora rubina (Vill.) Ach. Lich. Univ. 412. 1810. Lichen rubinus Vill. Hist. P1. Dauph. 3: 977. 1789. Material seen - SUFFOLK COUNTY: Brodo 1804 (127), 8888 (134). This species was found on exposed granitic boulders within a quarter mile of Long Island Sound. Distribution - Connecticut, Ontario, Michigan, Iowa, Minnesota, Black Hills, Arizona, Idaho, Washington, Alaska, Northern Saskatchewan: Arctic- boreal element, circumboreal (see Ahti, 1964). Lecanora subintricata (Nyl.) T. Fr. Lich. Scand. 1: 265. 1871. Lecanora varia var. subintricata Nyl. Flora 51: 478. 1868. Material seen - SUFFOLK COUNTY: 8Eshaug 25616 B (116); Brodo 795 (90B), 8888 (84), 8888 (57). Lecanora subintricata, although listed by Hale and Culberson (1960), was not listed by Fink (1935) and has not been mentioned in any recent North American literature which I have seen. The Long Island material fits the European descriptions very well. Lecanora fuscidula Degel. is a very similar species from Maine (Degelius, 1940). I examined the type specimen of 8, fuscidula kindly sent to me by Dr. Degelius and found it to differ from 8, subintricata chiefly in having a well developed gelatinous apothecial margin cortex, about 16-20 p thick. In addition, the thallus of 8, fuscidula is scurfy, ashy, and evanes- cent, whereas that of 8, subintricata is greenish and granulose, and usually is well developed. As in Europe (see Hillman & Grummann, 1957) the Long Island 8, 888: intricate was found on old wood and pine bark. It is often associated with Lecidea aeruginosa. Distribution - Europe; Asia (Vainio, 1928). 289 8§g§gpra symm8gga (Ach.) Ach. Syn. Lich. 340. 1814. Lecanora varia var. sxggicta Ach. Lich. Univ. 379. 1810. Material seen - SUFFOLK COUNTY: Brodo 59-261 (54). Lecanora symmicta is very similar to 8, conizaea. It is placed with Lecanora rather than in Lecidea as is often done, due to its apparent close ties with the other members of the Lecanora varia group, many of which lose their margins in maturity. Lecanora symmicta often resembles Lecidea vernalis in certain respects, but the latter has hemispherical apothecia in maturity, and larger spores (15-19 u long). Distribution - Maine, Connecticut, North Carolina, Tennessee, Michigan, Minnesota, Black Hills, Arizona, Washington, Manitoba; northeast Greenland (Lynge, 1940c): Arctic-boreal element (?); Europe; Asia (vainio, 1928). Lecanora cfr. 38888 (Ehrh.) Ach. Lich. Uhiv. 377. 1810. Lichen varius Ehrh. Pl. Crypt. Exs. no. 68. 1785. Material seen - SUFFOLK COUNTY: Orient, Long Beach, 85£ham 3917, 27 March 1927 (Latham). This specimen differs from all the other Lecanorae on Long Island not only in morphology but also in substrate (on bone). Bruce Fink, to whom this specimen was sent for identification many years ago, called it 8, 38888, The fact that its apothecial margin has a well developed gelatinous cortex puts it close to 8, 28888, The Long Island specimen, however, contains atranorine and is therefore KOH + yellow. Most authors regard 8, Zé£i§.33 a KOH - species, although some (e.g.,Hillmann &.Grummann, 1957) regard 8, XEEli as either KOH +Ior KOH -. The Long Island material also seems to be very close to Lecanora sarcopsis (Wehlenb. in Ach.) Rb‘hl. (8. 8881888 [Perm] Ach.) which, however, usually has an indistinct, ungelatinized apothecial cortex and slightly pruinose apothecial disks. 290 Distribution - Connecticut, North Carolina, Indiana, Iowa, Wisconsin, Minnesota, Black Hills, Arizona, Washington, Alaska: Temperate element, North Temperate subelement; Europe; Asia (Lynge, 1928). Lecanora sp. Material seen - SUFFOLK COUNTY: Brodo 1189 (101). This specimen is in the 8, subfusca group and closely resembles 8. subfuscata Magn. except that it does not have large crystals in the amphithecium and has a light grey rather than a dark grey to cinereous thallus. The thallus is granulose to almost sorediate in spots and smooth at the edges. The apothecial disks are deep mahogany brown, flat, 0.5-1.0 mm in diameter, with smooth to slightly crenulate margins. The epithecium is brownish (remaining so in KOH) and is not inspersed with granules (as in 8, chlarotera). The amphithecium, although it does not have large colorless crystals, is filled with smaller crystals. The apothecial cortex is 16-25 a thick and appears to be paraplechtenchymatous. The medulla and cortex are PD + yellow and KOH + yellow. The spores are 9-13 x 6-7 p. It was found on the bark of Quercus alba near the tree base. OCHROLECHIA Ochrg8ech8§.parella (L.) Mass. Ricerch. Anton. Lich. 32. 1852. Lichen parellus L. Mantissa l: 132. 1767. Material seen - SUFFOLK COUNTY: Imshaug 88188 (68), 888888_(86); 88888 22:22; (67). 1.11.3. <78). _1_6_1_2 <69). 29g (45)._£92_6. (78). mg (78). 3392 (66) Riverhead, 8888gg;1181_8, 1 May 1937 (Latham); Riverhead, Latham.36865, 16 May 1960 (Latham); Riverhead, Latham 36932, 16 May 1960 (Latham); Bay View, L££E£!.36953 B, 8 October 1960 (Latham); Orient, Long Beach, 888888, 26 April 1920 (Latham); Greenport, LEEEEE;ZQQ:' 27 September 1914 (FH); Eastport, Schrenk 15, 24 June 1894 (MICH). 291 Ochrolechia parella, the commonest species of Ochrolechia on Long Island, was also found in southern New Jersey and on Cape Cod. It is easily dis~ tinguished from the other species of Ochrolechia on Long Island by its C ~ thallus and its production of variolaric acid. This substance is most easily demonstrated by introducing KOH onto a water mount of apothecial sections and observing the thin colorless needles often in radiate clusters which are produced in the epithecial and amphithecial regions. Verseghy (1962) states that 8, parella is strictly saxicolous, but the Long Island specimens, all corticolous, compared favorably in morphology and chemistry with the type of 8, parella f. striata Vers. (leg. Szatala, Bulgaria, in herb. F). In Verseghy's monograph, my material keys down to 8.;pallescens (L.) Mass. (see discussion under 8, rosella). Ochrolechia parella is found on acid bark of both broad-leaf and coniferous trees, most frequently in bogs. It has also been seen on old wood. Distribution - North American distribution unclear; Asia (Zahlbruckner, 1930). Ochro8ech8g_rosella (M311. Arg.) Vers. Beih. Nova Hedw. l: 110. 1962. Pertusaria pallescens var. rosella Mfill.Arg. Flora 62: 483. 1879. Material seen - SUFFOLK COUNTY: 8ppgp_8881 (102). Verseghy (1962) published the combination Ochrolechia rosella using Pertusaria pallescens var. rosella M811. Arg. (from Asia) as her basionym. Tuckerman's combination, Lecanora pallescens v. rosella was listed as a synonym, but Verseghy mistakenly gave its first date of publication as 1882 (Syn. N. Am. Lich. l: 196) instead of 1872 (Gen. Lich. 125). Since Tuckerman's combination precedes that of filler, and the epithet rosella is at the varietal level in both, it would seem that verseghy's new combination should properly give Tuckerman's name as the basionym. Since it is possible that 292 Maller's rosella and Tuckerman's rosella are different species, especially in view of the fact that Verseghy does notlist Tuckerman's specimens in her notes on distribution and apparently did not see the Tuckerman material, it is difficult to defend considering Tuckerman's name as the true basionym of 8, rosella. From Verseghy's description, except for the single spore measurement given, which is slightly large, it would appear that the Long Island material is 8, rosella (M511. Arg) Vers., and that name will be used until the proper disposition of Tuckerman's name can be determined. The Long Island specimen was identical (except for the lack of sterile rays of tissue in the apothecia) with the material filed under Lecanora pallescens v. rosella in the Tuckerman herbarium. To aid in future discussions of the species, a lectotype should be designed for Tuckerman's epithet, since he did not cite any specimens in his original publication, and in 1882, only wrote "horthern and middle states, Mahlenberg, etc." A specimen collected in 1857 by Tuckerman in Hadley (Massachusetts?) has typical apothecia as well as a good portion of thallus. The apothecial and thalline reactions with hypochloride solution are as stated by Verseghy (1962) in her description of 8, rosella. I suggest that this specimen be regarded as the lectotype. This species has long been confused and misinterpreted in the literature. Although it has generally been considered as a variety of 8, pallescens, (L.) Mass. it actually is not similar to that species at all. True 8, pallescens is apparently a relatively uncommon oceanic species of the British Isles and France and has a C - thallus and a C -, KC«+ red apothecial margin, with conspicuously pruinose apothecial disks. Ochrolechia rosella, according to the material in the Tuckerman herbarium, has a C‘+ red reaction in the thallus and apothecial cortices. The apothecial disks are lightly or not at all pruinose, and sometimes show distinct "rays" of sterile tissue as 293 described by Tuckerman (1872). It is a relatively common northeastern species and is possibly synonomous with a very similar western species, 8. oregonensis Magn. These two taxa are closely related to 8, tartarea (L.) Mass. which also has an intensely C + red reaction in the thallus and apothecial cortices. According to Verseghy (1962), 8. tartarea is exclusively saxicolous. The specimen from Long Island is very well developed. The thallus is light grey to whitish and is rugose to thickly verrucose. Its cortex is C + red but the medulla is C - (and also KOH - and PD -). The apothecia are 10-20(-25) mm in diameter and are urn-shaped (i.e., with a narrow stipe and basal attachment). The apothecial disk is yellow-orange to orange pink, lightly pruinose, and appears very rough. No sterile rays were evident. With both C and KC, the disk turns red. The hymenium is about 200 p thick, and the spores are hyaline, very thin-walled, and measure 40-60 x 25-26 a. The Long Island specimen was collected in a bog on the bark of 8888 rubrum. Distribution - uncertain: Temperate element, "northern and middle states" (Tuckerman, 1882); Asia (Verseghy, 1962). Ochrolechia sp. Material seen - SUFFOLK COUNTY: Greenport, Latham 793, 28 March 1914 (Latham); Southold, Latham 973, 10 March 1922 (Latham). This species has usually been called 8, pallescens (L.) Mass., but ,9, pallescens is quite different in distribution, morphology, and chemistry (see page 292). In Verseghy (1962), the material agrees fairly well with descriptions of both 8, harmandi Vera. and 8, austroamericana (Malme) Vers. However, 8, E£EE£E§£,13 known only from Oceania and the Orient, and the thallus is —V_4-——- —:-— ————#—-- 294 stated to be not continuous. Ochrolechia austroamericana while agreeing better in thallus morphology (continuous, rugose) and being more logical from a phytogeographic viewpoint (from South America), cannot be used for nomen- clatural reasons. The name is a later homonym of 8, austroamericana (R33.) R53. Verseghy created the new combination apparently because the basionym of her taxon (8, tartarea var. austroamericana Malme, 1937) has priority over Rfisfinen's 8, pallescens var. austroamericana R53., 1939). Rgsgnen, however, raised his variety to species level in 1941. Since the Rules of Nomenclature state that only epithets of equal rank have priority over each other, Rasgnen's _O_. austroamerimnadearly has priority over Verseghy's combination. Without having examined any authentic material of either 8, austroamericana sensu Verseghy or 8, harmandi, I decline from introducing a new name since it may well be that such a common species already has a valid name. This species, while usually showing distinctly pruinose disks, often lacks pruina altogether. The C reaction of the thallus and apothecial margin is confined to the medullary regions and is negative in the cortices, exactly opposite from the situation in 8, rosella. It was found on oak and maple bark. Distribution - Cape Cod, southeastern United States, West Indies (seen in herb. MSC). HAEMATOMMA Haematomma ochrophaeum (Tuck.) Mass. Atti I. R. Istit. Veneto III. 5: 253. 1860. Biatora ochrgphaea Tuck. Proc. Amer. Acad. Arts Sci. 1: 253. 1848. Material seen - SUFFOLK COUNTY: Brodo 2125 (102). This species was collected a number of times in bogs on Cape Cod (Brodo 4174, 4205, 4342, 4371). Its morphology, especially the frequent 295 lack of septation in its spores, is discussed by Lamb (1954). The species differs from Haematomma sp. in ecology as well as in morphology being more characteristically found in bogs and swamps on bark and wood than in upland oak and pine forests. Distribution - Nova Scotia, Maine, Massachusetts, New Hampshire, Vermont, New York, North Carolina, West Virginia, Michigan, Ontario, Quebec: Temperate element, Appalachian subelement, Appalachian - Gnmu:Lakes unit, Japan (Culberson, 1963b). Cfr. Haematomma sp. Material seen - SUFFOLK COUNTY: 42 specimens collected by Imshaug and/or Brodo; Orient, Long Beach, Latham 22340, 7 December 1944 (Latham); Montauk, Hither Woods, Latham 27292, 17 April 1946 (Latham). The status of this material is far from clear. At first, it appeared to be identical with sterile material of Haematomma elatinum (Ach.) Mass. (see Culberson, 1963). With further study, mainly at the suggestion of Culberson, it became clear that the soralia were entirely different (originating in irregular breaks in the thallus, distinct, and punctiform in 8, elatinum, and in tiny hollow, globular to vermiform verrucae in this material), although both can produce a granular sorediate crust in the older parts of the thallus. In addition, 8, elatinum is generally found on coniferous bark, and 8, sp. is found on deciduous bark. Pertusaria trachythallina also contains thamnolic acid, and Imshaug (pers. comm.) pointed out that several specimens of that species show vermi- form, hollow sorediate verrucae. It seems odd, however, that no smooth, Pertusaria-like sterile thallus has yet been found among this material, and that only a few of the dozens of fertile Pertusaria specimens show any tendencoy towards the production of hollow verrucae. The distinctive, often thick, white to yellowish, fibrous prothalline margin seen on almost every specimen 296 of Haematomma sp. contrasts with the absent or, at most, very thin, white prothallus of Pertusaria trachythallina. Haematomma leprarioides (Vain.) Vain., described from South America, is a similar species usually found in the sterile condition. Its soredia are farinose, however, and are produced in distinct punctiform soralia not associated with verrucae. With what we now know about this species, it could as well be placed in Pertusaria as in Haematomma, and the only reason for choosing the latter is its superficial similarity to 8, elatinum. Distribution - Maine, North Carolina, Virginia (Culberson, 1963b); Massachusetts (Cape Cod), New Jersey: Temperate element, Appalachian subelement, Appalachian unit (?); Europe. CANDELARIACEAE CANDELARIELLA Candelariella aurella (Hoffm.) Zahlbr. Cat. Lich. Univ. 5: 790. 1928. Verrucaria aurella Hoffm. Deutch. Fl. 2: 197. 1796. Material seen - SUFFOLK COUNTY: Brodo 2799 (84), 8888 (115). From the descriptions given by Hakulinen (1954), these specimens represent var. aurella. This species commonly grows in association with species of Caloplaca on mortar in exposed situations. Distribution - Michigan, Indiana, Iowa, Kansas, Minnesota, Black Hills, California, Washington, Quebec, Canadian archipelago: Arctic-boreal element (?); arctic and temperate Europe (Hakulinen, 1954); Asia (Magnusson, 1940). Candelariella vitellina (Ehrh.) Mflll. Arg. Bull. Herb. Boiss. 2: 47. 1894. Lichen vitellinus Ehrh. Pl. Crypt. Exs. no. 155. 1785. Material seen - SUFFOLK COUNTY: Brodo 1802 (127), 1912 (114), 2368 (123), 297 8818 (108), 8888_(134). Candelariella vitellina was usually found sterile in the Long Island localities, although the few fertile specimens showed typical polysporous asci. Most of the Long Island specimens best fit the description of var. assericole R33. as given by Hakulinen (1954), the thallus being granular~ verruculose with the granules or verrucules becoming crowded into flattened or rounded patches sometimes becoming almost subsquamulose. Many grade into what is better referred to as var. vitellina with the thalline granules and verrucae more dispersed. For this reason, no segregation of the Long Island material into varieties was attempted. On Long Island, the species is found on exposed granitic boulders associated with Sargggyne spp. and Rinodina oreina. Distribution - Maine, Connecticut, Michigan, Indiana, Wisconsin, Minnesota, Black Hills, Arizona, Washington, northern Saskatchewan, Manitoba, Baffin Island: Arctic-boreal element; circumboreal. CANDELARIA Candelaria concolor (Dicks.) Arn. Flora 62: 364. 1879. Lichen concolor Dicks. Fasc. Pl. Crypt. Brit. 3: 18. 1793. var. concolor ’ Material seen - KINGS COUNTY: New Lots, Brainerd ? (with Phxscia millegrana) (BKL 032039). var. 888888_(Tuck.) Burnh. Bryologist 25: 73. 1922. 88889§chistes concolor var. 888888_Tuck. Syn. N. Am. Lich. 1: 52. 1882. Material seen - SUFFOLK COUNTY: Imshaug 88888 (52); £5292H§22232,(67)» 888 (77), 8888_(118), 8888 (67), 8118 (31), 8888.(65). With the exception of the New Lots specimen cited above (var. concolor), all the Long Island material of this species showed virtually no foliose lobes. It was found growing on the bark of various broadleaf trees usually at the 298 base or around raintracks. Distribution - Massachusetts, Connecticut, central New York, Arkansas, Missouri, Michigan, Indiana, Wisconsin, Minnesota, Black Hills, Arizona, Washington: Temperate element, North Temperate subelement; Europe; Asia (Zahlbruckner, 1930). PARMELIACEAE PARMELIOPSIS Parmeliopsis aleurites (Ach.) Nyl. Syn. Lich. 2:54. 1863. 888888 aleurites Ach. Lich. Suec. Prodr. 117. 1798. Material seen - NASSAU COUNTY: Brodo 547 (12), 3509 (10). SUFFOLK COUNTY: 37 specimens collected by Imshaug and/or Brodo; North Sea, Latham 36933c, 20 May 1954 (Latham). Parmeliopsis aleurites is found on the bark of various tree species, especially 88888 rigida and Chamaecyparis thyoides, but also oaks, and is occasionally found on lignum. It is most frequent in well lighted oak and pine forests. 4 Distribution - Maine, Connecticut, Massachusetts, New Jersey, North Carolina (mountains and piedmont), Tennessee, Alabama, Michigan, Minnesota, Black Hills, Arizona, boreal Ontario: Temperate element, East Temperate subelement; Europe; Asia (Vainio, 1928). Parmeliopsis ambigua (Whlf. in Jacq.) Nyl. Syn. Lich. 2: 54. 1863. Lichen ambiguus Wu1f. in Jacq. Coll. Bot. 4: 239. 1790. Material seen - SUFFOLK COUNTY: Imshaug 25806 (86), 88888 (86); 88888 8888.(78), 8818_(87). Both usnic and divaricatic acids were demonstrated in the Long Island material making it 8, ambigua sens. str. (or "chemical race A”) in the treat- ” ment by Culberson (1955c). 299 Parmeliopsis ambigua was usually found on Chameecyparis (occasionally on_88888) in bogs. As with the other species of Parmeliopsis, this one seems to have a strong specificity for conifers throughout its range. It is found abundantly on pine on the coastal plain of North Carolina (Culberson, 1958a), on white cedars farther north, and on spruce and fir in boreal forests. Distribution - Nova Scotia, Maine, Connecticut, New Jersey, Alabama, Michigan, Wisconsin, Black Hills, Arizona, Washington, Alaska, northern Saskatchewan, Manitoba, Baffin Island, arctic Ontario: Arctic-boreal element; circumboreal. Parme18opsis p1acorodia (Ach.) Nyl. Syn. Lich. 2: 55. 1863. Parmelia placorodia Ach. Syn. Lich. 196. 1814. Material seen - 24 specimens collected by Imshaug and/or Brodo; Orient, 888888, 1 April 1920, on Juniperus (Latham); Manorville, Latham 7767, 20 May 1937 (Latham); Riverhead, Latham 8196, 9 March 1938 (Latham); Napeague, Latham 8624, 11 June 1938 (Latham); Napeague, Latham 25985, 11 March 1947 (Latham); Napeague, LEEEE!:§$QZ§: 1 April 1956 (Latham). This species is the most conspicuous foliose member of the pine bark community. In some pine forests the ascending often subfruticose, finely divided and abundantly fruiting thalli of 8, placorodia can be seen on almost every pine tree, especially dead ones where the loose bark has ceased to slough off (see page 30). The substrate specificity of this species has been discussed in detail by Culberson (1955c). Parmeliopsis placorodia is almost exclusively a Elflfléf dwelling species but in various parts of the country can grow on various species within the genus. In the east, the substrate is 8, 883888, in the Great Lakes Region it is 8, banksiana, and in the west it is 8, ponderosa (Culberson, 1961b). On Long Island, ParmeliOpsis p1acorodia has also been collected on Chamaecyparis (twice), Vaccinium corymbosum (once) see page 57) and on 300 Quercgg,coccinea-velutina (twice). Rare occurrences on fence rails and shingles have been noted as well. Distribution - Northeastern United States (map: Culberson, 1955C), Black Hills, Arizona, Ontario: Temperate element, Appalachian subelement, Appalachian - Great Lakes - Rocky Mountain unit; endemic. PARMELIA Parmelia appalechensis W. Culb., Nova Hedw. 4(3-4): 571. 1962. Material seen - Brodo 59-270 (53). This species has long been included in the complex of pseudocyphellate Parmeliae collectively called 8, bolliana M311. Arg. (see Culberson and Culberson, 1956). In his description of the new species, Culberson (1962) indicated how it can be separated from the very similar 8, frondifera Merr. I have seen and collected much material of 8, frondifera in central New York (Madison County, Bridgeport) where it is always very fertile and always has an entirely pale undersurface with numerous pale buff rhizines. The Long Island specimen has a pitch black undersurface only becoming pale tan at the margins, and is covered with black or dark brown rhizines. It is essentially identical with the isotype of 8, appalachensis (Hale, Lich. Amer. Exs. 63 @887). The lobules so characteristic of 8. appalachensis are not well developed on the Long Island specimen but are distinctly present. The Long Island specimen was collected on the mossy base of a Quercus alba in an oak woods. Distribution - Nova Scotia south to North Carolina (figure .27: Temperate element, Appalachian subelement, Appalachian unit (map: Culberson, 1962). . 301 Parmelia arseneana Gyeln. Ann. Mycol. 36: 269. 1938. Material seen - SUFFOLK COUNTY: Brodo 3025 (50), 3853 (76), 3870 (62); Orient, Latham 942B, 25 April 1921 (Latham). This species has been variously treated by different authors. It mainly comprises what Hale (1955b) called Parmelia congpersa, chemical strain no. 1. It is found on granitic boulders. Distribution - Uncertain. Parmelia aurulenta TUck. Am. Jour. Sci. Arts. 11. 25: 424. 1858. Material seen - KINGS COUNTY: Gowanus, G. B. Brainerd, (1866?) (BKL O31946 . sp. Arthothelium taediosum (Nyl.) M511. Arg. Micarea melaena (Nyl.) Hedl. M, prasina (Fr.) K5rb. Opegraphaceae Qpegrapha cinerea Chev. Q, rufescens Pers. Subc lass ASCOMYCETES Order Caliciales 409 Caliciaceae Chaenotheca phaeocephala (Turn.) T.Fr. Order Sphaeriales Verrucariaceae Verrucaria microspora Nyl. y, muralis Ach. y, nigrescens Pers. 1. silicicola Fink in Hedr. Dermatocarpon miniatum (L.) Mann Pyrenulaceae Pygenula nitida (Weig.) Ach. Melanotheca cruenta (Mont.) Mfill. Arg. Trypethelipg.virens Tuck. in W. Darl. Porinaceae Porina cestrensis (Tuck. in W. Darl.) Mull. Arg. g, hibernica James & Swins. in Swins. g, nucula Ach. Order Lecanorales Graphidaceae Xylographa opegraphella Will. in Rothr. Graphis scripta (L.) Ach. Phaeographis dendritica (Ach.) Mull. Arg. 410 Diploschistaceae Diploschistes scruposus (Schreg.) Norm. Gyalectaceae Dimerella diluta (Pers.) Trev. Q. lutea (Dicks.) Trev. Collemataceae Collema subfurvum (Mfill. Arg.) Degel. Leptogium corticola (Tayl.) Tuck. in Lea L, cyanescens (Ach.) K6rb. Pannariaceae Placynthium nigrum (Huds.) 8. Gray Pannaria lurida (Mont.) Nyl. Stictaceae Lobaria pulmonaria (L.) Hoffm. L, quercizans (Ach.) Michx. Nephromaceae Nephroma laevigatum Ach. Peltigeraceae Solorina saccata (L.) Ach. Peltigera aphthosa (L.) Willd. L, canina (L.) Willd. P, polydactyla_(Neck.) Hoffm. P, praetextata (Fldrke in Sommerf.) Vain. Lecideaceae Lecidea aeruginosa Borr. in Hook. & Sowerby Lecidea albocaerulescens (wulf. in Jacq.) Ach. anthracophila Nyl. “4 Ir botrxosa (Fr.) T. Fr. coarctata (Turn. in Sm. & It“ Sowerby) Nyl. L, cyrtidia Tuck. L, erratica K3rb. L, macrocarpp (DC. in Lamb & DC.) Steud. L, myriocarpoides Nyl. L, pylanderi (Anzi) T. Fr. L, guadricolor (Dicks.) Borr. ex Hook. in Sm. L, scalaris (Ach.) Ach. L, uliginosa (Schrad.) Ach. L, varians Ach. L, vernalis (L.) Ach. L, viridescens (Schrad. in Gmel.) Ach. Catillaria glauconigrans (Tuck.) Hasse Bacidia atrogrisea (Del. in Hepp) Kdrb. chlorantha (Tuck.) Fink. In: no chlorococca (Graewe in Stizenb.) Lett. no: chlorostica (Tuck.) Schneid. . intermedia (Hepp in Stizenb.) Arn. It” 411 Bacidia inundata (Fr.) Kdrb. Cladonia calycantha Nyl. L, schweinitzii (Tuck. in W. Darl.) Q, capitata (Michx.) Spreng. Schneid. Q, carassensis Vain. [O B, trisepta (Naeg. in Mflll. Arg.) cariosa (Ach.) Spreng. Zahlbr. IO carneola (Fr.) Fr. ‘B. umbrina (Ach.) Bausch. IO carolipiana Tuck. Rhizocarpon cinereovirens (Mull. Arg.) Q, chlorophaea (F13rke in Sommerf.) Vain. ' Spreng. R, gpapgg (Fldrke in Flot.) Arn. Q, clavulifera Vain. Q, intermedium Degel. Q, coniocraea (Fldrke) Spreng. em. 5, obscuratum (Ach.) Mass. Sandst. R, plicatile (Leight.) A.L. Sm. Q, conista (Ach.) Robb. Stereocaulaceae Q, cristatella Tuck. Pycnothelia papillaria (Ehrh.) Duf. Q, cylindrica (Evans) Evans Stereocaulon saxatile Magn. Q, deformis (L.) Hoffm. Baeomycetaceae Q, didygg (Fee) vain. Baeomyces £23523 Pers. Q, evansii Abb. Cladoniaceae Q, fagigacea (vain.) Evans Cladonia alpestris (L.) Rabenh. Q, fimbriata (L.) Fr. Q, apodocarpa Robb. Q, floerkeana (Fr.) Flbrke Q. arbuscula (Wallr.) Rabenh. Q, floridana Vain. Q, atlantica Evans Q, furcata (Huds.) Schrad. Q, bapgllaris (Ach.) Nyl. Q, incrassata Fldrke Q, beaumontii (Tuck.) Vain. Q, macilenta Hoffm. Q, PQEZL Tuck. Q, ggteocyatha Robb. Q, brevis Sandst. Q, mitis Sandst. caespiticia (Pers.) Fldrke .nultiformis Merr. In [C 412 Cladonia nemoxyna (Ach.) Arn. C. parasitica (Hoffm.) Hoffm. Q, piedmontensis Merr. [O pityrea (Fldrke) Fr. [0 pleurota (Fldrke) Schaer. IO . pflidata (L.) Hoffm. IO rangiferina (L.) Web. robbinsii Evans [0 santensis Tuck. IO [0 scabriuscula (Del. in Duby) Nyl. IO simulata Robb. IO . sguamosa (Scop.) Hoffm. IO strepsilis (Ach.) Vain. IO subcariosa Nyl. IO submitis Evans '0 subtenuis (Abb.) Evans uncialis (L.) Web. “ [0 IO verticillata (Hoffm.) Schaer. vulcanica 2011. no Umbilicariaceae Umbilicaria mammulata (Ach.) Tuck. Q, muhlenbergii (Ach.) Tuck. Q, papulosa (Ach.) Nyl. Acarosporaceae Sarcogype clavus (Ram. in Lam. & DC.) Kremp. Q, privigna (Ach.) Mass. Sarcogyne simplex (Dav.) Nyl. Acarospora fuscata (Schrad.) Arn. Pertusariaceae Pertusaria alpine Hepp g, amagg (Ach.) Nyl. P. multipuncta (Turn.) Nyl. propinque Mflll. Arg. ['6 subpertusa Brodo [’0 O trachythallina Erichs. in Degel. ['0 I'd tuberculifera Nyl. I'd velata (Turn.) Nyl. . xanthodes Mull. Arg. ['13 Melanaria macounii Lamb Lecanoraceae Ionaspis pgppg (Ach. in Schaer.) T. Fr. Lecanora atra_(Huds.) Ach. 'L. caesiocinerea Nyl. caesiorubella Ach. I? Is chlarotera Nyl. Ir cinerea (L.) Sommerf. 'E" conizaea (Ach.) Nyl. Ir cupressi Tuck. 'F‘ dispersa (Pers.) Sommerf. hageni (Ach.) Ach. 'E“ is cfr. insignis Degel. Ir laevis Poelt 'E‘ muralis (Schreb.) Rabenh. 413 Lecanora rubina (Vill.) Ach. L, subintricata (Nyl.) T. Fr. L, syggicta (Ach.) Ach. L. cfr. varia (Ehrh.) Ach. It" . sp. Ochrolechia parella (L.) Mass. Ochrolechia rosella (Mull. Arg.) Vers. 9, sp. Haematomma ochrophaeum (Tuck.) Mass. A. sp. Candelariaceae Candelariella aurella (Hoffm.) Zahlbr. Q, vitellina (Ehrh.) Mull. Arg. Candelaria concolor (Dicks.) Arn. Parmeliaceae Parmeliopsis aleurites (Ach.) Nyl. g, ambigua (Wulf. in Jacq.) Nyl. L, placorodia (Ach.) Nyl. Parmelia appalachensis W. Culb. g, arseneana Gyeln. g, aurulenta Tuck. P, borreri (Turn. ex Sm. in Sm. & Sowerby) Turn. I'd . caperata (L.) Ach. I’d . conppersa (Ach.) Ach. I’d . galbina Ach. Parmelia hypotropa Nyl. I'd livida Tayl. micheuxiana Zahlbr. yo ['0 olivetorum Nyl. I'd perforata (Wulf. in Jacq.) Ach. I'd perlata (Huds.) Ach. I'd plittii Gyeln. reticulata Tayl. in Mack. [*6 I'd robusta Degel. I'd rudecta Ach. saxatilis (L.) Ach. I'd stenophylla (Ach.) Heug. Pd ['6 subaurifera Nyl. Pd sulcata Tayl. in Mack. I'd . tasmanica Hook. & Tayl. Hypogymnia physodes (L.) Nyl. Pseudevernia furfuracea (L.) ZOpf Cetraria ciliaris Ach. Q, fendleri (Nyl.) Tuck. C. islandica (L.) Ach. Q, tuckermanii Oakes in Tuck. Q, viridis Schwein. Anzia colpodes (Ach.) Stizenb. Usneaceae Evernia mesomorpha Nyl. Alectoria glabra Motyka A, nidulifera Norrl. in Nyl. Ramalina complanata (Sw. in Ach.) Ach. 414 Ramalina fastigiate (Liljebl.) Ach. Q. stenospora M311. Arg. Q, willeyi Howe Usnea longissima Ach. Q, mutabilis Stirt. Q, strigosa (Ach.) Eaton IC‘. . subfusca sensu Mbtyka trichodea Ach. IC.‘ Teloschistacese Caloplaca aurantiaca (Lightf.) T. Fr. [0 camptidia (Tuck.) Zahlbr. IO cerina (Ehrh. in Hoffm.) T. Fr. In citrina (Hoffm.) T. Fr. In discolor (Will. in Tuck.) Fink feracissima Magn. IO IO flavovirescens (Wulf.) Dalla Torre & Sarnth. Q, pyracea (Ach.) T. Fr. Xanthoria fallax (Hepp in Arn.) Arn. A, parietina (L.) Beltram. Iglppchistes chrysophthalmus (L.) Beltram. 1, flavicans (Sw.) Norm. Physciaceae Buellia curtisii (Tuck.) Imsh. in Brodo Q, dialyta (Nyl.) Tuck. Buellia polysppra (Will.in Tuck.) Vain. Q, punctata (Hoffm.) Mass. ‘Q. stigmaea Tuck. IQ. stillingiana Stein. Q, turgescens Tuck. Rinodina applanata Magn. . confragosa (Ach.) Kdrb. IFC1 milliaria Tuck. I50 I” oreina (Ach.) Mass. I73 pachysperma Magn. I'r'U . palipg Degel. Pyxine sorediata (Ach.) Mont. in Sagra Physcia adscendens (T. Fr.) Oliv. L, aipolia (Ehrh. in Humb.) Hampe in Ffirnr. P'd millegrana Degel. orbicularis (Neck.) P3tsch in I'd P3tsch & Scheiderm. I'd stellaris (L.) Nyl. I'd subtilis Degel. I’d . tribacoides Nyl. Anaptychia obscurata (Nyl.) Vain. A. palmulata (Michx.) Vain. A, pseudospeciosa Kurok. Class FUNGI IMPERFECTI Lepraria incana (L.) Ach. L, zonata Brodo L, sp. LITERATURE CITED Abbayes, H. des 1934. La vegetation lichénique du Massif Amoricain. Rennes. 267 p. 1938. Some Cladoniae (Lichens) of the British Dominions: S. Africa, Australia, the Antilles; with a dichotomous key to species of the subgenus Cladina. J. Bot. 76: 346-352. 1951. Traité de lichenologie. Encyc10pedie Biologique XLI. 217 p. Acharius, E. 1810. Lichenographia universalis. Gdttingen. 696 p. 1814. Synopsis methodica lichenum. Lund. 392 p. Ahlner, S. 1940. 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Mbnographia Cladoniarum universalis. 1. Acta Soc. Faun. Fl. Fenn. 4: 1-509. 438 Vainio, E. A. 1890. Etude sur la classification naturelle et la morphologie des lichens du Brésil. Acta Soc. Faun. Fl. Fenn. 7 (1): 1-247; 7 (2): 1-256. 1909. Lichens in viciniis hibernae expeditionis vegae prope pagum Pitlekai in Sibiria septentrionali a Dzre E. Amquist collecti. Ark. Bot. 8 (4): 1-175. 1922. Lichenographia Fennica II. Baeomyceae et Lecideales. Acta. Soc. Faun. Fl. Fenn. 53 (1): 1-340. 1928. Enumeratio lichenum in viciniis fluminis Konda (circ. 60o lat. bor.) in Sibiria occidentali crescentium. 'Muistiinpanoja, prof. A. Ahlqvistin kolmannelta tutkimnsretkelti Lanai-Siperiassa (v. 1880). IV. A XXVII. 6: 65-122. ._.1934- Lichenographia Fennica IV. Lecideales II. Acta Soc. Faun. Fl. Fenn. 57 (2): 1-506. Verseghy, K. 1962. Die Gattung Ochrolechia. Beih. Nova Hedw. 1: 1-146. Wainio, E. A. (see vainio, E. A.) Watson, W. 1953. Census catalogue of British lichens. London. 91 p. Watt, A. S. 1947. Pattern and process in the plant community. J. Ecol. 35: 1'22. Weber, W. A. 1962. Environmental modification and the taxonomy of the crustose lichens. Svensk Bot. Tidsk. 56: 293-333. 1963. Lichens of the Chiricahua Mbuntains, Arizona. Univ. Colo. Stud., Ser. Biol. 10: 1-27. and S. Shushan. 1959. Lichens of the Queen Charlotte Islands, Canada, collected in 1957 by Dr. Herman Persson. Svensk Bot. Tidsk. 53: 299-306. wetmore, C. M. 1960. The lichen genus Nephroma in North and Middle America. Publ. Mus. - Mich. State Univ. Biol. Ser. 1 (11): 369-452. 439 Wetmore, C. M. 1963. Catalogue of the lichens of Tasmania. Rev. Bryol. Lichenol. 32: 223-264. (in prep.) Lichens of the Black Hills (South Dakota and Wyoming). Wood, G. S. 1905. Additions to the lichen flora of Long Island. Bryologist 8: 51. 1914. A preliminary list of the lichens found within a radius of 100 miles of New York City. Torreya 14: 73-95. Young, C. 1938. Acidity and moisture in tree bark. Proc. Ind. Acad. Sci. 47: 106-115. Zahlbruckner, A. 1922a-l940a. Catalogue lichenum universalis, vol. 1-10. Leipzig. 1926b. Lichenes, p. 61-270. In Engler and Prantl, Die naturlichen Pflanzen-familien. ed. 2. Leipzig. 1930b. Symbolae Sinicae. III. Lichenes. Wien. 254 p. Zschacke, H. 1933-1934. Epigloeaceae, Verrucariaceae und Dermatocarpaceae, p. 44-656. I2_Rabenhorst's Kryptogamen-Flora von Deutschland, Osterreich und der Schweiz. ed. 2. 9(l,l). 440 Table 1. Degree of similarity of the lichen vegetation growing on . v various species of oak in the red oak forest. Coefficients of 2w a+b association were based on the formula C: x 100, where a = the number of lichen species on one tree, b = the number of lichen species on the compared tree, and w = the number of species found in common on both trees. A value of 100 indicates perfect association (i.e., identity, as far as lichen vegetation is concerned). A low value indicates relatively little similarity. Q. 1e_l_. g. cocc., Q. cocc. Q. alba Q. rubra Q. prinus x rubra Quercus velutina --- 71 71 68 6O 42 Q, coccinea --- --- 82 73 70 46 Q, coccinea ? x rubra --- --- --- 74 73 46 Q, alba --- --- --- --- 81 61 Q, rubra --- --- --- --- --- 66 441 'Table 2. Coefficients of association of lichen vegetation on different tree species at base and breast height quadrats. The coefficients were computed as in table 1. a. RED OAK FOREST Base Breast height Quercus rubra - Q, £123 70 67 Q, velutina - Q, rubra 81 78 Q, velutina - Q, alba 83 74 b. PINE - OAK FOREST Base mes. miss; 9. velutina - 9. 5Q; 83 57 («50)10 Since the asymptote of the species sample curve for the Quercus velutina - breast height data under "pine-oak forest" was not as sharp as was seen with the other curves, an extrapolation from 45 to ca. 90 trees was made, which adds approximately 2 species to the Q, velutina flora. It can be assumed that one of the two is shared with Q, 3123' in the Q, velutina - Q, gl§g_comparison, raising the coefficient from S7 to 60. (See full discussion on pages 59-61). gonnfl I!!! mean 5: cu .IIJINI ea m - -vousa moum - «cause ha I I I 53H“ :35. scene: 30‘— : __ I 2450““ ¥U=~£ I I I . I .II. I .1 II A . y .11.r - I All'lll'lyr .. i. J .Ill's'l'i .l I. ill:[ I] II" III A il.‘ . madam madam sunny ad .n UHmmN . as, a a wmmnnm .d . .o a «new ad .mmm «a uaaam .m mans» ad .0000 ad .ooou ad .q ..me .a 3:. .d 4.6mm .m M51949 Ind-mum ..fld .d .N V .0000 ad mnaaa mafia: .H UHWMZ some oomwusm oa=Ho> unmask hum "Eoum «use saws .umosoH one as: 5 human: can .muwueemu eunuuuoa uuonwwm use as: H monasz .mm «Heme Scum one some use .zufioeamo ousuewoa xumn uwunu mo nacho a“ voxauu edema .nm «Ha-H cache wage cu acmuawwuw> magma, m - scope» zoum u a u - n n u : umuomfiaan uneven Cu 30 W (1!! H E I £4335” 13543 x--- I 443 Table 4. Sand and soil pH. All samples were from uniform surface material, mixed into a slurry with distilled water, equilibrated for about 15 minutes, and measured using a glass electrode pH meter. Only one sample from each source was studied. General category Specific source locality, pH Exposed, eroding Yaphank 4.3 ground, supporting Baeomyces; central Commack 4.2 Long Island Riverhead 4.1 Beach sand from Fire Island (Bellport) 4.5 south shore Napeague Beach 4.6 Sand from central near Manorville 4.2 part of island Sand from north Shoreham, beach sand 6.2 shore beaches and behind very low dunes dunes Rocky Point, top of high 6.1 bluff, facing L. I. Sound Rocky Point, over crest of 5.1 bluff, protected from full on-shore winds Rocky Point, on beach 5.8 Transplanted From: Fire Island (south shore); 4.5 samples of to: Bellport on Great South beach sand Bay; for one year From: Fire Island (south shore); 5.8 to: Shoreham (north shore, on beach behind very low dunes); for one year 444 Table 5. Vertical distribution of some corticolous lichens in red oak and pine-oak forests. Not all species listed were treated in the pine- oak forest data because some were absent or too infrequent, and some, due to recognition problems with sterile material, were not included in early sampling. Red oak forest data were collected from localities 7 - 11 in the north shore transect. The pine-oak forest data are from continuum segments A and B, in Brodo (1961a). RED OAK FOREST PINE-OAK FOREST Total fre- % of total Total fre- Z of total quency (Z quadrat quency (% quadrat of 300 trees) occurrences. __ of 300 trees) ocgurrences. Base l.§_m Base 1.3 m Cladonia chlorophaea 9 100 0 42 96 4 Q, coniocraea 62 93 7 Graphis scripts 10 22 78 Hypogyggia physodes 3 37 63 Lecanora caesiorubella 6 6 94 L. chlarotera 5 25 75 11 33 67 Parmelia caperata 24 76 24 4 67 33 g, rudecta 24 69 31 3 86 14 g, saxatilis 14 57 43 11 60 40 g. sulcata 22 9 91 45 21 79 Eggtusaria xanthodes 4 38 62 Physeia millegrana 3 4O 6O 8 16 84 PE, orbicularis 3 100 0 5 71 29 o o o o o o o + o o e xumm ueouom o o o o o t. + o + o 2 5: 25m .232 To + + + + ++ ++ ++ + ++ a... .: mcauem Boo .1... ++ .I. - ++ ++ ++ +1 +1. ++. 3 uoxsauom .I+ .I+ .1. TI ii are .1... i. .1... .1... +1. mm 3:305 .~.< o o o o o + + ++ +_ + c swam ueouom o o o o o o + ++ + ++ 2 ..a 28m .33.. o o + o o ++ ++ .I. + ++ mm 3358.5 + o + + 0 +1. ++ +I. +++ ++ em xoaaaoo .1. .I. +.I. .TI. ii. .I. .I. .TI. ++ .TI. mm ozfruoouz m e m N a 1m m e m N if @388 5:33 .3. Beam one“: mouse mnH enemasz .oowam cw howsoH o: xmwv .u "mommquuouov mfiouoaeaoo .o mouoamaou -fiuouom .+ "commenceuouom mace .++ ”commenceuouom manquoouoc o: .+++ .uooesmuu madman sumo: AN “AnHooH .omoum cw vouuoaouv economuu vcmaeH woo; Hmuucoo afi .cowumucmamncmuu .eoumowfieou ou uooaau oo«ueuo Mao can .ouono momma use» «no mcouum>uomno .m .COwumucdenamuu momma Anonaooon nwnounu wonaoueomv unucoa know acouum>ummno .< .mumuommo wwfioaumm wage: uuooaowuemxo ucmHeaemuu somewfi msofiooauuoo unmauunmo ecu mo mufiemmm .w magma new ~9~un~ \ 446 Table 7. Lichen species diversity in various habitat types in ten localities closest to New York City. The numbers under the habitat headings indicate the number of species found at each locality. The distance from central Brooklyn (in miles) is indicated in parentheses next to each locality. Localities Saxi- Terri- Corti- Corti- Ligni- No. Name colous colous colous colous colous (base) (above base) 1. Prospect Pk. (2) l - - - - 2. Forest Pk. (9) 3. Alley Pd. Pk. (13) 2 4 l - - 4. Sands Pt. (19) - 2 4 - - 5. North Hills (15) - 5 - - - 6. Valley Stream (14) - 2 - - - 7. Rockville Center (17) 1 - - - - 8. E. Meadow (20) - 9 - - - 9. Brookville (24) - l l l 4 10. Glen Cove (23) 5 4 4 l 2 TOTALS 9 27 10 2 6 447 Table 8. Growth forms and city tolerance along a north shore, red oak forest transect. Numbers under the growth form headings refer to the number of species of each type found at each locality. Lp = leprose crust; Cr = crustose; Sq = squamose (including Cladonia); P I Parmeliopsistype; Pa . Parmeliatype (after Barkman, 1958). Distances from central Brooklyn (in miles) are indicated in parentheses next to each locality. Total no. of Lp Cr Sq P Pa corticolous Locality species Forest Park (9) O 0 0 0 O 0 Alley Pond Park (13) l O 0 l 0 0 Split Rock (16) 0 y 0 O 0 0 0 Brookville (24) 3 0 0 1 l 1 Laurel Hollow (28) 4 l 0 2 0 1 Cold Spring Harbor(30) 9 l 2 l l 4 Centerport (35) 7 l 0 3 0 3 Vernon Valley (37) 13 2 l 3 2 5 Sunken Meadow (39) ll 1 l 3 1 5 St. James (42) 21 l 6 3 3 8 448 Table 9. Atmospheric pollutants in New York City (Manhattan) and a suburb (Glen Cove, Long Island) (from U.S.P.H.S., 1962). The sulfate data are not directly comparable with 802 levels since a large proportion of the sulfate concentration consists of particulate sulfate compounds (Corn and DeMaio, 1964). All values are in ug/m3. New York City Glen Cove Pollutant Maximum Minimum Average Maximum Minimum. Average sulfate 53.1 6.0 22.8 - - - (no data available) nitrates 7.4 0.4 2.2 - - - organic compounds (soluable in benzene) winter 33.1 5.8 15.9 36.0 1.5 9.5 spring 35.8 4.8 13.1 23.0 2.3 7.7 summer 56.0 3.6 14.9 21.4 1.5 6.5 autumn 32.1 5.9 13.6 37.8 2.9 10.5 particulate matter winter 389 116 203 327 26 119 spring 365 90 176 247 31 108 summer 532 73 182 305 22 106 autumn 330 79 168 268 39 117 449 Table 10. Phytogeographic categories represented in the Long Island lichen flora. Aspects of each species' world-wide distribution are noted as follows: A found in Asia; E ' found in Europe; N I North American endemic; X = not endemic, but absent from Europe and Asia. Species with oceanic tendencies but which cannot be placed in the Oceanic subelement are indicated by asterisks (*). Details of the distribution of each species and/or references to published summaries or maps are presented in the annotated list. I. ARCTIC-BOREAL ELEMENT A. Arctic-alpige sgbelement: no representatives on Long Island B. Boreal-temperatg sub t. Caloplaca pyracea AB C. pyxidata AE Candelariella aurella AB C. rangiferina AB C. vitellina AB C. scabriuscula AE Cetraria islandica AB C. squamosa AE Cladonia alpestris AB C. uncialis AE C. arbuscula AB C. verticillata AB C. cariosa AE Dermatocarpon miniatum AB C. carneola AE Diploschistes scruposus AB C. chlorophaea AE Hypogymnia physodes AB C. deformis AE Lecanora cinerea AE C. fimbriata AE L. dispersa AE C. furcata AE L. rubina AB C. mitis AE L. symmicta AB C. pleurota AE Lecidea macrocarpa AE 450 (Arctic element, Boreal-temperate subelement continued) Lecidea vernalis AE Placynthium nigrum AE Parmelia saxatilis AE Rhizocarpon grande AE P. sulcata AE Rinodina oreina E Parmeliopsis ambigua AE Sarcogyne simplex AE Peltigera aphthosa AE Solorina saccata AE P. canina AE Verrucaria muralis AE P. polydactyla AE Xanthoria fallax E P. praetextata AE II. TEMPERATE ELEMENT A. North_Temperate subelement Acarospora fuscata AE Cladonia coniocraea AE Alectoria glabra N C. conista AE Bacidia umbrina E C. macilenta AE Buellia punctata AB C. multiformis x B. stillingiana N C. nemoxyna AE B. turgescens N Evernia mesomorpha AE Caloplaca aurantiaca AE Graphis scripta AE C. cerina AE Lecanora atra AB C. citrina AE L. hageni AB C. flavovirescens AE L. muralis AE Candelaria concolor AE L. varia* AE Catillaria glauconigrans N Lecidea aeruginosa AE Cetraria ciliaris AE L. albocaerulescens AE Chaenotheca phaeocephala E L. botryosa AE Cladonia bacillaris AE L. coarctata E 451 (Temperate element, North Temperate subelement continued) Lecidea nylanderi AE P. stenophylla AE L. quadricolor AE Pertusaria amara E L. scalaris AE Physcia adscendens AE L. uliginosa AE Ph. aipolia AE L. viridescens AE Ph. orbicularis AE Lobaria pulmonaria* AE Ph. stellaris AE Micarea prasina AE Rinodina confragosa AE Parmelia caperata AE R. milliaria N P. conspersa E Sarcogyne clavus E P. reticulata AE Stereocaulon saxatile E Usnea longissima AE Verrucaria nigrescens AE B. East Temperate subelement Anzia colpodes X Cladonia cristatella N Bacidia atrogrisea AB C. floerkeana AE B. inundata E C. parasitica AE B. schweinitzii N C. strepsilis AE Buellia curtisii N C. subcariosa AE B. polyspora X C. subtenuis X Cladonia apodocarpa N Dimerella diluta AB C. brevis E D. lutea AE C. caespiticia AE Lecidea anthracophila E C. capitata AE L. cyrtidia N C. caroliniana N L. erratica E C. clavulifera A ‘ Leptogium corticola E 452 (Temperate element, East Temperate subelement continued) Leptogium cyanescens AE Physcia subtilis N Leptorhaphis epidermidis E Ph. tribacoides E Micarea melaena AE -Po1ybleeeéepeée—éallen-——N- Parmelia aurulenta A Pycnothelia papillaria E P. galbina A Pyxine sorediata A P. livida N Ramalina fastigiate AE P. perforata E Trypethelium virens N P. rudecta A Usnea mutabilis N Parmeliopsis aleurites AE U. strigosa A Phaeographis dendritica AE Physcia millegrana N C. Appalachian subelement l. Appalachian unit: Buellia dialyta N Haematomma sp. N B. stigmaea N Parmelia appalachensis N Cladonia piedmontensis N 2- Amhahian- Mania; Caloplaca camptidia N Parmelia hypotropa N 3. Appalachian - Great Lakes unit: Alectoria nidulifera AE Collema subfurvum* AE Anaptychia palmulata A Haematomma ochrOphaeum A Bacidia chlorantha N Lobaria quercizans* N B. chlorococca E Parmelia olivetorum AE Baeomyces roseus AE P. subaurifera AE Cetraria tuckermanii N Umbilicaria mammulata N U. muhlenbergii AE 453 4. Appalachian - Great Lakes - Rocky Mountain unit: Cetraria fendleri N Parmeliopsis placorodia N Cladonia mateocyatha N Pseudevernia furfuracea E Parmelia borreri AE Umbilicaria papulosa X D. Coastal Plain subelement Bacidia chlorostica N Lecanora caesiorubella subsp. lathamii N Cetraria viridis N L. cupressi N Cladonia atlantica N Melanotheca cruenta N C. beaumontii N Parmelia mi'ch'auXiana: N C. boryi A Pertusaria propinqua N C. evansii N P. xanthodes N C. floridana N Porina cestrensis N C. incrassata AE Ramalina stenospora N C. santensis N R. willeyi N C. simulata N Usnea trichodea A C. submitis A E. Oceanic subelement Nephroma laevigatum AE Xanthoria parietina AE Pertusaria velata AE Xylographa opegraphella N F. Maritime subelement Verrucaria microspora E Verrucaria silicicola N III. TROPICAL ELEMENT A. Coastal Plain subelement Cladonia calycantha AE Cladonia vulcanica A C. didyma A Parmelia robusta E 454 (Tropical element Coastal Plain subelement continued) ’ Pertusaria tuberculifera X Ramalina complanata X Porina nucula E Teloschistes flavicans AE B. Appalachian - Temperate subelement Anaptychia obscurata AE Pannaria lurida X A. pseudospeciosa A Parmelia perlata AE Cladonia cylindrica A P. plittii X C. pityrea AE Teloschistes chrysophthalmus E~ C. Oceanic subelement Cladonia carassensis AE 455 Table 11. Phytogeographic affinities of Long Island lichens. All percentages are Z of total sample (209 species, or 81% of total lichen flora.) Total 1 of In Europe: In Asia,|. Europe N. Amer. # of flora not Asia not Europe & Asia Endemic species iiii #7.#7. I. ARCTIC 43 21 2 l 0 0 41 20 0 O l. Arctic-alpine 0 O 0 0 O 0 0 0 0 O 2. Boreal-temperate 43 21 2 l 0 O 41 20 0 0 II. TEMPERATE 149 71 19 9 ll 5 63 30 50 24 1. N. Temperate 52 25 7 3 0 0 39 19 5 2 2. E. Temperate 44 21 9 4 6 3 13 6 12 6 3. Appalachian 26 12 2 1 2 1 7 3 14 7 4. Coastal Plain 21 10 O 0 3 l l l 17 8 5. Oceanic 4 2 0 0 0 0 3 l 1 1 6. Maritime 2 l 1 l 0 O O 0 l 1 III. TROPICAL 17 8 3 l 4 2 6 3 0 O 1. Coastal Plain 8 4 2 1 2 1 2 1 0 0 2. Appa1.-temp. 8 4 l l 2 l 3 1 O 0 3. Oceanic l 1 O 0 0 0 l l O 0 TOTALS: 209 100 24 11 15 7 110 53 50 24 456 Table 12. European-American vicarious sub-generic taxa in the Long Island lichen flora. In the cases with asterisks the parent or daughter populations have apparently continued to diverge and speciate producing double-taxon vicariants. The problem, while slightly more complicated, is basically the same. Alternative no. 1 of Degelius (1940) refers to "true Vicariants" with one species found exclusively in America and the other, equally abundant, found only in Europe. Alternative no. 2 refers to "sub-Vicariants" with the European species represented in the American flora as a rare or very local plant in addition to the more abundant American species. America Europe Degelius (1940) Alternate Number 1. Cladonia subtenuis C. tenuis 2 2. Hypogymnia furfuracea H. furfuracea l (lecanoric acid strain) (olivetoric & physodic acid strains)* 3. Lobaria quercizans L. amplissima 2 4. Umbilicaria papulosa U. pustulata 2 .mm~ we HeuOu a no use «as» :ofiumuowu> ummu mo mouuwamooH no names: one moumuuvau ammo nodumuowo> noes mo saw: emu soHon mononucoume :« pupae: 0:9 A.oauou oeonu mo :oaeeaouwv ouoanaoo ouoa m how men owed oomv .uomhu coauMuowo> womuo cmnu momma: %Hu:mo«mucwuu on cu unmade moofim> omonu can: .uomhu couumuom0> 03» no oao has a« condensed can ooaomoua noon cw NON :mnu ouoa mo uoaam> no momma ea :Auqsovwm: .AmuaamooH use coaaoodm moo wowssemmv mowooam umnu mo mooawoonn mo Hanan: Hmuou one no uao omhu coauwuowo> umasoauumd mam aw nuaoodu umfiaoauumn a mo maeawoome mo ammuooouom emu moumofimcw sosHm> AMP: any .eowoomm umasouuuua e wow>mn onhu coaumuowo> umasowuume hem mo moeuwamuoH mo ammucoouod emu mouoowveu sooaonoum: .vaoHuH wcoq co moewu :oHumuowo> meowum> :« ononoua canoo 080m «0 newunawuuuwn .m~ canes ..__.L( c: m...— 3. do mmsw uN we w© canoe. upon vwowuu. datum HH Nm Ha No Ho mo wens mundane vwsocomx monomn we we ww o mN Nu mm mm Nu 0 Ho Ho be No uN momnwon onow our monomn we NH um NH um HH Nu No No we NH Nu No am and our monomn No Nm Nm mN 0 up o Hm am we uu no Nm Nm Ho bu arena ocean aspen N N Zappa mammm u m mxnno o o Hm venom. mean 3.95? meson memo monsoon mwuooomr monemn monn~an chow our monoun _+ woe our monoun arena ocean «swam . mHo ama .oucam ocwm amc a o ea em a o Ha wemaxsmnoas cm on «H mm on m an mvw>aa c on m we «a e~ ca «mangoes; o o be ma mm 0 ea «canes» o 0 Ha a e m 0H manoeucoo RN on nm em um 5N mm mumuommu mH 0 NH 5 o 0 OH «pounce «fiaoaumm ma mm qH mm mm on Hm ncmwum> o o m m an «H ma muonawfia: o o o mm on 0 ma nuumeon o 0 mm mm NN o om moaueuuo o mN o aN me 0 mm «Haneouuueuea o 0 mm o o 0 HH msooeoaauouUOAHm o mN m o «a as Na auoaamsuou museums AN mm be an me mm so euououmamo mm on me ooH em He mm mHHonsuoweomo muosmuuq Ame «use Ame AmMe Awmv llnllmmmq. ANNV maoasumaeou wuuxm mamsm QEmSo umouom umoHOM umouow assoc mo .0: made: umpoo xmo mom xmo xomfin menocwe .nawmfld Hmuoa some: uuoaumom moouumn meme scam .eocso and £60 —.‘_ am.o ca.a - m~.o Ase ma.o ANV om.o BE muououmfinu «monsoon om.o m~.o - oo.o Aoav am.o Acme em.o mHNoADHOfiaomo muoemoog o~.~ me.~ - e~.o A~e oo.~ Ame Ha.o EB mcmuwuaada wwomNnm no wcwmm >~meow>no mo eucuaousnmua oz mononueoumm :« «Henson any mosam> one MOw uqooxm ch.~ ao.~ - Ne.a a~e we.~ any o¢.~ ewawumudm mdauahmm Noe Nw.H he." . ee.o Ame o~.~ Ame me.a a! weakened eaaoauam aN.~ oc.n 4 e~.o Ange Ne.a Away m~e.o ll «unease cumulhem .movsaoou one: “gamma weuuoeaooov I .xmo so wowsouw nsozowa escaoofiuuoo mean we momma museum Hoodoo oweuo>< mue 0» women enemas: one .owouu mo .sa «mace 463 ~ Figure 1. Soil types (after Cline, 1957). a) excessively drained hilly soil (Plymouth-Haven AssociationL b) excessively drained sandy soil (Colton-Adams Association), c) Bridgehampton fine sandy loam, d) well drained, prairie-type soil (Hempstead- Bridgehampton Association). Figure 2. Original vegetation. a) red oak forest, b) pine-oak forest, and pine barrens c) scarlet-black oak forest, d) Hempstead Plains grassland, e) downs grassland and dune heath. Figure 3a. Mean precipitation for growing season, May 1 - Sept. 30. :euiveiy £11 essively '2!!! mate: 1! (5:15:21!- b) PM 31'“: Id 1nd Figure 3b. Figure 4a. Figure 4b. 464 Mean annual precipitation. Average January temperature (in degrees Fahrenheit). Average July temperature (in degrees Fahrenheit). will“ 465 Figure 5. Average annual number of days of dense fog. Figure 6a. Average July relative humidity. —.--- 8:00 a.m.; noon; ———— 8:00 p.m. Figure 6b. Average January relative humidity. —- —. — 8:00 a.m.; noon; —-—-— 8:00 p.m. ( (\ O 00: 8369 (33: O O O o 00 Q) 0 0083 ,» 20 ‘ 4O 0 O 2'” 30 :m .moauouoa usesum hummus one omoHu emu so noon» any .umouou xmo may one «o aouuuoe a moon on one unmade on» «o uwaase emu u< .Aeuouowm one no uwua emu cuv venom vouHuH moon wswxoofiuo>o umwaan ouomn zuuoz .ofi .«Nuon ad was nauaansu muaovuao hgswaa .omueoano mouoHou unwafi emu mam AmmOucoEOu uwsouvsmv pompous oeauu .Amuafimm«~w>own mHasmoEEa mOstmmuuouou< madame we uo>oo mcsouo .x:Mu:oz use: commomez um ecfimam use» was madam vcem .w .moa:m>thccoe some»: use .mm0ucoEou mHGOmmom .muwanwwaw>oun wagnmoea< mo masque mdwueaeaoo .cmooo one memos“ .oswoso use: moose «mono nuson no cowumuowo> scam mcmm .5 .modhu soaueuowo> .oHon newsman 00¢ ..K.,~ «.1. . tr... ,7 .....s». .... fife .muonmouuwa wwumoauuo> mo cowuooaaoo a mo unease: use was news: Axooz UCNHQH EN”; UfiNHnH HOUHOSW GO £0M0£ H0>QH$ Una“ 00.2%“ VOHOUHflfiu < ..VH .ouon venom one: moaucmfium wacovmao wo evsmue snag .mvwmwu madam hp movaaouuae .uuoeuwmm um ouonm muse» one use: won owe No>mum Hanan < .MH .mmm Homewoom> mo nusoumuomca so was mocuoooo ad vas mcwuaHo> uaouoad flame sues .vcmHnH wood asuucoo cw o-u>uonmz use: macho“ xoo xosHm .Na .mocwoooo ad was .mnam eaouond .mvflmwu macaw he causewaov .vamHmH mcoq Hmuucoo CH mucumuomug amcowuqz no>mexooum um ue0u0m secoocwm .aH .moehu sowumuow0> .qanaa newsman nod if .mofiuwamoofi COwuooHHoo mow .me assume 00 “(S filitu 5 Figure I5 4*,“ 469 Figure 16. Population changes in a corticolous lichen conmunity: non-directional shifts. a) Parmelia sulcata, b) Physcia millegrana. ”q 470 Figure 17. Population changes in a corticolous lichen community: directional shifts (succession) or a portion of a cyclic change. a) Parmelia sulcata, b) Physcia millegrana, c) Lecanora caesiorubella subsp. lathamii. .NH mam .oH .q .N .H mowufiamooH a“ a: um» um: unusauoaxo unmammcmuu upon» some: Home 059 .NH nwsoumu H euopaon one mafimsue economuu unassummo one new mom: mowufiamooH unseen xmo men .ouonm sumo: any .qH .mH .m .N .H enemas: one Anaomfiv ovoum cu moo: nuwuflamooH memfima Hmuucmu 0:9 .mcwumamooa unmaaacmnu mom uoomamuu umo3nuumm .mH unawam and £32.. «.0— —.Q ...—E o— n m_ 0.53”. 70 472 60 50 40 30 Figure 19. Distribution of the common trees along the north shore, red oak forest transect. A, Quercus velutina; B, Q, rubra; C, Q, alba; D, Q, coccinea; E, Acer rubrum; F, Fagus grandifolia. Frequencies were derived from 50 (A) A tree samples in each locality. ch pERCENr FREOUE a: ,4: ‘ I 1 PERCENT FREQUENCY 70 6O 50 40 3O 4O 30 IO 30 20 IO l 2 3 4 5 6789 I0 II I2(b) e /\ \ \‘D u! \ o \/ \ ’/o T I1 H 50 60(0) I 2 3 4 5 67 89 I0 II I2(b) —“. I I. E7 I l I I I IO 20 30 40 so 60(0) 12 3 4 56789|0|| I2(b) DISTANCE FROM CENTRAL BROOKLYN (miles) (a) and LOCALITY NUMBERS (b) FIgure I9 473 40 35 ER 25 Figure 20. Tree size along the north shore, red oak forest transect. Diameters at breast height (dbh) of Quercus rubra-and g. velutina are plotted against mileage from industrial Brooklyn. A gap appears between localities 4 and 6 because no black oaks were found in locality 5. 40 35 30 25 20 I0 DBH (centimeters) G l I l I l l I I I I I I T I I I I I I0 20 30 4o 50 60 (a) I 2 3 4 5 67 8 9 I0 II I2 (b) DISTANCE FROM CENTRAL BROOKLYN (miles) (a) and LOCALITY NUMBERS (b) Figure 20 '1‘]! .mumuummo aM .x N mmHHaumxmm .m .eI:II.IIe "muoomou «afloaumm .eII.I.ae “mumwuom menamuo a D¢I<¢-¢«D ”mumogam mzospmm .4I..I..I< mwumuooacco macommao . o o .muom w hugamuoH can smegma mocua :dmuw use .Avcsom vcmHmH wood ob muweexoue mum cu out hanmnouav muon conoqfi Hmuoeow noon kaoauuuxo em on: w zuwfieooa omawoem .humeUOA use moouu om mo magmas» aw unwwo: unseen no omen pomufio um ouceuuoooo unwound no women one mofionuocoum .uoomamuu umouom xmo mop .ouorm some: emu macaw neomUAN canoo oaom mo cowusmwuumav 05H .mHN ouowwm and 3 3.2.5 U_N oeammm t: 23232 5:80.. .23 23100: 35:23 20$ 3252a .‘OO ...... I ADNJOOSXJ iNlDlad ”4 475 Figure 21b. The distribution of some lichens along the north shore, red oak transect (drawn on a double logarithmic scale). Frequencies are based on percent occurrence at either base or breast height in samples of 50 trees per locality. A regression line was estimated and drawn by eye through the points from the lowest value to the peak value. Points beyond the peak (mainly in localities 10-12) were ignored since they do not enter into the slope of the initial rate of increasing lichen frequency with decreasing city effect. o-—————-o, Cladonia coniocraea; A-n-"A, Parmelia sulcata; fl, e—~-e, Z. rudecta; e—-——«—o, P_. saxatilis; x... ..... x, P. caperata ..v~'-,. PERCENT FREQUENCY 50 30 20 I l l 14] (a) (b) DISTANCE FROM CENTRAL BROOKLYN (miles) (a) and LOCALITY NUMBERS (b) Figure 2|b .ww 476 Figure 22. Bark-borer. a) Assembled, ready for use, b) with blade removed to show additional features of the steel holder. MAIL. i 22 477 Figure: 23. Ikansplanted bark disks bearing Parmelia caperata. ‘ Edustographs at left were taken in the laboratory the evening of disk; removal in August, 1961. Photographs at right were taken in early September, 1962 g; gig, where the disks were transplanted. Disks were removed from a black oak in Shoreham on the north shore, and transplanted to Shoreham (no. 201), Setauket (no. 202), Cold Spring Harbor (no. 203), Alley Pond Park (no. 204), and Forest Park (no. 205). Distance frmm central Brooklyn is indicated by the numbers to the right of the photographs. i ...... .. . n i a. .3 . . .v . :4: R3 ll {1 we inks are .. “ 2‘: ‘Ji 1. 478 Figure 24. Schematic representation of a possible mechanism for the city-effect. The city or town is represented by the heavy black line on the abscissa, with the zero point being at the town center. The drought effect and pollution effect curves are hypothetical, and are based mainly on the observations of the authors cited below. They depict the percent of the total effect acting at any given point. Lichen abundance (including both degree of cover and number of species) is represented by the width of the lichen abundance block. The degree of abundance is varied by the position of the lower edge of the block. The percent of the corticolous lichen flora showing a change in normal vertical distribution is represented by the position of the upper edge of the lichen abundance block. The point at which the upper and lower edges of the block meet, corticolous lichens are no longer present. A. Based on the data of Rydzak (1958) from Polish resort towns; B. based on the data of Jones (1952) from the midlands of England; C. based on Long Island data. a) drought effect, b) air pollution effect. OR eFFECT 96 cuu3u6”*7 POLLUTION OR 9a pnOUGHT ,— O O FFFECT SO 8 O .— I: 0 :3 O 8 C3 6‘3 ‘;:5115': '-: 'fé 9' .- g,’ 30' ~.‘ .. '4‘ I': "'M " e: O .n'.‘ 3.5...- .— I! o '3! Z" 0 s ' D O .' 8 ;‘.Z "’ a , o\° 5:229: " .I U. .: :2 F v .-'.‘.1é A A; .f ”I '5' u j :w ' 8 39v 0 .— I: 0 :3 O 8 . :5 O 5 " 39 .r’ 4‘ v' POLLUTION EFFECT POLLUTION EFFECT POLLUTION EFFECT 100 ......m. 1 o 0.... Q,. '~.. lICl-IEN ABUNDANCE so .‘ A . so 3 '2 o‘. 1 E E E o ‘ ‘ A A a I” I 2 a 4 s 7 DISTANCE (miles) « o LICHEN AIUNDANCE B . so 4 A I ‘00 no 24 2s DISTANCE (miles) lICHEN AIUNDANCE 100 “S'J'suv-‘z'I'“HI-on...” W 0 Iv.,..... Inna...“ ....'. 0.. Q so . . so o_ 100 DISTANCE (miles) % VERTICAL CHANGE % VEImCAL CHANGE CHANGE % VERTICAL 479 Figure 25. Arctic-Boreal element; Boreal-Temperate subelement. ‘\V - Cladonia alpestris (after Ahti, 1961). .°\\ \\=_ ,' .3 \\\\\- K I; 5,1233: Under 200 m :I I:] 200 to 500 - 500m 2000 ‘I X \ - Over 2000 I, I I I g/ / \I:I::i: ,: , \‘777 Figure 25 480 Figure 26. Temperate element; North Temperate subelement. Physcia stellaris (after Thomson, 1963). ‘I7L [:1 Under 200 m ‘ l I I:] 200 to 500 ‘ I I - 50010 2000 "‘27-. ,1, \7 7 I \I - 0m 2000 ‘ ‘ ‘ la , Z. i 7 / ‘77 I 7, ’ - 74‘, I Figure 26 ....~ 481 Figure 27. Temperate element; Appalachian subelement; R Appalachian unit. Parmelia appalachensis (after Culberson, } 1962). 4! A ".3 -..‘v'l-’- .v'". Figure 27 :I Under 200m 200 I0 500 500 I0 2000 Over 2000 482 Figure 28. Temperate element; Appalachian subelement; Appalachian-Ozark unit. Anzia colpodes (after Hale, 1955c). , .¢~ M3 o O O 2 m 0 O 5 0 0 O 2 I a. v 0 28 igure F .*A 483 Figure 29. Temperate element; Appalachian subelement; Appalachian-Great Lakes unit. Parmelia olivetorum (after Culberson, 1958b). [::I Under 200m 20010500 500:02000 Over 2000 " 'v'vo. WEE . , s',‘ 64 ;.‘. 4. _ In . ‘ ",Is - ‘A‘ 0., ‘— v V. Figure 29 ..I’L‘ 484 Figure 30. Temperate element; Appalachian subelement; Appalachian-Great Lakes - Rocky Mountain unit. Pseudevernia_furfuracea (after Hale, 1955c). \ ,I _: \.«\ / , y/ 7\ fl I: Under 200 m I I I [:I 200 Io 500 I I n 500 to 2000 Ix‘ I\ - 0m 2000 Figure 50 .w-fi 485 Figure 31. Temperate element; Coastal Plain subelement. Ramalina willeyi (after Howe, 1914). 200 to 500 I 500 to 2000 Over 2000 Figure 51 aw .‘ 486 Figure 32. Temperate element; East Temperate subelement. Parmelia aurulenta (after Hale, 1958). ‘~- IN“ I ‘3 7 ”(A 9’ ,( rsl (dfigm L ~L [:I Under 200 m (::] 200 I0 500 - 500 Io zooo \ Over 2000 . - \ I"! I _/,_ ::::I\m:i____ r, , , A *1 Figure 52 M- 487 Figure 32 A. Temperate element; Oceanic subelement. Nephroma laevigatum (after Wetmore, 1960). —.-._..._ SCI? , . / / a / / ,e- //’: Ag!".€u "3'“ I at 4’; ’If'”'~:17‘ ‘\ i__———-’j [ZZZ] 200.0500 I -5001o 2000 x \ - I ” ,I I““«"*‘”—‘*‘“" Figure 52A Ila 488 Figure 33. Phytogeographic affinities of the Long Island lichen flora. The three floristic elements are depicted on the abscissa. Percentages on the ordinate were derived from a sample of 209 species (81% of the total lichen flora). a) Species also found in EurOpe alone, b) species also found in Asia alone, c) species also found in both Europe and Asia, d) species endemic to North America. PERCENT OF SPECIES STUDIED 75 70 65 60 55 50 45 40 35 30 25 20 \\\\fl W su""~ 489 Figure 34. Historic relationships between floristic elements, subelements, and units in eastern America. Arrows indicate the general direction of the migration of species from one area (or category) to another. The categories have been placed in quasi-geographical positions relative to each other. Thickness of an arrow indicates the relative extent of the migration; a dotted line arrow refers to a slight connection. Trapical and Arctic-boreal elements indicate their worldwide affinities whereas the Temperate element is relatively isolated, except through its tropical or boreal connections. . . 1 ; Arctic-alpine ‘ K ARCTIC-BétkEAL Boreal -tempe rate A “J 2 w """ I 0‘ Great Lakes I I.“ a a. region 1 2 I if \l’ Rocky N. Atlantic Mountains Southern ........ Coastal S. Atlantic Appalachians : Plain & Ozarks Gulf A i F ..l < U I é. > Tropical (— O a: '— Figure 34 \V 490 Figures 35-100. In the distribution maps following, each apen circle represents a locality where a collection of a complete set of lichen species was made, but where the species in question was absent. With the exception of figure 36 (see below), a black dot indicates that a specimen of the species was collected in that locality. Specimens collected by Latham, Brainerd, Hulst, and others are mapped whether or not they were recollected by me in the same locality. In some cases (e.g., the pree~l900 New York City collections), these old records are of considerable historic interest. All Brooklyn dots represent pre~l900 collections. Figure 35. Localities of oceanic species. Included are the Long Island localities of Collema subfurvum, Leptogium cyanescens, Lobaria pulmonaria, E, guercizans, Nephroma laevigatum, and Pertusaria yelata. Xanthoria parietina was excluded since its distribution appears in figure 84. Figure 36. Bag and swamp localities. Figures 37-45. Lichens found mainly in bogs and swamps. 37. Cetraria ciliaris 38. g, viridis 39. Cladonia beaumontii 40. Q, didyma 41. g. santensis 42. Pertusaria amara 43. Usnea trichodea 491 44. Alectoria nidulifera (also in pine localities) 45. Parmelia hypotropa (also in maritime localities) Figures 46~S6. Lichens found mainly in pinevoak forests. (46wSl: with few or no localities east of Shinnecock; 52~56, with eastern extension; 46~492 pine specific; 50~512 oak specific.) 46. 47. 48. 49. SO. 51. 52. 53. Figures 57-67° lecidea anthrac0phila L. scalaris Ochrolechia parella Parmelicpsis placorodia Physcia aipolia Eh. stellaris Cladonia calycantha Q, floridana lecidea_varians Parmelia galbina g. perforata Lichens found mainly in morainal areas. (S7w60; terricolous; 6l~64z saxicolous; 65~67z corticolous) 57. 58. 59. 60. 61. 62. 63. Baeomyces roseus Cladonia squamosa Cladonia caespiticia 92222253 Acarospora fuscata lecanora cinerea Lecidea albocaerulescens 492 64. lecidea erratica 6S. Graphis scripta 66. Opegrapha cinerea 67. Parmelia borreri Figures 68~74. Lichens found mainly in the humid "fog belt" region. (73»74: fog belt species collected in New York City prior to 1870) 68. Anzia colpsdes 69. Buellia turgescens 70. Caloplaca camptidia 71. Arthonia siderea 72. Candelariella vitellina 73. Cladonia conista 74. Peltigera aphthosa Figure 75. An avoidance of the red oak forest. Physcia millegrana. Figures 76~77. The scattered distribution of two terrestrial lichens. 76. Cladonia strepsilis 77. Pycnotheli§_papillaria Figures 78~80. Lichens found mainly on sand dunes and sand plains. 78. Cladoni§_boryi 790 go Smeitis 80. Q, uncialis 493 Figures 8l~85. Lichens having a maritime distribution. (78w8l: aerohaline; 82: hydrohaline.) 81. Caloplaca citrina 82. Rinodina milliaria 83. 3. pachysperma 84. Xanthoria parietina 85. Verrucaria silicicola Figures 86 100. Presentmday distributions of some lichens under the influence of the city effect. The lichens are in order of decreasing tolerance. In these maps, no historic (i.e., pre~l901) western Long Island collections are indicated. 86. Cladonia coniocraea 87. Q. chlorophaea 88. g. cristatella 89. g. bacillaris 90. Bacidia chlorococca 91. Parmelia saxatilis 92. E. sulcata 93. Phaeographis dendritica 94. lecanora_caesiorubella subsp. lathamii 95. Parmelia caperata 96. E, rudecta 97. E, subaurifera 98. Pertusaria xanthodes 99. Lecanora chlarotera 100. Buellia polyspora 35 36 Bag and swamp localities 37 Cetraria ci liaris 0| “.2 ll“. 38 39 40 Cetraria viridis C ladonia beaumontii Cladonia didyma I Ill“. 44 46 Alectoria nidulifera Lecidea anthracophi la 00 2 It“. 47 Lecidea scalaris 48 Ochrolechia parella 49 Parmeliopsis placorodia k 00 u 0 O 4) O 00 O O O 0 m 0 o o 0 0 O . /a/ 5 IODLES OI IZKL“. 50 Physcia aipolia 5| Physcia stellaris 52 Cladonia calycantha 5 Wilt! II ‘ZKKW 53 54 55 Cladonia floridana Lecidea varians Parmelia galbina “.2 Kl“. u .n .1 . Ila-11!]. ulilulinltllll'ltl . 'Illr I‘IlIP, 59 Cladonia caespiticia 60 Cladonia pleurota 6l Acarospora fuscata W .0 ‘Z‘lw. 62 63 64 Lecanora ci nerea Lecidea albocaerulescens Lecidea erratica l0 ILES w—a “IKE“, 65 Graphis scripta 66 Opeg rapha cinerea 67 Parmelia borreri /’ 4 ' s IOMDLES I I I! "L“. 68 Anzia colpodes 69 Buellia turgescens 7| 72 73 Arthonia siderea Candelariella vitellina Cladonia conista IOZIILON 74 Peltigera aphthosa 75 Physcia millegrana . J‘s: .Q y 76 Cladonia strepsilis OI uzmou 77 Pycnothelia papillaria 78 Cladonia boryi 79 Cladonia submitis 0 ° '_ § .-‘2 .‘ K.‘() ,1) r) , . a L O o 0.0 DEE-3‘, “'0 00'- 0 ° C 0 ‘5’ o 80 0 oo o/, O 0 Q) 0 0 V»; N ,r ' [ 00 F i 5 l0 HOLES OJ I. Z K101 80 Cladonia uncialis 8| Caloplaca citrina v- k , ) o o o ”(O O O OO O o O O O O 0 O O O . O C O o o / 82 Rinodina milliaria B 2 It“. 86 Cladonia coniocraea 87 Cladonia chlorophaea o r )h 00 ( . , . ... 0° N .0 r ' 0.0 a ) . . ° 9. ° 0 o. 88 Cladonia cristatella . /‘——'—‘ 5 no nuts 8' IIZItW. 89 Cladonia bacillaris k % 7 90 Bacidia chlorococca . cg . W ...-x: , .b .09. M 0' 9| Parmelia saxatilis 5 l0 MILES Bl I62 IILOI 92 Parmel ia sul cata / 0' 93 Phaeographis dendritica . W“; C .1 J k {0 a 0’. , / (so 000 0 06;: (8‘70 0. - O ‘4 OO . 8p x.) : 0%. O . Cb O O y/ o 0 V’" 94 Lecanora caesiorubel la 5 I0 IKES II t! ZKILOI 7 95 Parmelia caperata *‘ fi 0 I 0 a o s ' o . G ’° . O . 0") v..‘ ( ‘ 0 ..O .. O O .. .0 .o .633 o: 0’ ... O. ‘/./ O 0 Q3 0 O W / . _’ o a. 0 ,./’ O / / ‘4/0/7 <7 96 Parmelia rudecta . 9 % y 97 Parmelia subaurifera 5 l0 MILES / n—o.--c-¢——-—4 0| ‘6 Z KILOM 98 99 IOO Pe rtusa ria xanthodes Lecanora chlarotera pt 0 1) OO 7'0 Buellia polyspora l0 IKES I62 KILOI .mUmaoN swamped; .mofi mfioowoucaa mammowummflnmflom .NOH mmamuomnam mwummauuam .HQH .pcuoewafima oao ma aHaum anu no scamwpda gamEe Loam .mmaoamm 30c manna mo mcafiauaaw emmH .mQHeHoH mausmfim .Auouma a“ vauasoev sauna Au .Amom ca wouaaoav snowmoumm An .nvaounu c«o»%£amuaa Am .AaamaoHonv maoofiouoaa ofimQOMuanoNHom .qu auawwm was ¢ 0 F .anuun .uum manque .fium “accounu pacemnawuma .uwm mafiaavoa_.vaa .Eaaauuaaao>aw .>afi "Eafiuosuoazn .mwn “anaashs .mN: moamwuxo .oxo "Esauazuwau .Hmo “xauuoo .poo maawuunuqsafim .smEm “nomad Human .mHa .mfioofiouoau mammoaummfimwaam a“ ma .Eafiuonuocaomm .o “mafiauonwn weapon a“ no .anoanufiuom .o m.aam sapwood nu ma .aawuoSuoau oaaokuoa .m m.dao muoamooq a« as .Eaaoucuoaa oawuoawooa .< .mapauoun< .moH unawwm ome 3— .....— h.‘ ‘ 7" “'\ --- ." ‘\)\.\ . fl . $295 282 o :2 85%,.“ 497 Figure 1.06. Thallus types. A. Hypophloedal; B. epiphloedal. bk, bark, Egg, necrotic layer of thallus; thal, living thallus tissue. Figure 10.7. Lichen phycobionts (camera lucida drawings). A. Trentepohlia (from Graphis scripts); B. Trebouxia fl (from Cladonia sp.); C. Nostoc, (from Leptogium cm). Figure 108. Some ascospore types. A. Polarilocular; B. mischoblastiomorphic; C. pachySPOI'OUS; D. murifom,